Proximity of restored hedgerows interacts with local floral diversity and species' traits to shape long‐term pollinator metacommunity dynamics

Disconnected habitat fragments are poor at supporting population and community persistence; restoration ecologists, therefore, advocate for the establishment of habitat networks across landscapes. Few empirical studies, however, have considered how networks of restored habitat patches affect metacommunity dynamics. Here, using a 10‐year study on restored hedgerows and unrestored field margins within an intensive agricultural landscape, we integrate occupancy modelling with network theory to examine the interaction between local and landscape characteristics, habitat selection and dispersal in shaping pollinator metacommunity dynamics. We show that surrounding hedgerows and remnant habitat patches interact with the local floral diversity, bee diet breadth and bee body size to influence site occupancy, via colonisation and persistence dynamics. Florally diverse sites and generalist, small‐bodied species are most important for maintaining metacommunity connectivity. By providing the first in‐depth assessment of how a network of restored habitat influences long‐term population dynamics, we confirm the conservation benefit of hedgerows for pollinator populations and demonstrate the importance of restoring and maintaining habitat networks within an inhospitable matrix.     

Matrix context and patch quality jointly determine diversity in a landscape‐scale experiment

The biodiversity of a habitat patch is predicted to be driven in part by interactions between patch quality and landscape context (i.e. type of regional matrix), but these interactions are rarely explored experimentally. Understanding the interaction between patch quality and matrix context can provide insight into the kind of dynamics that best describe a metacommunity and help predict how the diversity of a patch will respond to environmental change at different scales. We conducted a landscape‐scale experiment to examine how regional and local aspects of the terrestrial matrix interact to affect biodiversity within artificial ponds designed to mimic generic features of freshwater ephemeral ponds. We manipulated both the kind of matrix surrounding ponds (open canopy grassland, pine forest, and hardwood forest) and pond quality (three different types of leaf litter substrate). Ponds were left open to natural colonization for three months by aquatic insects and amphibians. The terrestrial matrix had consistent and strong effects on biodiversity throughout the experiment: ponds in open canopy areas had more animal morphotypes than ponds in pine or hardwood forests. Leaf litter type affected biodiversity during the experiment, with more animal morphotypes in ponds with higher quality litter than ponds with lower quality litter, and this effect was stronger in open canopy areas. The effect of leaf litter, however, disappeared by the end of the experiment. Our results suggest that the matrix surrounding patches has strong effects on community dynamics and biodiversity within patches, and conservation efforts aimed at maintaining biodiversity requires simultaneous consideration of both matrix habitats and habitat patches.     

生态保护区外斑块对景观连接度的影响评价

快速的城市化过程带来的生境斑块破碎化及损失会影响物种迁移、捕食等生态活动,对生物多样性构成威胁。然而,现有生态保护区可能无法覆盖其内生物的必要活动范围。生态保护区外的生境斑块对于维持生态过程也具有重要作用,因此识别生态保护区外的关键斑块并加以保护非常重要。以北京市延庆区为研究区,划分两种生境斑块,即核心生境斑块和潜在生境斑块,并基于图论构建生境网络。考虑地表覆盖类型、坡度、人类活动等因素构建生境阻力面。结合未来土地利用类型变化的模拟,研究城市化过程对区域生境网络和景观连接度的影响,选用CLUE-S模型模拟土地利用类型变化的格局。结合生境斑块特征和未来城市土地利用变化情况设计了3种未来生境变化情景。利用连接概率指数(PC)和网络连接度变化率(dI)评价不同生境变化情景下生态保护区外潜在生境斑块的景观连接度重要性,判断保护优先顺序,并分析景观格局变化对不同迁移能力物种的影响。结果表明：生态保护区外的全部潜在生境斑块对维持生境整体景观连接度有最大2.15%的影响,单个潜在生境斑块对维持景观连接度有最大0.28%的影响。此外,景观格局及其变化对不同迁移能力物种的影响差异显著,因此需针对保护物种和城市生境特征设计保护方案,研究区需要优先保护大中型斑块和位于关键位置的小型斑块。为了满足对生物多样性保护的需求,建议在区分生境斑块保护优先顺序时考虑生境斑块对景观连接度的贡献和城市化扩展过程的压力。研究为城市生物多样性保护和生境管理提供了方法参考。     

Landscape-scale habitat fragmentation is positively related to biodiversity,despite patch-scale ecosystem decay

Positive effects of habitat patch size on biodiversity are often extrapolated to infer negative effects of habitat fragmentation on biodiversity at landscape scales. However, such cross-scale extrapolations typically fail. A recent, landmark, patch-scale analysis (Chase et al., 2020, Nature 584, 238–243) demonstrates positive patch size effects on biodiversity, that is, ‘ecosystem decay’ in small patches. Other authors have already extrapolated this result to infer negative fragmentation effects, that is, higher biodiversity in a few large than many small patches of the same cumulative habitat area. We test whether this extrapolation is valid. We find that landscape-scale patterns are opposite to their analogous patch-scale patterns: for sets of patches with equal total habitat area, species richness and evenness decrease with increasing mean size of the patches comprising that area, even when considering only species of conservation concern. Preserving small habitat patches will, therefore, be key to sustain biodiversity amidst ongoing environmental crises.     

Effects of landscape and patch-level attributes on regional population persistence

Species responses are influenced by processes operating at multiple scales, yet many conservation studies and management actions are focused on a single scale. Although landscape-level habitat conditions (i.e., habitat amount, fragmentation and landscape quality) are likely to drive the regional persistence of spatially structured populations, patch-level factors (i.e., patch size, isolation, and quality) may also be important. To determine the spatial scales at which habitat factors influence the regional persistence of endangered Ord's kangaroo rats (Dipodomys ordii) in Alberta, Canada, we simulated population dynamics under a range of habitat conditions. Using a spatially-explicit population model, we removed groups of habitat patches based on their characteristics and measured the resulting time to extinction. We used proportional hazards models to rank the influence of landscape and interacting patch-level variables. Landscape quality was the most influential variable followed by patch quality, with both outweighing landscape- and patch-level measures of habitat quantity and fragmentation/proximity. Although habitat conservation and restoration priorities for this population should be in maximizing the overall quality of the landscape, population persistence depends on how this goal is achieved. Patch quality exerted a significant influence on regional persistence, with the removal of low quality road margin patches (sinks) reducing the risk of regional extinction. Strategies for maximizing overall landscape quality that omit patch-level considerations may produce suboptimal or detrimental results for regional population persistence, particularly where complex local population dynamics (e.g., source-sink dynamics) exist. This study contributes to a growing body literature that suggests that the prediction of species responses and future conservation actions may best be assessed with a multi-scale approach that considers habitat quality and that the success of conservation actions may depend on assessing the influences of habitat factors at multiple scales.     

Metacommunity,mainland‐island system or island communities? Assessing the regional dynamics of plant communities in a fragmented landscape

Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among‐patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape‐scale. In this study, we used extensive field data from a fragmented, semi‐arid landscape in Israel to parameterize a multi‐species incidence‐function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics – the metacommunity, the mainland‐island, or the island communities type – best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch‐matrix study landscape is best represented as a system of highly isolated ‘island’ communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33–60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.     

Habitat quality of source patches and connectivity in fragmented landscapes

Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.     

Single large AND several small habitat patches: A community perspective on their importance for biodiversity

The debate whether single large or several small (SLOSS) patches benefit biodiversity has existed for decades, but recent literature provides increasing evidence for the importance of small habitats. Possible beneficial mechanisms include reduced presence of predators and competitors in small habitat areas or specific functions such as stepping stones for dispersal. Given the increasing amount of studies highlighting individual behavioral differences that may influence these functions, we hypothesize that the advantage of small versus large habitat patches not only depends on patch functionality but also on the presence of animal personalities (i.e., risk-tolerant vs. risk-averse). Using an individual-based, spatially-explicit community model, we analyzed the diversity of mammal communities in landscapes consisting of a few large habitat islands interspersed with different amounts and sizes of small habitat patches. Within these heterogeneous environments, individuals compete for resources and form home-ranges, with only risk-tolerant individuals using habitat edges. Results show that when risk-tolerant individuals exist, small patches increase species diversity. A strong peak occurs at approximately 20% habitat cover in small patches when those small habitats are only used for foraging but not for breeding and home-range core position. Additional usage as stepping stones for juvenile dispersal further increases species persistence. Overall, our results reveal that a combination of a few large and several small habitat patches promotes biodiversity by enhancing landscape heterogeneity. Here, heterogeneity is created by pronounced differences in habitat functionality, increasing edge density, and variability in habitat use by different behavioral types. The finding that a combination of single large AND several small (SLASS) patches is needed for effective biodiversity preservation has implications for advancing landscape conservation. Particularly in structurally poor agricultural areas, modern technology enables precise management with the opportunity to create small foraging habitats by excluding less profitable agricultural land from cultivation.     

Temporal patch occupancy dynamics of the Siberian flying squirrel in a boreal forest landscape

Ability to predict species distribution in a landscape is of crucial importance for natural resource management and species conservation. Therefore, the understanding of species habitat requirements and spatio-temporal dynamics in occurrence is needed. We examined patch occupancy patterns of the Siberian flying squirrel Pteromys volans in northern Finland across a seven year study period. Forest patches dominated by mature spruce ( Picea abies ) in a study area (375 km²) were surveyed to monitor the presence or absence of the flying squirrel. The patch occupancy pattern was dynamic: about half of the habitat patches were occupied at least once during the study period and more patches were colonised than were abandoned. Patches that were continuously occupied (i.e. occupied during all sample periods) were typically of high quality (based on habitat and landscape characteristics), continuously unoccupied patches were usually of low quality, and intermediate quality patches were occupied intermittently. The variables explaining patch occupancy were similar each year, and a statistical model based on data from the year 2000 also predicted occupancy in 2004 with similar accuracy. However, data from a single survey were inadequate for identifying patches used intermittently by flying squirrels. Despite inconsistent occupancy, these patches may be important for the local persistence of flying squirrels. The dynamic occupancy pattern may thus affect estimates of suitable habitat area and identification of functional patch networks for landscape planning. These results emphasise the need for follow-up studies to better understand population patterns and processes in time.     

Extinction–immigration dynamics lag behind environmental filtering in shaping the composition of tropical dry forests within a changing landscape

The impact of rapid habitat loss and fragmentation on biodiversity is a major issue. However, we still lack an integrative understanding of how these changes influence biodiversity dynamics over time. In this study, we investigate the effects of these changes in terms of both niche-based and neutral dynamics. We hypothesize that habitat loss has delayed effects on neutral immigration–extinction dynamics, while edge effects and environmental heterogeneity in habitat patches have rapid effects on niche-based dynamics. We analyzed taxonomic and functional composition of 100 tree communities in a tropical dry forest landscape of New-Caledonia subject to habitat loss and fragmentation. We designed an original, process-based simulation framework, and performed Approximate Bayesian Computation to infer the influence of niche-based and neutral processes. Then, we performed partial regressions to evaluate the relationships between inferred parameter values of communities and landscape metrics (distance to edge, patch area, and habitat amount around communities), derived from either recent or past (65 yr ago) aerial photographs, while controlling for the effect of soil and topography. We found that landscape structure influences both environmental filtering and immigration. Immigration rate was positively related to past habitat amount surrounding communities. In contrast, environmental filtering was mostly affected by present landscape structure and mainly influenced by edge vicinity and topography. Our results highlight that landscape changes have contrasting spatio-temporal influences on niche-based and neutral assembly dynamics. First, landscape-level habitat loss and community isolation reduce immigration and increase demographic stochasticity, resulting in slow decline of local species diversity and extinction debt. Second, recent edge creation affects environmental filtering, incurring rapid changes in community composition by favoring species with edge-adapted strategies. Our study brings new insights about temporal impacts of landscape changes on biodiversity dynamics. We stress that landscape history critically influences these dynamics and should be taken into account in conservation policies.     

Island theory, matrix effects and species richness patterns in habitat fragments

Island biogeography theory, created initially to study diversity patterns on islands, is often applied to habitat fragments. A key but largely untested assumption of this application of theory is that landscape matrix species composition is non‐overlapping with that of the islands. We tested this assumption in successional old field patches in a closely mowed matrix, and because our patches are appropriately viewed as sets of contiguous habitat units we studied patterns of species richness per unit area. Previous studies at our site did not find that diversity patterns on patch ‘islands’ conformed to predictions of island biogeography theory. Our results indicate that when matrix species are removed from the patch samples, diversity patterns conform better to theory. We suggest that classical island theory remains an appropriate tool to study diversity patterns in fragmented habitats, but that allowances should be made for spill‐over colonization of ‘islands’ from the ‘sea’.     

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.     

Persistence and vulnerability: Retaining biodiversity in the landscape and in protected areas

An objective of biodiversity conservation activities is to minimize the exposure of biodiversity features to threatening processes and to ensure, as far as possible, that biodiversity persists in the landscape. We discuss how issues of vulnerability and persistence can and should be addressed at all stages of the conservation planning and implementation process. Procedures for estimating the likelihood of persistence and for measuring degrees of vulnerability at different spatial and temporal scales using subjective assessments, rules of thumb and analytical and simulation models are reviewed. The application of information on vulnerability and persistence to conservation planning and management is discussed under the headings of natural dynamics, replication of protection, levels of representation, source and sink population structures, refuges and critical resources, reserve design, habitat fragmentation and levels of management.     

基于MSPA与最小路径方法的巴中西部新城生态网络构建

目前快速城市化导致了生境斑块的日益破碎化,景观之间的连通性不断降低。构建生态网络可以连接破碎的生境斑块,增加绿地景观的连通性,对生物多样性保护具有重要意义。以高度景观破碎化的四川省巴中西部新城为研究区,采用形态学空间格局分析(MSPA)方法,提取出对研究区生态网络构建具有重要生态意义的核心区和桥接区两类景观要素,并选用整体连通性(IIC)、可能连通性(PC)和斑块重要性(d I)等景观指数,分别对核心区和桥接区进行景观连接度评价,遴选出对维持景观连通性贡献最大的10个核心区生境斑块作为生态网络的源地,并根据斑块对维持景观连通的重要性程度将其他核心区和桥接区进行类型划分,以此作为景观阻力的赋值依据,融入消费面模型中,最后采用最小路径方法构建了研究区潜在的生态网络,并基于重力模型对重要生态廊道进行了识别与提取,在此基础上有针对性地提出了生态网络优化的对策。研究结果表明,MSPA方法能够科学的辨识出研究区内对生态保护具有重要意义的结构性要素,例如作为物种栖息地的核心区和物种迁移通道的桥接区,这些要素是生态网络的重要组成部分;景观连通性的计算,明确了研究区景观要素的保护重点,为最小路径方法中的景观阻力赋值提供了重要的参考信息;基于MSPA与最小路径方法的生态网络分析框架综合了现有景观结构性要素识别、连通性分析以及物种潜在迁移路径分析等方法,将景观中潜在的生态源地和结构性廊道的连通性作为构建生态网络的重要基础和主要依据,从而使得生态网络的构建更科学。研究结果可为高度破碎化地区生态网络的构建提供重要的参考与依据,对其他地区生态网络的构建也具有一定的借鉴意义。     

Habitat selection behaviour links local and regional scales in aquatic systems

The role of habitat selection behaviour in the assembly of natural communities is an increasingly important theme in ecology. At the same time, ecologists and conservation biologists are keenly interested in scale and how processes at scales from local to regional interact to determine species distributions and patterns of biodiversity. How important is habitat selection in generating observed patterns of distribution and diversity at multiple spatial scales? In theory, habitat selection in response to interacting species can generate both positive and negative covariances among species distributions and create the potential to link processes of community assembly across multiple scales. Here I demonstrate that habitat selection by treefrogs in response to the distribution of fish predators functions at both the regional scale among localities and the local scale among patches within localities, implicating habitat selection as a critical link between local communities and the regional dynamics of metacommunities in complex landscapes.     

Metapopulation theory for fragmented landscapes

We review recent developments in spatially realistic metapopulation theory, which leads to quantitative models of the dynamics of species inhabiting highly fragmented landscapes. Our emphasis is in stochastic patch occupancy models, which describe the presence or absence of the focal species in habitat patches. We discuss a number of ecologically important quantities that can be derived from the full stochastic models and their deterministic approximations, with a particular aim of characterizing the respective roles of the structure of the landscape and the properties of the species. These quantities include the threshold condition for persistence, the contributions that individual habitat patches make to metapopulation dynamics and persistence, the time to metapopulation extinction, and the effective size of a metapopulation living in a heterogeneous patch network.     

Spatially structured metapopulation models: global and local assessment of metapopulation capacity

We model metapopulation dynamics in finite networks of discrete habitat patches with given areas and spatial locations. We define and analyze two simple and ecologically intuitive measures of the capacity of the habitat patch network to support a viable metapopulation. Metapopulation persistence capacity lambda(M) defines the threshold condition for long-term metapopulation persistence as lambda(M)>delta, where delta is defined by the extinction and colonization rate parameters of the focal species. Metapopulation invasion capacity lambda(I) sets the condition for successful invasion of an empty network from one small local population as lambda(I)>delta. The metapopulation capacities lambda(M) and lambda(I) are defined as the leading eigenvalue or a comparable quantity of an appropriate "landscape" matrix. Based on these definitions, we present a classification of a very general class of deterministic, continuous-time and discrete-time metapopulation models. Two specific models are analyzed in greater detail: a spatially realistic version of the continuous-time Levins model and the discrete-time incidence function model with propagule size-dependent colonization rate and a rescue effect. In both models we assume that the extinction rate increases with decreasing patch area and that the colonization rate increases with patch connectivity. In the spatially realistic Levins model, the two types of metapopulation capacities coincide, whereas the incidence function model possesses a strong Allee effect characterized by lambda(I)=0. For these two models, we show that the metapopulation capacities can be considered as simple sums of contributions from individual habitat patches, given by the elements of the leading eigenvector or comparable quantities. We may therefore assess the significance of particular habitat patches, including new patches that might be added to the network, for the metapopulation capacities of the network as a whole. We derive useful approximations for both the threshold conditions and the equilibrium states in the two models. The metapopulation capacities and the measures of the dynamic significance of particular patches can be calculated for real patch networks for applications in metapopulation ecology, landscape ecology, and conservation biology.     

基于图论的两栖类生物栖息地网络规划——以黑斑侧褶蛙为例

生物多样性保护和生物栖息地网络建设是目前我国国土空间规划的重要内容,提升生物栖息地网络的景观功能连接度对生物多样性保护具有重要作用。目前,已有研究对生物栖息地网络规划进行了探索,但在实际规划层面仍缺乏可操作性强的技术方法支撑。本研究采用图论方法,聚焦国土空间规划中生物多样性保护和生态网络建设涉及的进行生物栖息地斑块重要性评价,以确定斑块优先保护次序;寻找最优新增斑块位置,以改善生物栖息地网络景观功能连接度;依据景观功能连接度的降低程度,判断建设项目的影响或评价规划新增建设项目的潜在影响3方面内容,在雄安新区两栖类生物黑斑侧褶蛙(Pelophylax nigromaculata)栖息地网络规划中进行应用研究。结果表明: 图论方法可以有效解决上述3方面的问题;本研究识别的5个最优新增黑斑侧褶蛙栖息地位置使栖息地网络整体景观功能连接度提升19%;通过评价G45高速公路对两栖类生物栖息地网络功能连接度的影响,找出了4个穿越通道设置以减弱G45高速公路的影响。     

Landscape-level comparison of genetic diversity and differentiation in a small mammal inhabiting different fragmented landscapes of the Brazilian Atlantic Forest

Habitat loss and fragmentation can have detrimental effects on all levels of biodiversity, including genetic variation. Most studies that investigate genetic effects of habitat loss and fragmentation focus on analysing genetic data from a single landscape. However, our understanding of habitat loss effects on landscape-wide patterns of biodiversity would benefit from studies that are based on quantitative comparisons among multiple study landscapes. Here, we use such a landscape-level study design to compare genetic variation in the forest-specialist marsupial Marmosops incanus from four 10,000-hectare Atlantic forest landscapes which differ in the amount of their remaining native forest cover (86, 49, 31, 11 %). Additionally, we used a model selection framework to evaluate the influence of patch characteristics on genetic variation within each landscape. We genotyped 529 individuals with 12 microsatellites to statistically compare estimates of genetic diversity and genetic differentiation in populations inhabiting different forest patches within the landscapes. Our study indicates that before the extinction of the specialist species (here in the 11 % landscape) genetic diversity is significantly reduced in the 31 % landscape, while genetic differentiation is significantly higher in the 49 and 31 % landscapes compared to the 86 % landscape. Results further provide evidence for non-proportional responses of genetic diversity and differentiation to increasing habitat loss, and suggest that local patch isolation impacts gene flow and genetic connectivity only in the 31 % landscape. These results have high relevance for analysing landscape genetic relationships and emphasize the importance of landscape-level study designs for understanding habitat loss effects on all levels of biodiversity.     

Effect of Landscape Pattern on Insect Species Density within Urban Green Spaces in Beijing,China

Urban green space is an important refuge of biodiversity in urban areas. Therefore, it is crucial to understand the relationship between the landscape pattern of green spaces and biodiversity to mitigate the negative effects of urbanization. In this study, we collected insects from 45 green patches in Beijing during July 2012 using suction sampling. The green patches were dominated by managed lawns, mixed with scattered trees and shrubs. We examined the effects of landscape pattern on insect species density using hierarchical partitioning analysis and partial least squares regression. The results of the hierarchical partitioning analysis indicated that five explanatory variables, i.e., patch area (with 19.9% independent effects), connectivity (13.9%), distance to nearest patch (13.8%), diversity for patch types (11.0%), and patch shape (8.3%), significantly contributed to insect species density. With the partial least squares regression model, we found species density was negatively related to patch area, shape, connectivity, diversity for patch types and proportion of impervious surface at the significance level of p < 0.05 and positively related to proportion of vegetated land. Regression tree analysis further showed that the highest species density was found in green patches with an area <500 m². Our results indicated that improvement in habitat quality, such as patch area and connectivity that are typically thought to be important for conservation, did not actually increase species density. However, increasing compactness (low-edge) of patch shape and landscape composition did have the expected effect. Therefore, it is recommended that the composition of the surrounding landscape should be considered simultaneously with planned improvements in local habitat quality.