Green Veining: Landscape Determinants of Biodiversity in European Agricultural Landscapes

Many semi-natural landscape elements, the so-called green veining, are disappearing from the intensively used agricultural landscapes of Europe. In order to develop or restore biodiversity in these networks, it is necessary to quantify the relation between biodiversity and amount, spatial arrangement and management intensity of green veining elements. In this review, we investigate whether biodiversity increases with the amount of green veining in an agricultural landscape following the species–area relationship, and whether a certain level of biodiversity can be reached at lower densities of green veining if green veining elements are better connected (higher spatial connectivity) or if they are managed less intensively (lower management intensity). We reviewed studies on aboveground biodiversity in green veining structures in 39 scientific papers on field and experimental studies within Europe. More of these studies focussed on management intensity than on amount or spatial configuration of green veining. Also more studies focussed on the spatial scale of individual landscape elements than on the farm or landscape scale, which may be caused by the large number of studies focussing on plant or invertebrate species. Species living at larger spatial scales, e.g. mammals and birds were not often studied at the level of green veining elements as they also use agricultural fields as part of their habitat. We could not verify the species–area relation for green veining, nor the effect of amount, spatial configuration or management intensity on this relation, because only few studies quantified the found effects and no studies were found on the effect of management intensity or spatial configuration on the species–area curve in green veining. We addressed the most important challenges for future field and model research in order to fill the identified gaps in knowledge.     

Sacrificing patches for linear habitat elements enhances metapopulation performance of woodland birds in fragmented landscapes

It is generally assumed that large patches of natural habitat are better for the survival of species than the same amount of habitat in smaller fragments or linear elements like hedges and tree rows. We use a spatially explicit individual-based model of a woodland bird to explore this hypothesis. We specifically ask whether mixtures of large, small and linear habitat elements are better for population performance than landscapes that consist of only large elements. With equal carrying capacity, metapopulations perform equally or better in heterogeneous landscape types that are a mix of linear, large and small habitat elements. We call this increased metapopulation performance of large and small elements “synergy”. These mixed conditions are superior because the small linear elements facilitate dispersal while patches secure the population in the long run because they have a lower extinction risk. The linear elements are able to catch and guide dispersing animals which results in higher connectivity between patches leading to higher metapopulation survival. Our results suggest that landscape designers should not always seek to conserve and create larger units but might better strive for more variable landscapes with mixtures of patch sizes and shapes. This is especially important when smaller units play a key role in connecting patches and dispersal through the matrix is poor.     

Extinction risk of wood-living model species in forest landscapes as related to forest history and conservation strategy

Dead wood is a critical resource for biodiversity in boreal forests. We analysed the persistence of five model species inhabiting dead wood. By parameterising a metapopulation model (the incidence function model), the model species were all assigned characteristics that makes it likely that they have disappeared from some (20%) forest landscapes with a long history of forest management. In the metapopulation model, a forest stand (5 ha) was regarded as a habitat patch. The amount of habitat in each patch was obtained from models of dead wood dynamics of Norway spruce in central Sweden. Dead wood generated by altered management over the entire landscape was found to be less efficient in reducing extinction risks in comparison to the same amount of dead wood generated by protecting reserves. Because generation of dead wood by altered management is often less expensive than setting aside reserves, it is difficult to determine which conservation measure is most cost-efficient. In a landscape subjected to forestry for the first time, it was better to preserve a few large reserves than many small ones. However, in a managed, highly fragmented forest landscape it was better to set aside many small reserves. The reason for this was that small plots with high habitat quality could be selected, while large reserves originally contained habitats both of high and low quality, and the rate of habitat quality increase was low. A strategy for biodiversity conservation in a managed forest landscape should include information about the history of the landscape, the current amount and spatial distribution of forest habitats, and the potential for rapid restoration of forest habitats, both on managed and unmanaged forest land.     

Functional rather than structural connectivity explains grassland plant diversity patterns following landscape scale habitat loss

Context

Functional connectivity is vital for plant species dispersal, but little is known about how habitat loss and the presence of green infrastructure interact to affect both functional and structural connectivity, and the impacts of each on species groups.

Objectives

We investigate how changes in the spatial configuration of species-rich grasslands and related green infrastructure such as road verges, hedgerows and forest borders in three European countries have influenced landscape connectivity, and the effects on grassland plant biodiversity.

Methods

We mapped past and present land use for 36 landscapes in Belgium, Germany and Sweden, to estimate connectivity based on simple habitat spatial configuration (structural connectivity) and accounting for effective dispersal and establishment (functional connectivity) around focal grasslands. We used the resulting measures of landscape change to interpret patterns in plant communities.

Results

Increased presence of landscape connecting elements could not compensate for large scale losses of grassland area resulting in substantial declines in structural and functional connectivity. Generalist species were negatively affected by connectivity, and responded most strongly to structural connectivity, while functional connectivity determined the occurrence of grassland specialists in focal grasslands. Restored patches had more generalist species, and a lower density of grassland specialist species than ancient patches.

Conclusions

Protecting both species rich grasslands and dispersal pathways within landscapes is essential for maintaining grassland biodiversity. Our results show that increases in green infrastructure have not been sufficient to offset loss of semi-natural habitat, and that landscape links must be functionally effective in order to contribute to grassland diversity.

     

Habitat structure mediates spatial segregation and therefore coexistence

Understanding the mechanisms driving diversity in nature is an important and ongoing challenge in our changing world. To efficiently protect ecosystem diversity it is crucial to explain why and how species coexist. Over the last decades models explaining species coexistence have increased in complexity but usually don’t incorporate a detailed spatial context. However, spatial structure has been shown to affect species coexistence and habitat deterioration is one of the biggest threats to biodiversity. We therefore explore a spatially explicit two-species model and assess the effects of habitat structure on species coexistence using a wide diversity of fractal landscapes. Each species is specialized in a particular habitat type. We find that landscape structure has a major influence on the stability and constitution of a two species system and may be sufficient to explain the coexistence of two species. Well connected and highly structured habitat configurations allow spatial segregation of both species and this decreases local interspecific competition; in our model this is the most important process stabilizing coexistence.     

Landscape cohesion: an index for the conservation potential of landscapes for biodiversity

In urbanising landscapes, planning for sustainable biodiversity occurs in a context of multifunctional land use. Important conditions for species persistence are habitat quality, the amount and configuration of habitat and the permeability of the landscape matrix. For planning purposes, these determinants should be integrated into simple indicators for spatial conditions of persistence probability. We propose a framework of three related indices. The cohesion index is based on the ecology of metapopulations in a habitat network. We discuss how an indicator for species persistence in such a network could be developed. To translate this network index into an area index, we propose the concept of spatial cohesion. Habitat cohesion and spatial cohesion are defined and measured for single species or, at best, for species profiles. Since species differ in their perception of the same landscape, different species will rate different values of these indices for the same landscape. Because landscapes are rarely planned for single species, we further propose the index of landscape cohesion, which integrates the spatial cohesion indices of different species. Indices based on these concepts can be built into GIS tools for landscape assessment. We illustrate different applications of these indices, and emphasise the distinction between ecological and political decisions in developing and applying such tools. This revised version was published online in August 2006 with corrections to the Cover Date.     

Quantifying determinants contributing to plant species richness in mosaic landscapes: a single- and multi-patch perspective

Despite good theoretical knowledge about determinants of plant species richness in mosaic landscapes, validations based on complete surveys are scarce. We conducted a case study in a highly fragmented, traditional agricultural landscape. In 199 patches of 20 representative multi-patch-plots (MPPs, 1 ha) we recorded a total of 371 plant species. In addition to an additive partitioning of species diversity at the (a) patch- and (b) MPP-scale, we adopted the recently proposed ‘specificity’ measure to quantify the contribution of a spatial subunit to landscape species richness (subunit-to-landscape-contribution, SLC). SLC-values were calculated at both scales with respect to various spatial extents. General regression models were used to quantify the relative importance of hypothesis-driven determinants for species richness and SLC-values. At the patch scale, habitat type was the main determinant of species richness, followed by area and elongated shape. For SLC-values, area was more important than habitat type, and its relevance increased with the extent of the considered landscape. Influences of elongated shape and vegetation context were minor. Differences between habitat types were pronounced for species richness and also partly scale-dependent for SLC-values. Relevant predictors at the MPP-scale were nonlinear habitat richness, the gradient from anthropogenic to seminatural vegetation, and the proportions of natural vegetation and rare habitats. Linear elements and habitat configuration did not contribute to species richness and SLC. Results at the MPP-scale were in complete accordance with the predictions of the mosaic concept. Hence, our study represents its first empirical validation for plant species diversity in mosaic landscapes.     

The influence of landscape characteristics and home-range size on the quantification of landscape-genetics relationships

A common approach used to estimate landscape resistance involves comparing correlations of ecological and genetic distances calculated among individuals of a species. However, the location of sampled individuals may contain some degree of spatial uncertainty due to the natural variation of animals moving through their home range or measurement error in plant or animal locations. In this study, we evaluate the ways that spatial uncertainty, landscape characteristics, and genetic stochasticity interact to influence the strength and variability of conclusions about landscape-genetics relationships. We used a neutral landscape model to generate 45 landscapes composed of habitat and non-habitat, varying in percent habitat, aggregation, and structural connectivity (patch cohesion). We created true and alternate locations for 500 individuals, calculated ecological distances (least-cost paths), and simulated genetic distances among individuals. We compared correlations between ecological distances for true and alternate locations. We then simulated genotypes at 15 neutral loci and investigated whether the same influences could be detected in simple Mantel tests and while controlling for the effects of isolation-by-distance using the partial Mantel test. Spatial uncertainty interacted with the percentage of habitat in the landscape, but led to only small reductions in correlations. Furthermore, the strongest correlations occurred with low percent habitat, high aggregation, and low to intermediate levels of cohesion. Overall genetic stochasticity was relatively low and was influenced by landscape characteristics.     

Interactive effects of area and connectivity on the diversity of tachinid parasitoids in highly fragmented landscapes

Although many empirical and theoretical studies have elucidated the effects of habitat fragmentation on the third trophic level, little attention has been paid to the impacts of this driver on more generalist groups of non-hymenopteran parasitoids. Here, we used the highly-diverse group of tachinid flies as an alternative model to test the effects of landscape fragmentation on insect parasitoids. Our aims were: (i) to evaluate the relative importance of habitat area and connectivity losses and their potential interaction on tachinid diversity, (ii) to test whether the effects of habitat fragmentation changes seasonally, and (iii) to further assess the effect of habitat diversity on tachinid diversity and whether different parasitoid-host associations modify the species richness response to fragmentation. In 2012 a pan-trap sampling was conducted in 18 semi-natural grasslands embedded in intensive agricultural landscapes along statistically orthogonal gradients of habitat area, connectivity and habitat diversity. We found an interaction between habitat area and connectivity indicating that tachinid abundance and species richness were more negatively affected by habitat loss in landscapes with low rather than with relatively large habitat connectivity. Although tachinid communities exhibited large within-year species turnover, we found that the effects of landscape fragmentation did not change seasonally. We found that habitat diversity and host association did not affect tachinid species diversity. Our results have important implications for biodiversity conservation as any attempts to mitigate the negative effects of habitat loss need to take the general level of habitat connectivity in the landscape into account.     

Habitat specificity of patches in modern agricultural landscapes

Habitat specificity analysis provides a tool for partitioning landscape species diversity on landscape elements by separating patches with many rare specialist species from patches with the same number of species, all of which are common generalists and thus provide information of relevance to conservation goals at regional and national levels. Our analyses were based upon species data from 2201 patch elements in SE Norwegian modern agricultural landscapes. The context used for measuring habitat specificity strongly influences the results. In general the gamma diversity contribution and core habitat specificity calculated from the patch data set were correlated. High values for both measures were observed for woodland, pastures and road verges whereas midfield islets and boundary transitional types were ranked low, as opposed to findings in traditional, extensively managed agricultural landscapes. This is due to our study area representing intensively used agricultural landscape elements holding a more trivial species composition, in addition to ruderals being favoured by fertility and disturbance, a finding also being supported by the semi-natural affiliation index. Results obtained by use of checklist data from the same study area diverged from patch data. Caution is needed in interpretation of habitat specificity results obtained from checklist data, because modern agricultural landscapes contain several land types which are seldom surveyed by botanists, thus being under-represented in the data set. We propose the use of core habitat specificity and gamma diversity contribution in parallel to obtain a value neutral diversity assessment that addresses patch uniqueness and other properties of conservation interests.     

Effects of forest patch size on avian diversity

The effects of landscape patchiness on the diversity of birds of the Georgia Piedmont were investigated during 1993. Birds were sampled along line transects within relatively large (10–13.25 ha) and small (less than 3.25 ha) forest patches located within nonforest agricultural landscapes. Patterns of habitat use in these patches were compared to those in contiguous forest patches larger than 13.25 ha. Analysis of variance revealed significant differences in diversity between large and small woodlots and between contiguous and fragmented landscapes, especially in terms of the numbers of edge and interior species and winter-resident, summer-resident, and year-round birds observed.     

Biodiversity functions of urban cemeteries: Evidence from one of the largest Jewish cemeteries in Europe

As the world becomes more urbanized, urban cemeteries may become increasingly valuable for biodiversity conservation as cemeteries are ubiquitous elements of the green infrastructure in cities worldwide. By implementing a multi-taxon approach at different spatial extents, we analyzed habitat functions of a large urban cemetery in Berlin (Weiÿensee Jewish Cemetery) and explored related environmental variables. This cemetery is an outstanding cultural heritage site but it also stands for old urban cemeteries that have progressed to urban woodland, an ecosystem type that exists in many regional and religious contexts. The cemetery provided a habitat for 604 species; species of conservation concern comprised 1.6⿿100% of total species among different groups of taxa (in decreasing order: bats, birds, lichens, bryophytes, carabids, vascular plants, spiders). Species richness and species composition at the plot level were significantly related to differences in management intensity and resulting vegetation structures but differed among taxonomic groups. In vascular plants, carabids and spiders, the species composition varied significantly with habitat age, and there was a set of characteristic species for different age classes in each species group. Our results thus support the use of differentiated management approaches to maintain habitat heterogeneity by allowing wilderness development in some parts of a cemetery while keeping others more open. Since these aims can be combined with efforts to preserve outstanding grave architectures and allow access to visitors, our study indicates ways of reconciling conflicting aims of heritage preservation and biodiversity conservation, a promising perspective for biodiversity conservation in culturally shaped urban landscapes. We conclude that cemeteries provide important cultural ecosystem services within the urban green infrastructure.     

Butterfly assemblages in residential gardens are driven by species’ habitat preference and mobility

Context

Understanding the factors contributing to maintaining biodiversity is crucial to mitigate the impact of anthropogenic disturbances. Representing large proportions of green area in highly modified landscapes, residential gardens are often seen as local habitats that can contribute to larger networks of suitable environments at the landscape scale.

Objectives

We investigated the impact of the landscape context on butterfly communities observed in residential gardens, taking into account garden characteristics, land-use types and presence of linear features in the surrounding landscape. We examined how species traits affected butterflies’ response to landscape context and habitat quality.

Methods

We performed a cross-scale study, based on citizen science data documenting butterfly species composition and abundance in 920 gardens across France. We examined the effect of garden quality, the area of different land-use types and the length of linear elements measured at three scales within the surrounding landscape. Species were grouped according to their habitat preference and mobility.

Results

Urbanization negatively affected total species richness and the abundance of butterfly in each group. This was related to declining habitat quality and reduced area of suitable habitat in the surrounding landscape. The magnitude of this effect, however, was negatively correlated with mobility, a trait related to habitat preference. The spatial scale at which landscape context best explained variation in butterfly abundance changed with species’ habitat preference.

Conclusions

This study highlights the importance of preserving high quality habitats in altered landscapes and considering species’ mobility and habitat preference when assessing the impact of landscapes on butterfly communities.

     

Urban-rural and temporal differences of woody plants and bird species in Harbin city,northeastern China

Urbanization has been greatly accelerated by the economic growth in China, while its possible effects on woody plants, bird species and their associations are not well defined yet. Here, we analyzed urban-rural gradients (landscape level: urban-farmland-forest-natural reserves; city level: ring road and urban build-up history) and temporal data (1955–1980–2014 for woody plants; 1980s–2010 s for birds) in Harbin city, China, to investigate the changes in the composition and diversity of woody plants and birds during urbanization. Both landscape gradient and temporal data confirmed that urbanization had the function of species conservations with sharp increases of alien species and tropical type plants. In the case woody species, 60-yr urbanization in Harbin had induced increases of 9 families and 17 genera, and there were 7–20 more families, 12–35 more genera, 1.6–2.6 higher Margalef richness in urban areas than those in nature reserves and local forest farms; Increases in alien species (4-fold in 60-yr urbanization; 21% in urban area vs <2% for non-urban region) and tropical type plants (1.6-fold in 60-yr urbanization; temperate/tropical ratio at 1.2 in urban area vs >1.6 in non-urban area) were mainly responsible for these compositional changes, which can be proved by their significant correlations. Moreover, moderate disturbance had peak values in alien species, tropical type plants, Shannon-wiener diversity, Margalef richness index and Pielou evenness index, and both ring road- and buildup history gradients showed the similar tendency. Compared with those in 1980s, forest- and eurytopic-habitats birds increased 9–11 species (23–39%), and omnivorous, insect-eating, and phytophagous bird increased 5–9 species (14.1–29.4%) in those in 2010s, indicating that bird temporal changes were closely related with the changes in urban forests owing to food supply and habitat provision. Our findings could provide data for biodiversity evaluation of urbanization effects, and is also useful for ecological re-construction of local cities in China.     

Species Dynamics in Disturbed Landscapes: When does a Shifting Habitat Mosaic Enhance Connectivity?

Although landscape ecology emphasizes the effects of spatial pattern on ecological processes, most neutral models of species–habitat relationships have treated habitat as a static constraint. Do the working hypotheses derived from these models extend to real landscapes where disturbances create a shifting mosaic? A spatial landscape simulator incorporating vegetation dynamics and a metapopulation model was used to compare species in static and dynamic landscapes with identical habitat amounts and spatial patterns. The main drivers of vegetation dynamics were stand-replacing disturbances, followed by gradual change from early-successional to old-growth habitats. Species dynamics were based on a simple occupancy model, with dispersal simulated as a random walk. As the proportion of available habitat (p) decreased from 1.0, species occupancy generally declined more rapidly and reached extinction at higher habitat levels in dynamic than in static landscapes. However, habitat occupancy was sometimes actually higher in dynamic landscapes than in static landscapes with similar habitat amounts and patterns. This effect was most pronounced at intermediate amounts of habitat (p = 0.3?0.6) for mobile species that had high colonization rates, but were unable to cross non-habitat patches. Differences between static and dynamic landscapes were contingent upon the initial metapopulation size and the shapes of disturbances and the resulting habitat patterns. Overall, the results demonstrate that dispersal-limited species exhibit more pronounced critical behavior in dynamic landscapes than is predicted by simple neutral models based on static landscapes. Thus, caution should be exercised in extending generalizations derived from static landscape models to disturbance-driven landscape mosaics.     

Impacts of selective logging on tree diversity across a rainforest landscape: the importance of spatial scale

Selective logging of tropical forests imposes spatial pattern on the landscape by creating a mosaic of patches affected by different intensities of disturbance. To understand the ecological impacts of selective logging it is therefore necessary to explore how patterns of tree species composition are affected by this patchy disturbance. This study examines the impacts of selective logging on species composition and spatial patterns of vegetation structure and tree diversity in Sabah, Borneo. We compare tree diversity between logged and unlogged forest at three scales: species richness within plots, species turnover among plots, and total species richness and composition of plots combined. Logging had no effect on tree diversity measured at the smallest scale. Logged forest had a greater rate of species turnover with distance, so at a large spatial scale it supported more tree species than the relatively homogeneous unlogged area. Tree species composition also differed significantly between the two types of forest, with more small dipterocarps and large pioneers in logged forest, and more large dipterocarps in unlogged forest. Our results emphasize the importance of sampling at a sufficiently large scale to represent patterns of biodiversity within tropical forest landscapes. Large areas of production forest in SE Asia are threatened with conversion to commercial crops; our findings show that selectively logged forest can retain considerable conservation value.     

Spiders associated with the meadow and tree canopies of orchards respond differently to habitat fragmentation

The response of animal communities to habitat quality and fragmentation may vary depending on microhabitat associations of species. For example, sensitivity of species to woody habitat fragmentation should increase with their degree of association with woody plants. We investigated effects of local and landscape factors on spider communities in different microhabitats within Swiss apple orchards. We expected a stronger negative effect of woody habitat fragmentation on spiders inhabiting tree canopies compared to spiders living in the meadow. The 30 orchards that we sampled varied in woody habitat amount and isolation at landscape and patch scales. Local factors included management intensity and plant diversity. Spiders associated with meadow were affected by plant diversity, but not by fragmentation. In contrast, spiders associated with canopies responded to isolation from other woody habitats. Surprisingly, we found both positive and negative effects of habitat isolation on local abundance. This indicates that differences in dispersal and/or biotic interactions shape the specific response to habitat isolation. The relative importance of local and landscape factors was in accordance with the microhabitat of the spiders. Thus, considering microhabitat associations can be important for identifying processes that would be overlooked if sampling were pooled for the whole habitat.     

Responses of Chilean forest birds to anthropogenic habitat fragmentation across spatial scales

Although it is recognized that anthropogenic forest fragmentation affects habitat use by organisms across multiple spatial scales, there is uncertainty about these effects. We used a hierarchical sampling design spanning three spatial scales of habitat variability (landscape > patch > within-patch) and generalized mixed-effect models to assess the scale-dependent responses of bird species to fragmentation in temperate forests of southern Chile. The abundances of nine of 20 bird species were affected by interactions across spatial scales. These interactions resulted in a limited effect of within-patch habitat structure on the abundance of birds in landscapes with low forest cover, suggesting that suitable local habitats, such as sites with dense understory cover or large trees, are underutilized or remain unused in highly fragmented landscapes. Habitat specialists and cavity-nesters, such as tree-trunk foragers and tapaculos, were most likely to exhibit interactions across spatial scales. Because providing additional sites with dense understory vegetation or large habitat trees does not compensate the negative effect of the loss of forest area on bird species, conservation strategies should ensure the retention of native forest patches in the mixed-use landscapes.     

Functional connectivity of lynx at their southern range periphery in Ontario,Canada

Peripheral populations are often small and isolated compared to those in the range core, in part due to the patchy distribution of suitable habitats at range margins. It follows that peripheral populations typically occur at lower densities and are more susceptible to extinction, but their persistence may be facilitated through connectivity with core areas. Relationships between connectivity and the distribution of animal populations have not yet been fully evaluated, especially for large carnivores having extensive spatial needs and specialized habitat requirements. Using observations of snow tracks, we modeled occurrence of Canada lynx (Lynx canadensis) in relation to landscape characteristics along their southern range periphery in Ontario, Canada; we sought to assess functional connectivity of lynx habitat along the southern margins of the range. As observed in other studies, young coniferous forests had the highest probability of lynx occurrence, likely due to their association with snowshoe hares (Lepus americanus). We used the occurrence model to parameterize a resistance surface and then circuit theory to predict functional connectivity along the southern periphery of lynx distribution. Lynx typically travelled through landscapes with higher connectivity than random paths, implying that lynx habitat requirements in their southern range likely extend beyond habitat composition, and that conservation efforts should seek to preserve metapopulation dynamics through functional connectivity of suitable habitat across larger spatial scales.