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.

Context

The classical theory of island biogeography explains loss of species in fragmented landscapes as an effect of remnant patch size and isolation. Recently this has been challenged by the habitat amount and habitat continuum hypotheses, according to which persistence in modified landscapes is related to total habitat amount rather than habitat configuration or the ability of species to use all habitats to varying degrees. Distinguishing between these theories is essential for effective conservation planning in modified landscapes.

Objective

Identify which factors of habitat type, amount and configuration predict the persistence of a keystone woodland specialist, the eastern bettong Bettongia gaimardi, in a fragmented landscape.

Method

In the Midlands region of Tasmania we carried out camera surveys at 62 sites in summer and winter. We included habitat and landscape features to model whether habitat amount or patch size and isolation influenced the presence of the eastern bettong, and to measure effects of habitat quality.

Results

Habitat amount within a 1 km buffer was a better predictor of occupancy than patch size and isolation. Occupancy was also affected by habitat quality, indicated by density of regenerating stems.

Conclusion

Our results support the habitat amount hypothesis as a better predictor of presence. For a species that is able to cross the matrix between remnant patches and utilise multiple patches, the island biogeography concept does not explain habitat use in fragmented landscapes. Our results emphasize the value of small remnant patches for conservation of the eastern bettong, provided those patches are in good condition.

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.

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.

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.

Context

The relative influence of habitat loss versus configuration on avian biodiversity is poorly understood. However, this knowledge is essential for developing effective land use strategies, especially for grassland songbirds, which have experienced widespread declines due to land use changes. Habitat configuration may be particularly important to grassland songbirds as configuration of habitat affects the extent of edge effects on the landscape, which strongly influences habitat use by grassland birds.

Objectives

We examined the relative influence of grassland amount and a measure of grassland configuration per se (Landscape Shape Index; LSI) on the relative abundance and richness of grassland songbirds.

Methods

In 2013, 361 avian point counts were conducted across 47, 2.4 km radii landscapes in south-west Manitoba, Canada, selected to minimize the correlation between grassland amount and configuration. We used generalized linear mixed-effects models within a multi-model inference framework to determine the relative importance of grassland amount and configuration on songbird response variables.

Results

Effects of grassland amount and configuration were generally weak, but effects of configuration were greater than grassland amount for most species. Relative abundance and richness of obligate species, and Savannah sparrows, showed a strong negative response to LSI, while grasshopper sparrows responded positively to grassland amount.

Conclusion

Our results suggest that habitat configuration must be considered when managing landscapes for conservation of grassland songbirds. Maintaining large, intact tracts of grasslands and limiting development of roads that bisect grassland parcels may be an effective means of maintaining grassland songbird diversity and abundance in northern mixed-grass prairies.

Context

Urban environments create a wide range of habitats that harbour a great diversity of plant species, many of which are of alien origin. For future urban planning and management of the green areas within the city, understanding of the spatial distribution of invasive alien species is of great importance.

Objectives

Our main aim was to assess how availability of different ecosystem types within a city area, as well as several parameters describing urban structure interact in determining the cover and identity of invasive alien species.

Methods

We studied the distribution of chosen invasive plant species in a mid-sized city in the Czech Republic, central Europe, on a gradient of equal sized cells from the city centre to its outskirts.

Results

A great amount of variation was explained by spatial predictors but not shared with any measured variables. The species cover of invasive species decreased with increasing proportion of urban greenery and distance from the city centre, but increased with habitat richness; road margins, ruderal sites, and railway sites were richest in invasive species. In contrast, the total number of invasive species in cells significantly decreased with increasing distance from the city centre, but increased with habitat richness.

Conclusions

Our results suggest that different invasive species prefer habitats in the vicinity of the city centre and at its periphery and the spatial structure and habitat quality of the urban landscape needs to be taken into account, in efforts to manage alien plant species invasions in urban environments.

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.

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.

Context

Hoverflies are often used as bio-indicators for ecosystem conservation, but only few studies have actually investigated the key factors explaining their richness in woodlands.

Objectives

In a fragmented landscape in southwest France, we investigated the joint effects of woodland area, structural heterogeneity, connectivity and history on the species richness of forest-specialist hoverflies, and whether there was a time lag in the response of hoverflies to habitat changes, and tested the effect of spatiotemporal changes.

Methods

Current species richness was sampled in 48 woodlands using 99 Malaise traps. Structural variables were derived from a rapid habitat assessment protocol. Old maps and aerial photographs were used to extract past and present spatial patterns of the woodlands since 1850. Relationships between species richness and explanatory variables were explored using generalized linear models.

Results

We show that current habitat area, connectivity, historical continuity and the average density of tree-microhabitats explained 35 % of variation in species richness. Species richness was affected differently by changes in patch area between 1979 and 2010, depending on woodland connectivity. In isolated woodlands, extinction debt and colonization credit were revealed, showing that even several decades are not sufficient for hoverflies to adapt to landscape-scale habitat conditions.

Conclusions

These findings emphasise the importance of maintaining connectedness between woodlands, which facilitates the dispersion in a changing landscape. Our results also highlight the benefits of using a change-oriented approach to explain the current distribution patterns of species, especially when several spatial processes act jointly.

Context

Beyond the recognized importance of protecting large areas of contiguous habitat, conservation efforts for many species are complicated by the fact that patch suitability may also be affected by characteristics of the landscape within which the patch is located. Currently, little is known about the spatial scales at which species respond to different aspects of the landscape surrounding an occupied patch.

Objectives

Using grassland bird point count data, we describe an approach to evaluating scale-specific effects of landscape composition on patch occupancy.

Methods

We used data from 793 point count surveys conducted in idle and grazed grasslands across Wisconsin, USA from 2012 to 2014 to evaluate scale-dependencies in the response of grassland birds to landscape composition. Patch occupancy models were used to evaluate the relationship between occupancy and landscape composition at scales from 100 to 3000 m.

Results

Bobolink (Dolichonyx oryzivorus) exhibited a pattern indicating selection for grassland habitats in the surrounding landscape at all spatial scales while selecting against other habitats. Eastern Meadowlark (Sturnella magna) displayed evidence of scale sensitivity for all habitat types. Grasshopper Sparrow (Ammodramus savannarum) showed a strong positive response to pasture and idle grass at all scales and negatively to cropland at large scales. Unlike other species, patch occupancy by Henslow’s Sparrow (A. henslowii) was primarily influenced by patch area.

Conclusions

Our results suggest that both working grasslands (pasture) and idle conservation grasslands can play an important role in grassland bird conservation but also highlight the importance of considering species-specific patch and landscape characteristics for effective conservation.

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.

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.

Context

Loss and fragmentation of semi-natural grasslands has critically affected many butterfly species in Europe. Habitat area and isolation can have strong effects on the local biodiversity but species may also be strongly affected by the surrounding matrix.

Objectives

We explored how different land cover types in the landscape explained the occurrence of butterfly species in semi-natural grasslands.

Methods

Using data from 476 semi-natural grasslands in Sweden, we analysed the effect of matrix composition on species richness and occurrence. Additionally, we analysed at which spatial scales butterflies responded to matrix types (forests, semi-natural grasslands, arable land and water).

Results

Forest cover showed the strongest positive effect on species richness, followed by semi-natural grasslands. Forest also had a positive effect on red-listed species at local scales. Responses to matrix composition were highly species-specific. The majority of the 30 most common species showed strong positive responses to the amount of forest cover within 200–500 m. There was a smaller group of species showing a positive response to arable land cover within 500–2000 m. Thirteen species showed positive responses to the amount of semi-natural grasslands, generally at larger scales (10–30 km).

Conclusions

Our study showed that surrounding forest is beneficial for many grassland butterfly species and that forests might mitigate the negative effects of habitat loss caused by agricultural intensification. Also, semi-natural grasslands were an important factor for species richness at larger spatial scales, indicating that a landscape consisting mainly of supporting habitats (i.e. forests) are insufficient to sustain a rich butterfly fauna.

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.

Context

Native vegetation is often used as a proxy for habitat to estimate habitat availability in landscapes. This approach may lead to incorrect estimates of the impacts of habitat loss and fragmentation on species, which have not been thoroughly quantified so far.

Objectives

We quantified to what extent the loss of native vegetation reflect actual habitat loss by native species in landscapes. We tested the hypothesis that habitat availability declines at greater rates than native vegetation and thus is overestimated when it is quantified on the basis of native vegetation.

Methods

Using simulations, we quantified how the loss of native vegetation in artificial and real landscapes affects habitat availability for species with different habitat requirements. We contrasted a generalist species, which uses all native vegetation, with 10 habitat-specialist species classified into three categories (interior, patchy and riparian species).

Results

Habitat availability generally declined at greater rates than native vegetation for all specialist species. This pattern was apparent for different specialist species in a broad range of landscape types. Interior species always lost habitat availability more rapidly than the generalist species. Most riparian species lost habitat availability more rapidly than the generalist species. Responses of patchy species were more complex, depending on their dispersal abilities and landscape structure.

Conclusions

Habitat availability is likely to be overestimated when native vegetation is used as proxy for habitat, because habitat availability will generally decline at greater rates than native vegetation. Therefore, a species-centered approach should be adopted when estimating habitat availability in landscapes.

Context

Habitat destruction is the leading threat to terrestrial biodiversity, isolating remnant habitat in a matrix of modified vegetation.

Objectives

Our goal was to determine how species richness in several broad taxonomic groups from remnant forest was influenced by matrix quality, which we characterized by comparing plant biomass in forest and the surrounding matrix.

Methods

We coupled data on species-area relationships (SARs) in forest remnants from 45 previously published studies with an index of matrix quality calculated using new estimates of plant biomass derived from satellite imagery.

Results

The effect size of SARs was greatest in landscapes with low matrix quality and little forest cover. SARs were generally stronger for volant than for non-volant species. For the terrestrial taxa included in our analysis, matrix quality decreased as the proportion of water, ice, or urbanization in a landscape increased.

Conclusions

We clearly demonstrate that matrix quality plays a major role in determining patterns of species richness in remnant forest. A key implication of our work is that activities that increase matrix quality, such as active and passive habitat restoration, may be important conservation measure for maintaining and restoring biodiversity in modified landscapes.

Context

Despite decades of research, there is an intense debate about the consistency of the hump-shaped pattern describing the relationship between diversity and disturbance as predicted by the intermediate disturbance hypothesis (IDH). Previous meta-analyses have not explicitly considered interactive effects of disturbance frequency and intensity of disturbance on plant species diversity in terrestrial landscapes.

Objective

We conducted meta-analyses to test the applicability of IDH by simultaneously examining the relationship between species richness, disturbance frequency (quantified as time since last disturbance as originally proposed) and intensity of disturbance in forest landscapes.

Methods

The effects of disturbance frequency, intensity, and their interaction on species richness was evaluated using a mixed-effects model.

Results

We found that species richness peaks at intermediate frequency after both high and intermediate disturbance intensities, but the richness-frequency relationship differed between intensity classes.

Conclusions

Our study highlights the need to measure multiple disturbance components that could help reconcile conflicting empirical results on the effect of disturbance on plant species diversity.

Context

In modern agricultural landscapes, fragmentation of partial habitats is a significant filter for multi-habitat users, reducing local taxonomic and functional diversity. There is compelling evidence that small species are more susceptible than large species. The impact of habitat fragmentation on intraspecific body-size distribution, however, is yet unexplored.

Objectives

We tested habitat fragmentation, a major driver of pollinator loss, for its impact on intraspecific body-size distributions of solitary wild-bee species. Subsequently, we tested individual body size for its impact on pollination services.

Methods

We sampled 1272 individuals of the four most common Andrena wild bee species in 22 newly established flowering fields (0.21–0.41 ha) in Hessen, Central Germany, over two consecutive years. Study sites were located in a ca. 80 ha landscape context of increasing habitat fragmentation. We analysed the pollen loads of the most abundant species.

Results

Body size within local populations of the two medium-sized bees increased with fragmentation, suggesting intraspecific selection for higher dispersal capacity. Pollen analysis carried out for the most common species revealed that larger individuals visited a significantly smaller plant spectrum. Habitat fragmentation may thus alter pollination services without necessarily affecting species richness or composition.

Conclusions

Systematic body-size variation at the population level thus explains the considerable variability between simple community measures and ecosystem functioning. Filtering processes at the individual level require increased understanding for targeting pollination services under current and future land-use change.

Context

Ecological theory suggests that large habitat fragments should harbour more species than small fragments. However, this may depend on the surrounding matrix. Matrices in fragmented landscapes may either amplify or reduce area effects, which could influence predicted extinctions based on species-area relationships (SARs).

Objective

To determine the influence of matrix type on SARs.

Methods

We surveyed birds within 59 coastal forest fragments in two matrix types, anthropogenic (South Africa) and natural (Mozambique). We classified species as forest specialists or habitat generalists and fitted species-area models to compare how SAR slopes differed among matrix types. We also calculated nestedness and evenness to determine if these varied among matrix type and used logistic regressions to identify species-specific responses to matrix type.

Results

For habitat generalists, SARs were weak within both matrices, while for forest specialists it was strong in the anthropogenic but weak in the natural matrix. In the former, the SAR was similar to those recorded for real islands within archipelagos. Forest specialist assemblages were nested by area within anthropogenic, but not natural matrices. Matrix type did not influence evenness. Area only affected the occurrence of one species when the matrix was natural, compared to 11 species when it was anthropogenic.

Conclusions

Forest specialist bird species conformed to island biogeographic predictions of species loss in forest fragments embedded in anthropogenic, but not natural matrices. Extinctions from small forest fragments might be prevented by conserving natural- or restoring anthropogenic matrices, as well as by increasing forest area.