Toward ecologically scaled landscape indices

Nature conservation is increasingly based on a landscape approach rather than a species approach. Landscape planning that includes nature conservation goals requires integrated ecological tools. However, species differ widely in their response to landscape change. We propose a framework of ecologically scaled landscape indices that takes into account this variation. Our approach is based on a combination of field studies of spatially structured populations (metapopulations) and model simulations in artificial landscapes. From these, we seek generalities in the relationship among species features, landscape indices, and metapopulation viability. The concept of ecological species profiles is used to group species according to characteristics that are important in metapopulations' response to landscape change: individual area requirements as the dominant characteristic of extinction risk in landscape patches and dispersal distance as the main determinant of the ability to colonize patches. The ecological profiles and landscape indices are then integrated into two ecologically scaled landscape indices (ESLI): average patch carrying capacity and average patch connectivity. The field data show that the fraction of occupied habitat patches is correlated with the two ESLI. To put the ESLI into a perspective of metapopulation persistence, we determine the viability for six ecological profiles at different degrees of habitat fragmentation using a metapopulation model and computer-generated landscapes. The model results show that the fraction of occupied patches is a good indicator for metapopulation viability. We discuss how ecological profiles, ESLI, and the viability threshold can be applied for landscape planning and design in nature conservation.     

Metapopulation models for extinction threshold in spatially correlated landscapes

Simple analytical models assuming homogeneous space have been used to examine the effects of habitat loss and fragmentation on metapopulation size. The models predict an extinction threshold, a critical amount of suitable habitat below which the metapopulation goes deterministically extinct. The consequences of non-random loss of habitat for species with localized dispersal have been studied mainly numerically. In this paper, we present two analytical approaches to the study of habitat loss and its metapopulation dynamic consequences incorporating spatial correlation in both metapopulation dynamics as well as in the pattern of habitat destruction. One approach is based on a measure called metapopulation capacity, given by the dominant eigenvalue of a "landscape" matrix, which encapsulates the effects of landscape structure on population extinctions and colonizations. The other approach is based on pair approximation. These models allow us to examine analytically the effects of spatial structure in habitat loss on the equilibrium metapopulation size and the threshold condition for persistence. In contrast to the pair approximation based approaches, the metapopulation capacity based approach allows us to consider species with long as well as short dispersal range and landscapes with spatial correlation at different scales. The two methods make dissimilar assumptions, but the broad conclusions concerning the consequences of spatial correlation in the landscape structure are the same. Our results show that increasing correlation in the spatial arrangement of the remaining habitat increases patch occupancy, that this increase is more evident for species with short-range than long-range dispersal, and that to be most beneficial for metapopulation size, the range of spatial correlation in landscape structure should be at least a few times greater than the dispersal range of the species.     

生境破碎化对动物种群存活的影响

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

The use of spatial concepts as a basis for designing a viable-habitat network: Conserving redstart (Phoenicurus phoenicurus) populations in Sherwood Forest, England

This paper presents a method for the design of a habitat network in a fragmented landscape. It employs metapopulation dynamics and connectivity to develop spatial rules which, together with the requirements of a target species, can be used to guide the creation of a network to improve population persistence. Using these guidelines, two conservation strategies of patch enlargement and corridor creation are developed. The spatial models used to produce the habitat network are described. The resulting maps show the network consisting of the source patches that could be occupied by stable populations and a number of patches, which could support viable local populations, situated in the proximity of the sources or linked to them. The approach could be used to inform decision-making in nature conservation and landscape planning.     

Spatial modelling of the barn owl Tyto alba habitat using landscape characteristics derived from SPOT data

Landscape parameters extracted from classified SPOT satellite imagery are used as independent variables for predicting potential habitat areas of the barn owl Tyto alba in a landscape north-east of Brussels, Belgium Field data on the nest sites, recorded during 9 yr, are used as the dependent variable A canonical correlation analysis of the landscape characteristics of 'successful breeding and non-breeding sites' selects a set of most significant parameters contributing to the distinction between suitable and unsuitable breeding habitat parameters measuring the spatial configuration and fragmentation of landscape elements, such as deciduous woods and grasslands, in combination with some visual characteristics of the open spaces in the landscape The selected set of parameters formed the basis for the calculation of a habitat model, whereby potential breeding sites could be located throughout the complete study area, at specific levels of confidence The results showed the relevance of this method for landscape ecological research and nature conservation planning     

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.     

多尺度结合的西安市浐灞河湿地水鸟生境保护规划

城市湿地及其物种多样性的保护是城市生态建设的重要内容.结合西安市浐灞河湿地,对城市湿地水鸟生境的保护规划问题进行了探讨.经调查,西安市浐灞河段共发现水鸟20种,分属8目12科,孕育了丰富的水鸟多样性资源.同时保留有本地湿地植物种类,生境类型多样,具有很高的保护价值.浐灞河湿地水鸟生境的保护规划应以景观生态学、保护生物学、复合种群等相关理论为指导,区域/景观-局地-群落多尺度结合,保护与开发结合进行规划,具体从以下方面着手:生境网络的构建;小生境的设计,滨水地带土地利用格局优化及生境水深的控制;生境植物群落的构建.     

Viability analyses with habitat-based metapopulation models

Population viability analysis (PVA) models incorporate spatial dynamics in different ways. At one extreme are the occupancy models that are based on the number of occupied populations. The simplest occupancy models ignore the location of populations. At the other extreme are individual-based models, which describe the spatial structure with the location of each individual in the population, or the location of territories or home ranges. In between these are spatially structured metapopulation models that describe the dynamics of each population with structured demographic models and incorporate spatial dynamics by modeling dispersal and temporal correlation among populations. Both dispersal and correlation between each pair of populations depend on the location of the populations, making these models spatially structured. In this article, I describe a method that expands spatially structured metapopulation models by incorporating information about habitat relationships of the species and the characteristics of the landscape in which the metapopulation exists. This method uses a habitat suitability map to determine the spatial structure of the metapopulation, including the number, size, and location of habitat patches in which subpopulations of the metapopulation live. The habitat suitability map can be calculated in a number of different ways, including statistical analyses (such as logistic regression) that find the relationship between the occurrence (or, density) of the species and independent variables which describe its habitat requirements. The habitat suitability map is then used to calculate the spatial structure of the metapopulation, based on species-specific characteristics such as the home range size, dispersal distance, and minimum habitat suitability for reproduction. Received: April 1, 1999 / Accepted: October 29, 1999     

Landscape characteristics, spatial extent, and breeding bird diversity in Ohio, USA

Abstract. Avian communities are often used by ecologists as indicators of environmental decline over large spatial areas, because of the ease with which birds can be monitored by nonprofessionals and the availability of continent‐wide breeding bird data. The influence of scale on the relationship between bird diversity and the characteristics of the landscape, which can serve as proxies for decline, is receiving greater attention but is still not well understood. We combined data from the Breeding Bird Survey with landscape characteristics derived from the National Land Classification Data for Ohio, USA, to determine the effects of landscape extent on relationships between birds and landscape characteristics. These relationships were determined through previous work to be correlated with avian richness and diversity. We created areas of varying sizes using buffers around each of 58 routes, and calculated diversity for several groups of birds: all birds, five habitat guilds, and three migration guilds. The landscape extent over which landscape characteristics were considered affected the relationship between these characteristics and bird richness and diversity overall, as well as richness and diversity for several of the habitat and migratory guilds. Diversity of woodland birds, Neotropical migrants, and richness of short‐distance migrants were best explained by the landscape characteristics examined here, possibly due to a less homogeneous collection of species in the other guild groups. These results suggest that more attention is required in selecting the appropriate scale when using landscape characteristics to predict or manage avian communities, as some characteristics may be more useful for management activities over small areas versus efforts over larger areas.     

A formula for the mean lifetime of metapopulations in heterogeneous landscapes

We present a formula for the mean lifetime of metapopulations in heterogeneous landscapes. This formula provides new insights into the effect of the spatial structure of habitat networks on metapopulation survival, with consequences for modeling, landscape evaluation, and metapopulation management. In the whole study, the spatially realistic metapopulation model of Frank and Wissel is taken as a basis. First, as a key result on the way toward the desired formula, it is shown that a simple nonspatial (Levins-type) model is able to reproduce the behavior of the complex spatial model considered regarding the mean lifetime, provided its parameters appropriately summarize all the relevant details of spatial heterogeneity. Second, the formula presented reveals how data from species and landscape have to be combined to estimate the survival chance of a metapopulation without having to run any simulation or to solve numerically any model equation. Third, by taking the formula as a basis, landscape measures are derived that allow dissimilar habitat networks to be evaluated, compared, and ranked in terms of their effect on metapopulation survival. Fourth, a combination of analytical, nonlinear regression as well as aggregation techniques was used to deduce the formula presented. The potential of these techniques for simplifying (meta)population models that are complex due to spatial heterogeneity is discussed.     

Spatial genetic structure in a metapopulation of the land snail Cepaea nemoralis (Gastropoda: Helicidae)

Habitat fragmentation is a major force affecting demography and genetic structure of wild populations, especially in agricultural landscapes. The land snail Cepaea nemoralis (L.) was selected to investigate the impact of habitat fragmentation on the spatial genetic structure of an organism with limited dispersal ability. Genetic and morphological patterns were investigated at a local scale of a 500 m transect and a mesoscale of 4 x 4 km in a fragmented agricultural landscape while accounting for variation in the landscape using least-cost models. Analysis of microsatellite loci using expected heterozygosity (HE), pairwise genetic distance (FST/1-FST) and spatial autocorrelograms (Moran's I) as well as shell characteristics revealed spatial structuring at both scales and provided evidence for a metapopulation structure. Genetic diversity was related to morphological diversity regardless of landscape properties. This pointed to bottlenecks caused by founder effects after (re)colonization. Our study suggests that metapopulation structure depended on both landscape features and the shape of the dispersal function. A range of genetic spatial autocorrelation up to 80 m at the local scale and up to 800 m at the mesoscale indicated leptokurtic dispersal patterns. The metapopulation dynamics of C. nemoralis resulted in a patchwork of interconnected, spatially structured subpopulations. They were shaped by gene flow which was affected by landscape features, the dispersal function and an increasing role of genetic drift with distance.     

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.     

生境破坏的模式对集合种群动态和续存的影响

构建了空间关联的集合种群模型,该模型不但包含了种群的空间结构信息,而且引入了破坏生境的全局密度和局部密度两个指标,它们描述了破坏生境的模式.模型揭示了破坏生境的空间分布格局复杂地影响了集合种群的动态和续存,破坏和未破坏生境斑块的均匀混合不利于集合种群的增长和续存,而生境类型聚集分布可以促进集合种群的快速增长和长期续存;对于两种斑块类型相对均匀混合的生境来说,均匀场假设可能会高估集合种群的续存,对于相对斑块类型高度聚集的生境,均匀场假设可能会低估集合种群的续存;物种的迁移范围也会影响集合种群的续存,迁移范围越大的物种越容易抵御生境的破坏而免遭灭绝.这意味着在生物保护中不能仅仅考虑生境的恢复和斑块质量的改善,生境结构的构建也是很重要的,加强生境斑块之间的连通性也有利于物种的长期续存.     

基于鸟类视角的城市生态廊道构建方法研究综述

基于鸟类移动特征与栖息地需求的生态廊道不仅有利于减少由生境丧失与生境破碎化导致的物种灭绝,进而促进物种的基因扩散与交流,而且可以改善城市居民的身心健康,推动城市不动产增值。然而,迄今为止,由国外学者研发的主要生态廊道构建方法没有内在地考虑鸟类的移动与生境特征,国内学者的鸟类生态廊道研究则侧重于对技术方法的探索而缺乏对研究鸟种特征的认知。因此,基于鸟类视角的城市生态廊道构建研究尚有较大的改进空间。本文通过对1975—2020年的国内外相关研究进行分析,识别出了可结合鸟类生态的7种廊道构建方法及3种廊道优化方法,并总结了相关方法的优缺点及其适用情况。最后,本文认为将典型鸟类物种的观测数据、真实运动能力与景观结构相融合,研发出计算高效便捷而且能够模拟廊道生态效益的计算机模型,将会是鸟类生态廊道构建方法的发展趋势。     

Merging spatial and temporal structure within a metapopulation model

Current research recognizes that both the spatial and temporal structure of the landscape influence species persistence. Patch models that incorporate the spatial structure of the landscape have been used to investigate static habitat destruction by comparing persistence results within nested landscapes. Other researchers have incorporated temporal structure into their models by making habitat suitability a dynamic feature of the landscape. In this article, we present a spatially realistic patch model that allows patches to be in one of three states: uninhabitable, habitable, or occupied. The model is analytically tractable and allows us to explore the interactions between the spatial and temporal structure of the landscape as perceived by the target species. Extinction thresholds are derived that depend on habitat suitability, mean lifetime of a patch, and metapopulation capacity. We find that a species is able to tolerate more ephemeral destruction, provided that the rate of the destruction does not exceed the scale of its own metapopulation dynamics, which is dictated by natural history characteristics and the spatial structure of the landscape. This model allows for an expansion of the classic definition of a patch and should prove useful when considering species inhabiting complex dynamic landscapes, for example, agricultural landscapes.     

Habitat fragmentation: island v landscape perspectives on bird conservation

Fragments of habitat are often viewed as islands and are managed as such; however, habitat fragmentation includes a wide range of spatial patterns of environments that may occur on many spatial scales. Fragments exist in a complex landscape mosaic, and dynamics within a fragment are affected by external factors that vary as the mosaic structure changes. The simple analogy of fragments to islands, therefore, is unsatisfactory. Understanding how birds respond to these complexities of fragmentation requires mechanistic studies focused on habitat selection and movement behaviour. Conservation efforts must be based on viewing fragmentation as a range of conditions that occurs in a landscape mosaic, and management should be directed toward the mosaics rather than focusing solely on reserves.     

Breeding bird species diversity across gradients of land use from forest to agriculture in Europe

Loss, fragmentation and decreasing quality of habitats have been proposed as major threats to biodiversity world‐wide, but relatively little is known about biodiversity responses to multiple pressures, particularly at very large spatial scales. We evaluated the relative contributions of four landscape variables (habitat cover, diversity, fragmentation and productivity) in determining different components of avian diversity across Europe. We sampled breeding birds in multiple 1‐km² landscapes, from high forest cover to intensive agricultural land, in eight countries during 2001?2002. We predicted that the total diversity would peak at intermediate levels of forest cover and fragmentation, and respond positively to increasing habitat diversity and productivity; forest and open‐habitat specialists would show threshold conditions along gradients of forest cover and fragmentation, and respond positively to increasing habitat diversity and productivity; resident species would be more strongly impacted by forest cover and fragmentation than migratory species; and generalists and urban species would show weak responses. Measures of total diversity did not peak at intermediate levels of forest cover or fragmentation. Rarefaction‐standardized species richness decreased marginally and linearly with increasing forest cover and increased non‐linearly with productivity, whereas all measures increased linearly with increasing fragmentation and landscape diversity. Forest and open‐habitat specialists responded approximately linearly to forest cover and also weakly to habitat diversity, fragmentation and productivity. Generalists and urban species responded weakly to the landscape variables, but some groups responded non‐linearly to productivity and marginally to habitat diversity. Resident species were not consistently more sensitive than migratory species to any of the landscape variables. These findings are relevant to landscapes with relatively long histories of human land‐use, and they highlight that habitat loss, fragmentation and habitat‐type diversity must all be considered in land‐use planning and landscape modeling of avian communities.     

Using connectivity metrics in conservation planning – when does habitat quality matter?

Aim The objective of conservation planning is often to prioritize patches based on their estimated contribution to metapopulation or metacommunity viability. The contribution that an individual patch makes will depend on its intrinsic characteristics, such as habitat quality, as well as its location relative to other patches, its connectivity. Here we systematically evaluate five patch value metrics to determine the importance of including an estimate of habitat quality into the metrics. Location We tested the metrics in landscapes designed to represent different degrees of variability in patch quality and different levels of patch aggregation. Methods In each landscape, we simulated population dynamics using a spatially explicit, continuous time metapopulation model linked to within patch logistic growth models. We tested five metrics that are used to estimate the contribution that a patch makes to metapopulation viability: two versions of the probability of connectivity index, two versions of patch centrality (a graph theory metric) and the metapopulation capacity metric. Results All metrics performed best in environments where patch quality was very variable and high quality patches were aggregated. Metrics that incorporated some measure of patch quality did better in all environments, but did particularly well in environments with high variance of patch quality and spatial aggregation of good quality patches. Main conclusions Including an estimate of patch quality significantly increased the ability of a given connectivity metric to rank correctly habitat patches according to their contribution to metapopulation viability. Incorporating patch quality is particularly important in landscapes where habitat quality is highly variable and good quality patches are spatially aggregated. However, caution should be used when applying patch metrics to homogeneous landscapes, even if good estimates of patch quality are available. Our results demonstrate that landscape structure and the degree of variability in patch quality need to be assessed prior to selecting a suitable method for estimating patch value.     

Under‐representation of avian studies in landscape genetics

Landscape genetics is a rapidly growing discipline that examines how heterogeneous landscapes and other environmental factors influence population genetic variation. We conducted a systematic review of the landscape genetic literature which demonstrates that birds are severely under‐represented relative to their species diversity and general publication prevalence. Most avian studies were on species that have relatively low dispersal ability, and we suggest that this reflects an assumed high vagility of birds that precludes spatial genetic variation relatable to landscape heterogeneity. However, spatial genetic variation exists in several bird species with very high dispersal ability, but this has not been considered in the context of landscape features. Genetic patterns may also relate to landscape due to breeding habitat selection and territorial behaviour, despite the fact that species may be able to move throughout different landscape elements with minimal movement costs. Habitat loss and fragmentation are continuing globally and are strongly related to declines in bird populations. Landscape genetic studies provide a means to understand, predict and mitigate the effects of anthropogenic landscape change on birds. This review promotes the need for landscape genetic studies of birds, such that a greater understanding of the drivers of their genetic structuring can be developed and generalizations can be made from landscape genetic studies that apply more broadly across taxa.     

Consequences of forest fragmentation for the dynamics of bird populations: conceptual issues and the evidence

This paper reviews the consequences of forest fragmentation for the dynamics of bird populations. Owing to high mobility and large home ranges, birds usually perceive fragmented forests in a finegrained manner, i.e. embrace several forest fragments in functional home ranges. On a regional scale, however, coarse-grained clusters of fine-grained fragments (hierarchical fragmentation may sub-divide bird populations into isolated demes, which enter a domain of metapopulation dynamics. Distinctions are made between pure distance-area or population-level effects and more indirect community-level effects due to changes in landscape composition. Distance-area effects, such as insularization and decreasing fragment size, directly prevent dispersal and reduce population size. Landscape effects, such as reduced fragment-matrix and interior-edge ratios, increase the pressure from surrounding predators, competitors, parasites and disease. In short, forest fragmentation can be viewed as a two-step process. Initially, fine-grained fragmentation triggers distance-area and landscape effects on a local scale, which in turn, results in a range retraction of a population to non-fragmented or less fragmented parts of a region. At a certain point, non-fragmented areas become so widely spaced out that regional distance-area effects come into operation, giving rise to metapopulation dynamics. Although few bird metapopulations have yet been documented, metapopulation dynamics probably is a common characteristic of bird populations confined to 'hierarchical' fragmented forests.