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1.
Context
The relative importance of habitat area and connectivity for species richness is often unknown. Connectivity effects may be confounded with area effects or they may be of minor importance as posited by the habitat-amount hypothesis.Objectives
We studied effects of habitat area and connectivity of linear landscape elements for plant species richness at plot level. We hypothesized that connectivity of linear landscape elements, assessed by resistance distance, has a positive effect on species richness beyond the effect of area and, further, that the relative importance of connectivity varies among groups of species with different habitat preferences and dispersal syndromes.Methods
We surveyed plant species richness in 50 plots (25 m2) located on open linear landscape elements (field margins, ditches) in eight study areas of 1 km2 in agricultural landscapes of Northwest Germany. We calculated the area of linear landscape elements and assessed their connectivity using resistance distance within circular buffers (500 m) around the plots. Effects of area and connectivity on species richness were modelled with generalised linear mixed models.Results
Species richness did not increase with area. Resistance distance had significant negative effects on total richness and on the richness of typical species of grasslands and wetlands. Regarding dispersal syndromes, resistance distance had negative effects on the richness of species with short-distance, long-distance and aquatic dispersal. The significant effects of resistance distance indicated that species richness increased with connectivity of the network of linear landscape elements.Conclusions
Connectivity is more important for plant species richness in linear landscape elements than area. In particular, the richness of plant species that are dispersal limited and confined to semi-natural habitats benefits from connective networks of linear landscape elements in agricultural landscapes.2.
Elsa Aggemyr Alistair G. Auffret Love Jädergård Sara A. O. Cousins 《Landscape Ecology》2018,33(12):2273-2284
Context
Understanding how landscape patterns affect species diversity is of great importance in the fields of biogeography, landscape ecology and conservation planning, but despite the rapid advance in biodiversity analysis, investigations of spatial effects on biodiversity are still largely focused on species richness.Objectives
We wanted to know if and how species richness and species composition are differentially driven by the spatial measures dominating studies in landscape ecology and biogeography. As both measures require the same limited presence/absence information, it is important to choose an appropriate diversity measure, as differing results could have important consequences for interpreting ecological processes.Methods
We recorded plant occurrences on 112 islands in the Baltic archipelago. Species richness and composition were calculated for each island, and the explanatory power of island area and habitat heterogeneity, distance to mainland and structural connectivity at three different landscape sizes were examined.Results
A total of 354 different plant species were recorded. The influence of landscape variables differed depending on which diversity measure was used. Island area and structural connectivity determined plant species richness, while species composition revealed a more complex pattern, being influenced by island area, habitat heterogeneity and structural connectivity.Conclusions
Although both measures require the same basic input data, species composition can reveal more about the ecological processes affecting plant communities in fragmented landscapes than species richness alone. Therefore, we recommend that species community composition should be used as an additional standard measure of diversity for biogeography, landscape ecology and conservation planning.3.
Balázs Deák Orsolya Valkó Péter Török András Kelemen Ádám Bede András István Csathó Béla Tóthmérész 《Landscape Ecology》2018,33(7):1117-1132
Context
Landscape and habitat filters are major drivers of biodiversity of small habitat islands by influencing dispersal and extinction events in plant metapopulations.Objectives
We assessed the effects of landscape and habitat filters on the species richness, abundance and trait composition of grassland specialist and generalist plants in small habitat islands. We studied traits related to functional spatial connectivity (dispersal ability by wind and animals) and temporal connectivity (clonality and seed bank persistence) using model selection.Methods
We sampled herbaceous plants, landscape (local and regional isolation) and habitat filters (inclination, woody encroachment and disturbance) in 82 grassland islands in Hungary.Results
Isolation decreased the abundance of good disperser specialist plants due to the lack of directional vectors transferring seeds between suitable habitat patches. Clonality was an effective strategy, but persistent seed bank did not support the survival of specialist plants in isolated habitats. Generalist plants were unaffected by landscape filters due to their wide habitat breadth and high propagule availability. Clonal specialist plants could cope with increasing woody encroachment due to their high resistance against environmental changes; however, they could not cope with intensive disturbance. Steep slopes providing environmental heterogeneity had an overall positive effect on species richness.Conclusions
Specialist plants were influenced by the interplay of landscape filters influencing their abundance and habitat filters affecting species richness. Landscape filtering by isolation influenced the abundance of specialist plants by regulating seed dispersal. Habitat filters sorted species that could establish and persist at a site by influencing microsite availability and quality.4.
Context
Anthropogenic landscape simplification and natural habitat loss can negatively affect wild bees. Alternatively, anthropogenic land-use change may diversify landscapes, creating complementary habitats that maintain overall resource continuity and diversity.Objectives
We examined the effects of landscape composition, including land-cover diversity and percent semi-natural habitat, on wild bee abundance and species richness within apples, a pollinator-dependent crop. We also explored whether different habitats within diverse landscapes can provide complementary floral resources for bees across space and time.Methods
We sampled bees during apple bloom over 2 years within 35 orchards varying in surrounding landscape diversity and percent woodland (the dominant semi-natural habitat) at 1 km radii. To assess habitat complementarity in resource diversity and temporal continuity, we sampled flowers and bees within four unique habitats, including orchards, woodlands, semi-natural grasslands, and annual croplands, over three periods from April–June.Results
Surrounding landscape diversity positively affected both wild bee abundance and richness within orchards during bloom. Habitats in diverse landscapes had different flower communities with varying phenologies; flowers were most abundant within orchards and woodlands in mid-spring, but then declined over time, while flowers within grasslands marginally increased throughout spring. Furthermore, bee communities were significantly different between the closed-canopy habitats, orchards and woodlands, and the open habitats, grasslands and annual croplands.Conclusions
Our results suggest that diverse landscapes, such as ones with both open (grassland) and closed (woodland) semi-natural habitats, support spring wild bees by providing flowers throughout the entire foraging period and diverse niches to meet different species’ requirements.5.
Context
Habitat loss and habitat fragmentation negatively affect amphibian populations. Roads impact amphibian species through barrier effects and traffic mortality. The landscape variable ‘accessible habitat’ considers the combined effects of habitat loss and roads on populations.Objectives
The aim was to test whether accessible habitat was a better predictor of amphibian species richness than separate measures of road effects and habitat loss. I assessed how accessible habitat and local habitat variables determine species richness and community composition.Methods
Frog and tadpole surveys were conducted at 52 wetlands in a peri-urban area of eastern Australia. Accessible habitat was delineated using a highway. Regressions were used to examine relationships between species richness and eleven landscape and local habitat variables. Redundancy analysis was used to examine relationships between community composition and accessible habitat and local habitat variables.Results
Best-ranked models of species richness included both landscape and local habitat variables. There were positive relationships between species richness and accessible habitat and distance to the highway, and uncertain relationships with proportion cover of native vegetation and road density. There were negative relationships between species richness and concreted wetlands and wetland electrical conductivity. Four species were positively associated with accessible habitat, whereas all species were negatively associated with wetland type.Conclusions
Barrier effects caused by the highway and habitat loss have negatively affected the amphibian community. Local habitat variables had strong relationships with species richness and community composition, highlighting the importance of both availability and quality of habitat for amphibian conservation near major roads.6.
Jodie Martin Gwenaël Vourc’h Nadège Bonnot Bruno Cargnelutti Yannick Chaval Bruno Lourtet Michel Goulard Thierry Hoch Olivier Plantard A. J. Mark Hewison Nicolas Morellet 《Landscape Ecology》2018,33(6):937-954
Context
Routine movements of large herbivores, often considered as ecosystem engineers, impact key ecological processes. Functional landscape connectivity for such species influences the spatial distribution of associated ecological services and disservices.Objectives
We studied how spatio-temporal variation in the risk-resource trade-off, generated by fluctuations in human activities and environmental conditions, influences the routine movements of roe deer across a heterogeneous landscape, generating shifts in functional connectivity at daily and seasonal time scales.Methods
We used GPS locations of 172 adult roe deer and step selection functions to infer landscape connectivity. In particular, we assessed the influence of six habitat features on fine scale movements across four biological seasons and three daily periods, based on variations in the risk-resource trade-off.Results
The influence of habitat features on roe deer movements was strongly dependent on proximity to refuge habitat, i.e. woodlands. Roe deer confined their movements to safe habitats during daytime and during the hunting season, when human activity is high. However, they exploited exposed open habitats more freely during night-time. Consequently, we observed marked temporal shifts in landscape connectivity, which was highest at night in summer and lowest during daytime in autumn. In particular, the onset of the autumn hunting season induced an abrupt decrease in landscape connectivity.Conclusions
Human disturbance had a strong impact on roe deer movements, generating pronounced spatio-temporal variation in landscape connectivity. However, high connectivity at night across all seasons implies that Europe’s most abundant and widespread large herbivore potentially plays a key role in transporting ticks, seeds and nutrients among habitats.7.
Context
The habitat amount hypothesis has rarely been tested on plant communities. It remains unclear how habitat amount affect species richness in habitat fragments compared to island effects such as isolation and patch size.Objectives
How do patch size and spatial distribution compared to habitat amount predict plant species richness and grassland specialist plant species in small grassland remnants? How does sampling area affect the prediction of spatial variables on species richness?Methods
We recorded plant species density and richness on 131 midfield islets (small remnants of semi-natural grassland) situated in 27 landscapes in Sweden. Further, we tested how habitat amount, compared to focal patch size and distance to nearest neighbor predicted species density and richness of plants and of grassland specialists.Results
A total of 381 plant species were recorded (including 85 grassland specialist species). A combination of patch size and isolation was better in predicting both density and richness of species compared to habitat amount. Almost 45% of species richness and 23% of specialist species were explained by island biogeography parameters compared to 19 and 11% by the amount of habitat. A scaled sampling method increased the explanation level of island biogeography parameters and habitat amount.Conclusions
Habitat amount as a concept is not as good as island biogeography to predict species richness in small habitats. Priority in landscape planning should be on larger patches rather than several small, even if they are close together. We recommend a sampling area scaled to patch size in small habitats.8.
Kaisa J. Raatikainen Anna Oldén Niina Käyhkö Mikko Mönkkönen Panu Halme 《Landscape Ecology》2018,33(12):2169-2187
Context
Habitat loss is a major threat to biodiversity. It can create temporal lags in decline of species in relation to destruction of habitat coverage. Plant species specialized in semi-natural grasslands, especially meadows, often express such extinction debt.Objectives
We studied habitat loss and fragmentation of meadows and examined whether the changes in meadow coverage had caused an extinction debt on vascular plants. We also studied whether historical or present landscape patterns or contemporary environmental factors were more important determinants of species occurrence.Methods
We surveyed the plant species assemblages of 12 grazed and 12 mown meadows in Central Finland and detected the meadow coverages from their surroundings on two spatial scales and on three time steps. We modelled the effects of functional connectivity, habitat amount, and isolation on species richness and community composition.Results
We observed drastic and dynamic meadow loss in landscapes surrounding our study sites during the last 150 years. However, we did not find explicit evidence for an extinction debt in meadow plants. The observed species richness correlated with contemporary factors, whereas both contemporary factors and habitat availability during the 1960s affected community composition.Conclusions
Effective conservation management of meadow biodiversity builds on accurate understanding of the relative importance of past and present factors on species assemblages. Both mown and grazed meadows with high species richness need to be managed in the future. The management effort should preferably be targeted to sites located near to each other.9.
Katherina Ng Philip S. Barton Sarina Macfadyen David B. Lindenmayer Don A. Driscoll 《Landscape Ecology》2018,33(1):109-125
Context
Farming practices influence the degree of contrast between adjoining habitats, with consequences for biodiversity and species movement. Little is known, however, on insect community responses to different kinds of edges over time, and the extent of cross-habitat movement in agricultural landscapes.Objective
To determine temporal changes in beetle responses to different farmland-woodland edges, and document cross-habitat movement.Methods
We examined species richness, abundance, and movement across edges between remnant woodlands and four farmland uses (plantings, fallow, annual crops, woody debris applied over crops post-harvest) in southeastern Australia. We used directional pitfall traps to infer movement, and sampled at edges, and 20 and 200 m on both sides of edges, during spring and summer.Results
Detritivore and predator abundance varied between seasons across the edge between woodlands and all farmlands, but seasonal differences were weaker for fallow-woodland and woody debris-woodland edges. Detritivores moved from farmlands towards woodlands, but not across fallow-woodlands and woody debris-woodlands edges during summer. During summer, predators showed short-range movement towards edges from all farmlands except plantings, and towards woody debris from woodlands. Edges showed temporally stable predator richness and higher herbivore richness than adjoining habitats.Conclusions
Farmland use and season interactively affect beetle abundance across farmland-woodland edges. Woody debris can reduce seasonal fluctuations in beetle edge responses and increase permeability for cross-habitat movement, while plantings provide habitat during summer. Edges provide important resources for beetles in adjoining habitats, however, seasonal movement of predators specifically into edges may affect prey assemblages—a link requiring further study.10.
Context
A recent hypothesis, the habitat amount hypothesis, predicts that the total amount of habitat in the landscape can replace habitat patch size and isolation in studies of species richness in fragmented landscapes.Objectives
To test the habitat amount hypothesis by first evaluating at which spatial scale the relationship between species richness in equal-sized sample quadrats and habitat amount was the strongest, and then test the importance of spatial configuration of habitat—measured as local patch size and isolation—when habitat amount was taken into account.Methods
A quasi-experimental setup with 20 habitat patches of dry calcareous grasslands varying in patch size, patch isolation and habitat amount at the landscape scale was established in the inner Oslo fjord, Southern Norway. We recorded species richness of habitat specialists of vascular plants in equal-sized sample quadrats and analysed the relationship between species richness, habitat amount in the landscape and patch size and isolation.Results
Although the total amount of habitat in a 3 km-radius around the local patch was positively related to species richness in the sample quadrats, local patch size had an additional positive effect, and the effect of patch size was higher when the amount of habitat within the 3 km-radius was high than when it was low.Conclusions
In our study system of specialist vascular plants in dry calcareous grasslands, we do not find support for the habitat amount hypothesis.11.
Olivia Dondina Valerio Orioli Lorenza Colli Massimiliano Luppi Luciano Bani 《Landscape Ecology》2018,33(2):275-287
Context
Ecological networks are often designed based on the degree of suitability and permeability of land cover classes, as obtained by estimating the statistical relationships between occurrence data and classes coverage using habitat suitability models (HSMs). Considering only the classes coverage, but not their spatial arrangement, frequently prevents HSMs from correctly identifying nodes and connectivity elements.Objectives
We propose a new approach in the design of ecological networks starting from the relationship between occurrence data and both land cover classes coverage and spatial arrangement, as calculated for different simulated species perceptions of the landscape (SSPLs, corresponding to different combinations of classes alternatively assuming the role of nodes, connectivity elements, or matrix).Methods
The approach consists of comparing the ability to explain the observed species occurrence of both the nodes coverage and the connectivity degree provided by both nodes and connectivity elements, calculated for each SSPL. The better performing SSPL will provide information about the land cover classes that should be considered in designing an ecological network for the species, as well as their role in the network.Results
When applied to the Hazel Dormouse in an agricultural landscape in northern Italy, the method proved effective and allowed us to identify woodlands and hedgerows as nodes, and poplar cultivations, biomasses and reforestations as connectivity elements.Conclusions
The proposed method can be adopted to identify nodes and connectivity elements for virtually every species sensitive to fragmentation, and has important practical implications when integrated in landscape management plans developed to guarantee ecological connectivity.12.
Context
Theory predicts that habitat loss and fragmentation may have drastic consequences on species’ interactions. To date, however, little empirical evidence exists on the strength of interspecific competition in shaping animal communities in fragmented landscapes.Objectives
Our aim was to measure the degree of ongoing competitive interference between species in fragmented landscapes. Our model system was the community of ground-dwelling rodents in deciduous woodlands in central Italy, composed of a habitat generalist species (Apodemus sylvaticus) and two forest specialists (Apodemus flavicollis and Myodes glareolus). Our objectives were to test whether species were segregated among forest patches and whether spatial segregation was determined by interspecific competition or habitat and resource availability.Methods
We surveyed the populations inhabiting 29 woodland patches in a highly fragmented landscape using a capture-mark-recapture protocol, capturing >4500 individuals. First we modelled species’ distribution as a function of habitat, resource availability and landscape variables. The second stage of our analyses involved measuring the response of vital rate parameters (body mass, reproduction, survival, recruitment, population density) to competitor density.Results
The relative distribution of species reflected a spatial segregation of habitat generalists and specialists according to habitat quality, cover and connectivity. Interspecific competition mainly affected individual level vital rates, whereas we found no substantial effects at the population level.Conclusions
Competitive exclusion of specialist species by generalist species was occurring. However, when compared to other factors such as habitat connectivity and resource availability, interspecific competition played a relatively minor role in shaping the studied community.13.
Context
According to the trophic-rank hypothesis, species may be differentially affected by habitat isolation due to their trophic position in the food chain, i.e. high-order trophic levels may be more negatively affected than low-order levels.Objectives
The aim of this paper is to study how species richness, abundance and composition of saproxylic beetle communities are affected by patch (=tree) quality and small-scale patch connectivity. Following the trophic-rank hypothesis, we expected predators to be more negatively affected by patch isolation than wood-feeding beetles.Methods
We studied the beetle community, patch connectivity and patch quality on 28 large oaks. Different connectivity measures were calculated using 50 m-buffers around trees and using distances to the five nearest trees.Results
Beetle species richness increased with the diameter of oaks, i.e. patch quality. No evidence of the trophic-rank hypothesis was found for species richness patterns. In accordance with the trophic-rank hypothesis, abundance of predatory beetles increased with patch connectivity but lower trophic levels were unaffected or even decreased with patch connectivity.Conclusions
The structure of invertebrate communities on trees changes with small-scale patch connectivity due to a differential response of low-order and high-order trophic levels. Isolated trees are more exposed to the sun than the more connected trees, which may affect the beetles; however, it was impossible to distinguish the microclimatic from the spatial effects. Although scattered trees generally have a higher conservation value than trees in forests, we conclude that forest trees may be more important for certain trophic levels.14.
Milena F. Diniz Ricardo B. Machado Arthur A. Bispo Paulo De M. Júnior 《Landscape Ecology》2018,33(11):1911-1923
Context
The umbrella approach applied to landscape connectivity is based on the principle that the conservation or restoration of the dispersal habitats for some species also can facilitate the movement of others. Species traits alone do not seem to be enough to identify good connectivity umbrella species, showing the need to investigate the influence of additional factors on this property.Objectives
We test whether the potential of a species as a connectivity umbrella can be influenced by landscape composition and configuration.Methods
We simulated movement routes for eight hypothetical species in artificial patchy landscapes with different levels of fragmentation, habitat amount and matrix permeability. We determined the effectiveness of the connectivity umbrella of the virtual species using pairwise intersections of important habitats for their movements in all landscapes.Results
The connectivity umbrella performance of all species was affected by the interaction of fragmentation level and habitat amount. In general, species performance increased with decreasing fragmentation and increasing habitat amount. In most landscapes and considering the same dispersal threshold, species able to move more easily through the matrix showed higher umbrella performance than those for which the matrix offered greater resistance.Conclusions
The connectivity umbrella is not a static feature that depends only on the species traits, but rather a dynamic property that also varies according to the landscape attributes. Therefore, we do not recommend spatial transferability of the connectivity umbrella species identified in a landscape to others that have divergent levels of fragmentation and habitat quantity.15.
Context
Butterflies have been continuously declining for several decades in Europe due to many factors, such as farming intensification. Rural landscapes have undergone dramatic changes leading to homogenized landscapes.Objectives
In this study, we investigated how landscape composition, structure and connectivity impact butterfly communities according to their ecological and biological traits.Methods
We made use of 5669 Lepidoptera surveys performed at 4525 distinct locations in lowland Central France. We considered 19 ecological groups based on habitat specialization, mobility, diet, voltinism or overwintering strategy. Generalized linear mixed-effect models were used to relate the species richness of these groups to landscape variables defined in circular zones with radius from 250 m to 5 km.Results
Richness of most species groups co-varied with landscape variables, with the exception of mobile, imago-overwintering, monophagous and polyphagous species. Habitat proportion explained more variation in butterfly diversity than habitat connectivity or habitat diversity. Moreover, the best proportion models were generally found for the 250-m circular zones. Thirteen species groups were disfavored by cropland amount. Except for forest specialists and high mobility group, no other group was more diverse in landscapes dominated by a single land cover type. Rather, for total diversity and 14 groups, species richness peaked for forest proportions varying between 40 and 80%, and for total diversity and nine groups for grassland proportions ranging from 30 to 60%.Conclusions
These results indicate that landscape homogenization is contributing to the ongoing decline in butterflies, and support preserving and (re)creating mosaics of grasslands and forests.16.
Christina A. Buelow Ronald Baker April E. Reside Marcus Sheaves 《Landscape Ecology》2017,32(3):547-561
Context
Complex structural connectivity patterns can influence the distribution of animals in coastal landscapes, particularly those with relatively large home ranges, such as birds. To understand the nuanced nature of coastal forest avifauna, where there may be considerable overlap in assemblages of adjacent forest types, the concerted influence of regional landscape context and vegetative structural connectivity at multiple spatial scales warrants investigation.Objectives
This study determined whether species compositions of coastal forest bird assemblages differ with regional landscape context or with forest type, and if this is influenced by structural connectivity patterns measured at multiple spatial scales.Methods
Three replicate bird surveys were conducted in four coastal forest types at ten survey locations across two regional landscape contexts in northeast Australia. Structural connectivity patterns of 11 vegetation types were quantified at 3, 6, and 12 km spatial scales surrounding each survey location, and differences in bird species composition were evaluated using multivariate ordination analysis.Results
Bird assemblages differed between regional landscape contexts and most coastal forest types, although Melaleuca woodland bird assemblages were similar to those of eucalypt woodlands and rainforests. Structural connectivity was primarily correlated with differences in bird species composition between regional landscape contexts, and correlation depended on vegetation type and spatial scale.Conclusions
Spatial scale, landscape context, and structural connectivity have a combined influence on bird species composition. This suggests that effective management of coastal landscapes requires a holistic strategy that considers the size, shape, and configuration of all vegetative components at multiple spatial scales.17.
18.
Else Foster Jamie Love Romina Rader Nick Reid Michael J. Drielsma 《Landscape Ecology》2017,32(9):1837-1847
Context
A challenge devising revegetation strategies in fragmented landscapes is conserving for the widest spectrum of biodiversity. Habitat network reconstruction should improve landscape capacity to maintain species populations. However, the location of revegetation often fails to account for species occurrence and dispersal processes operating across spatial scales.Objectives
Our objective was to integrate metapopulation theory with estimates of landscape capacity and dispersal pathways to highlight connectivity gaps. Maintenance of populations could thereby be facilitated through reconnecting habitat networks across regional and broader scales, with assumed benefit for the dispersal needs of less sensitive species.Methods
Predicted occupancy and metapopulation capacity were calculated for a generic focal species derived from fragmentation-sensitive woodland birds, mammals and reptiles. A metapopulation connectivity analysis predicted regional dispersal links to identify likely routes through which individuals may move to contribute to the viability of the population. We used the revegetation programmes of the Brigalow–Nandewar Biolinks project, eastern New South Wales, Australia, to demonstrate our approach.Results
Landscape capacity of the current landscape varied across the region. Low-value links between populations provided greatest opportunities for revegetation and improved landscape capacity. Where regional connectivity did not indicate a pathway between populations, broader scale connectivity provided guidance for revegetation.Conclusions
The metapopulation-based model, coupled with a habitat dispersal network analysis, provided a platform to inform revegetation locations and better support biodiversity. Our approach has application for directing on-ground action to support viable populations, assess the impact of revegetation schemes or monitor the progress of staged implementations.19.
Charles A. Martin 《Landscape Ecology》2018,33(11):1831-1835
Context
The ecological literature is filled with studies highlighting the importance of both habitat loss and fragmentation on biodiversity. The patch concept has been central to these findings, being also at the heart of many ecological theories. Recently, the habitat amount hypothesis has been proposed as an alternative, where the patch concept is put to a rest, and both patch size and patch isolation effects on species richness are reduced to a single gradient: habitat loss in the landscape.Objectives
As this theory stated clear predictions that could be experimentally tested, many formal tests of the hypothesis have been published recently and this study aims at synthesizing their results.Methods
A meta-analysis of 13 tests of the habitat amount hypothesis was conducted, to produce a single combined test of the theory.Results
The 13 tests combined suggest that effects of patch size and isolation, while controlling for habitat amounts, do exist although their overall effect is weak (r?=?0.158).Conclusions
Literal interpretations of the habitat amount hypothesis, where patch size and isolation have absolutely no effect on species richness, are probably oversimplifications of the processes at work. Still, the theory could prove useful as a baseline of the effects of habitat loss, against which patch size and isolation effects must be contrasted.20.
Jessica L. Neumann Geoffrey H. Griffiths Andrew Hoodless Graham J. Holloway 《Landscape Ecology》2016,31(2):301-315