Context

Habitat loss, fragmentation and degradation are widespread drivers of biodiversity decline. Understanding how habitat quality interacts with landscape context, and how they jointly affect species in human-modified landscapes, is of great importance for informing conservation and management.

Objectives

We used a whole-ecosystem manipulation experiment in the Brazilian Amazon to investigate the relative roles of local and landscape attributes in affecting bat assemblages at an interior-edge-matrix disturbance gradient.

Methods

We surveyed bats in 39 sites, comprising continuous forest (CF), fragments, forest edges and intervening secondary regrowth. For each site, we assessed vegetation structure (local-scale variable) and, for five focal scales, quantified habitat amount and four landscape configuration metrics.

Results

Smaller fragments, edges and regrowth sites had fewer species and higher levels of dominance than CF. Regardless of the landscape scale analysed, species richness and evenness were mostly related to the amount of forest cover. Vegetation structure and configurational metrics were important predictors of abundance, whereby the magnitude and direction of response to configurational metrics were scale-dependent. Responses were ensemble-specific with local-scale vegetation structure being more important for frugivorous than for gleaning animalivorous bats.

Conclusions

Our study indicates that scale-sensitive measures of landscape structure are needed for a more comprehensive understanding of the effects of fragmentation on tropical biota. Although forest fragments and regrowth habitats can be of conservation significance for tropical bats our results further emphasize that primary forest is of irreplaceable value, underlining that their conservation can only be achieved by the preservation of large expanses of pristine habitat.

Context

The analysis of individual movement choices can be used to better understand population-level resource selection and inform management.

Objectives

We investigated movements and habitat selection of 13 bobcats in Vermont, USA, under the assumption individuals makes choices based upon their current location. Results were used to identify “movement-defined” corridors.

Methods

We used GPS-collars and GIS to estimate bobcat movement paths, and extracted statistics on land cover proportions, topography, fine-scale vegetation, roads, and streams within “used” and “available” space surrounding each movement path. Compositional analyses were used to determine habitat preferences with respect to landcover and topography; ratio tests were used to determine if used versus available ratios for vegetation, roads, and streams differed from 1. Results were used to create travel cost maps, a primary input for corridor analysis.

Results

Forested and scrub-rock land cover were most preferred for movement, while developed land cover was least preferred. Preference depended on the composition of the “available” landscape: Bobcats moved?>?3 times more quickly through forest and scrub-rock habitat when these habitats were surrounded by agriculture or development than when the available buffer was similarly composed. Overall, forest edge, wetland edge and higher stream densities were selected, while deep forest core and high road densities were not selected. Landscape-scale connectivity maps differed depending on whether habitat suitability, preference, or selection informed the travel cost map.

Conclusions

Both local and landscape scale land cover characteristics affect habitat preferences and travel speed of bobcats, which in turn can inform management and conservation activities.

Context

Understanding habitat selection can be challenging for species surviving in small populations, but is needed for landscape-scale conservation planning.

Objectives

We assessed how European bison (Bison bonasus) habitat selection, and particularly forest use, varies across subpopulations and spatial scales.

Methods

We gathered the most comprehensive European bison occurrence dataset to date, from five free-ranging herds in Poland. We compared these data to a high-resolution forest map and modelled the influence of environmental and human-pressure variables on habitat selection.

Results

Around 65% of European bison occurrences were in forests, with cows showing a slightly higher forest association than bulls. Forest association did not change markedly across spatial scales, yet differed strongly among herds. Modelling European bison habitat suitability confirmed forest preference, but also showed strong differences in habitat selection among herds. Some herds used open areas heavily and actively selected for them. Similarly, human-pressure variables were important in all herds, but some herds avoided human-dominated areas more than others.

Conclusions

Assessing European bison habitat across multiple herds revealed a more generalist habitat use pattern than when studying individual herds only. Our results highlight that conflicts with land use and people could be substantial if bison are released in human-dominated landscapes. Future restoration efforts should target areas with low road and human population density, regardless of the degree of forest cover. More broadly, our study highlights the importance of considering multiple subpopulations and spatial scales in conservation planning.

Context

The application of regional-level airborne lidar (light detection and ranging) data to characterize habitat patches and model habitat connectivity over large landscapes has not been well explored. Maintaining a connected network of habitat in the presence of anthropogenic disturbances is essential for regional-level conservation planning and the maintenance of biodiversity values.

Objectives

We quantified variation in connectivity following simulated changes in land cover and contrasted outcomes when different conservation priorities were emphasized.

Methods

First, we defined habitat patches using vegetation structural attributes identified via lidar. Second, habitat networks were constructed for different forest types and assessed using network connectivity metrics. And finally, land cover change scenarios were simulated using a series of habitat patch removals, representing the impact of implementing different spatial prioritization schemes.

Results

Networks for different forest structure types produced very different patch distributions. Conservation scenarios based on different schemes led to contrasting changes during land cover change simulations: the scheme prioritizing only habitat area resulted in immediate near-term losses in connectivity, whereas the scheme considering both habitat area and their spatial configurations maintained the overall connectivity most effectively. Adding climate constraints did not diminish or improve overall connectivity.

Conclusions

Both habitat area and habitat configuration should be considered in dynamic modeling of habitat connectivity under changing landscapes. This research provides a framework for integrating forest structure and cover attributes obtained from remote sensing data into network connectivity modeling, and may serve as a prototype for multi-criteria forest management and conservation planning.

Context

Habitat loss and fragmentation may alter habitat occupancy patterns, for example through a reduction in regional abundance or in functional connectivity, which in turn may reduce the number of dispersers or their ability to prospect for territories. Yet, the relationship between landscape structure and habitat niche remains poorly known.

Objectives

We hypothesized that changes in landscape structure associated with habitat loss and fragmentation will reduce the habitat niche breadth of forest birds, either through a reduction in density-dependent spillover from optimal habitat or by impeding the colonization of patches.

Methods

We surveyed forest birds with point counts in eastern Ontario, Canada, and analyzed their response to loss and fragmentation of mature woodland. We selected 62 landscapes varying in both forest cover (15–45%) and its degree of fragmentation, and classified them into two categories (high versus low levels of loss and fragmentation). We determined the habitat niche breadth of 12 focal species as a function of 8 habitat structure variables for each landscape category.

Results

Habitat niche breadth was narrower in landscapes with high versus low levels of loss and fragmentation of forest cover. The relative occupancy of marginal habitat appeared to drive this relationship. Species sensitivity to mature forest cover had no apparent influence on relative niche breadth.

Conclusions

Regional abundance and, in turn, density-dependent spillover into suboptimal habitat appeared to be determinants of habitat niche breadth. For a given proportion of forest cover, fragmentation also appeared to alter habitat use, which could exacerbate its other negative effects unless functional connectivity is high enough to allow individuals to saturate optimal habitat.

Context

Freshwater ecosystems depend on surrounding terrestrial landscape for resources. Most important are terrestrial leaf litter subsidies, which differ depending on land use. We lack a good understanding of the variation of these inputs across spatial scales.

Objectives

We sought to determine: (1) the relative importance of local versus catchment-level forestation for benthic leaf litter biomass in streams, (2) how landscape configuration alters these relationships, and (3) how land use affects the quality and diversity of leaf litter subsidies.

Methods

We measured biomass and identity of benthic leaf litter in 121 reaches in 10 independent catchments seasonally over the course of a year. We assessed direct and indirect effects of forestation, reach position, and seasonality on leaf litter biomass using structural equation models, and assessed how leaf litter diversity varied with land use.

Results

In catchments with forested headwaters, the degree of forestation and reach position in the catchment influenced benthic leaf litter biomass indirectly through local reach-scale forestation. In catchments where forest was only located downstream, or with minimal forest, none of these factors influenced reach-level benthic leaf litter. Leaf litter diversity peaked in fall in all land use types, but was generally lowest in forested reaches.

Conclusions

Not only habitat amount, but its location relative to other habitats is important for ecosystem function in the context of cross-ecosystem material flows. Here, lack of upstream forest altered spatial patterns of leaf litter storage. Studies with high spatiotemporal resolution may further reveal effects of landscape configuration on other ecosystems.

Context

Primates are an important component of biodiversity in tropical regions. However, many studies on the effects of habitat change on primates ignore the relative influence of landscape composition and configuration.

Objectives

This study addresses the question: how important are landscape-scale forest area and composition relative to patch-scale (1–1080 ha) and site-scale (transect of 1 km) habitat variables for the occupancy and abundance of four primate species in the Colombian Llanos.

Methods

Using a randomly stratified survey design, 81 fragments were surveyed for primate occupancy and abundance. We used zero-inflated models to test the relative influence of landscape-scale, patch-scale and site-scale variables on occupancy and abundance for each species. A 95% confidence set of models was constructed using the cumulative Akaike weight for each model and the relative importance of each set of variables calculated for each primate species.

Results

Occupancy was determined by a combination of site-scale, patch-scale and landscape-scale variables but this varied substantially among the primate species.

Conclusion

Our study highlights the importance of managing primates at a range of scales that considers the relative importance of site-, patch- and landscape-scale variables.

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

The positive correlation between landscape area of semi-natural habitat and wild pollinator richness and abundance in agroecosystems has been well studied. However, we lack a deep understanding of local scale floral resource and nest provisioning for wild bees necessary to optimize implementation of pollinator conservation practices.

Objectives

The primary objective of this study was to use a spatially interactive landscape pollination model (hereafter, the Lonsdorf model) to represent field scale spatial patterns of wild bee abundance and richness within a heterogeneous landscape in the mid-Atlantic USA.

Methods

We parameterized the Lonsdorf model with high resolution aerial imagery and insight from a previously published floristic study. To test the Lonsdorf model predictions, field studies were conducted to measure wild bee abundance and species richness in apple orchards as a function of distance from a forest edge.

Results

Field measurements indicated apple pollinator abundance and species richness significantly decreased with increasing distance from the forest edge. The Lonsdorf model pollination service score was highly sensitive to changes in resource provisioning in orchard and non-crop areas, and including resource rich forest and forest edge habitats in the model significantly improved pollination service estimates.

Conclusions

We demonstrated a novel application of the Lonsdorf model at a field scale to predict trends in pollination service provisioning as a factor of local habitat features. With sufficiently detailed inputs, the Lonsdorf model is a promising tool to quantify field scale pollination service deficits, guiding more cost effective habitat supplementation and other conservation efforts.

Context

Species site-occupancy patterns may be influenced by habitat variables at both local and landscape scales. Although local habitat variables influence whether the site is suitable for a given species, the broader landscape context can also influence site occupancy, particularly for species that are sensitive to land-use change.

Objectives

To examine the relative importance of local versus landscape variables in explaining site occupancy of eight bat species within the Brazilian Cerrado, a Neotropical savanna that is experiencing widespread habitat loss and fragmentation.

Methods

Bats were surveyed within 16 forest patches over two years. We used a multi-model information-theoretic approach, adjusted for species detection bias, to assess whether landscape variables (percent cover and number of patches of natural vegetation within a 2- and 8-km radius of each forest site) or local site variables (canopy cover, understory height, number of trees, and number of lianas) best explained site occupancy in each species.

Results

Landscape variables were among the best models (ΔAIC_c or ΔQAIC_c < 2) for four species (top-ranked model for black myotis), whereas local variables were among the best for five species (top-ranked model for vampire bats). Neither local nor landscape variables explained site occupancy in two frugivorous species.

Conclusion

Species associated with a particular habitat type will not respond similarly to the amount, distribution or relative suitability of that habitat, or even at the same scale. This reinforces the challenge of species distribution modelling, especially in the context of forecasting species’ responses to future land-use or climate-change scenarios.

Context

Landscape-scale population dynamics are driven in part by movement within and dispersal among habitat patches. Predicting these processes requires information about how movement behavior varies among land cover types.

Objectives

We investigated how butterfly movement in a heterogeneous landscape varies within and between habitat and matrix land cover types, and the implications of these differences for within-patch residence times and among-patch connectivity.

Methods

We empirically measured movement behavior in the Baltimore checkerspot butterfly (Euphydryas phaeton) in three land cover classes that broadly constitute habitat and two classes that constitute matrix. We also measured habitat preference at boundaries. We predicted patch residence times and interpatch dispersal using movement parameters estimated separately for each habitat and matrix land cover subclass (5 categories), or for combined habitat and combined matrix land cover classes (2 categories). We evaluated the effects of including edge behavior on all metrics.

Results

Overall, movement was slower within habitat land cover types, and faster in matrix cover types. Butterflies at forest edges were biased to remain in open areas, and connectivity and patch residence times were most affected by behavior at structural edges. Differences in movement between matrix subclasses had a greater effect on predictions about connectivity than differences between habitat subclasses. Differences in movement among habitat subclasses had a greater effect on residence times.

Conclusions

Our findings highlight the importance of careful classification of movement and land cover in heterogeneous landscapes, and reveal how subtle differences in behavioral responses to land cover can affect landscape-scale outcomes.

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.

Context

Golden-cheeked warblers (Setophaga chrysoparia), an endangered wood-warbler, breed exclusively in woodlands co-dominated by Ashe juniper (Juniperus ashei) in central Texas. Their breeding range is becoming increasingly urbanized and habitat loss and fragmentation are a main threat to the species’ viability.

Objectives

We investigated the effects of remotely sensed local habitat and landscape attributes on point occupancy and density of warblers in an urban preserve and produced a spatially explicit density map for the preserve using model-supported relationships.

Methods

We conducted 1507 point-count surveys during spring 2011–2014 across Balcones Canyonlands Preserve (BCP) to evaluate warbler habitat associations and predict density of males. We used hierarchical Bayesian models to estimate multiple components of detection probability and evaluate covariate effects on detection probability, point occupancy, and density.

Results

Point occupancy was positively related to landscape forest cover and local canopy cover; mean occupancy was 0.83. Density was influenced more by local than landscape factors. Density increased with greater amounts of juniper and mixed forest and decreased with more open edge. There was a weak negative relationship between density and landscape urban land cover.

Conclusions

Landscape composition and habitat structure were important determinants of warbler occupancy and density, and the large intact patches of juniper and mixed forest on BCP (>2100 ha) supported a high density of warblers. Increasing urbanization and fragmentation in the surrounding landscape will likely result in lower breeding density due to loss of juniper and mixed forest and increasing urban land cover and edge.

Context

Conservation research often focuses on individual threats at a single spatial scale, but population declines can result from multiple stressors occurring at different spatial scales. Analyses incorporating alternative hypotheses across spatial scales allow more robust evaluation of the ecological processes underlying population declines.

Objectives

Populations of many aerially insectivorous birds are declining, yet conservation efforts remain focused on habitat due to an absence of data on changes in prey availability. We evaluate the potential for prey and habitat availability at multiple spatial scales to influence a population of eastern whip-poor-wills (Antrostomus vociferous).

Methods

We assess relationships between landcover (topographical map and satellite imagery) and insect abundance (moths and beetles from blacklight traps), and whip-poor-will distribution and abundance within eastern Canada using Ontario breeding bird atlas data (1980s and 2000s), acoustic recordings (regional), and point counts (local).

Results

Whip-poor-will occurrence in both atlas time periods was positively associated with forest area and fragmentation, but only a delayed effect of urban area explained reductions in detection. Contemporary regional whip-poor-will presence was positively related to moth abundance, and local whip-poor-will abundance was best predicted by area of open-canopy forest, anthropogenic linear disturbance density, and beetle abundance. Our finding that bird presence and abundance were associated with human activity and insect abundance across spatial scales suggests factors beyond habitat structure are likely driving population declines in whip-poor-wills and other aerial insectivores.

Conclusions

This study demonstrates the importance of examining multiple hypotheses, including seasonally and locally variable food availability, across a range of spatial scales to direct conservation efforts.

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

The anthropocene is characterised by global landscape modification, and the structure of remnant habitats can explain different patterns of species richness. The most pervasive processes of degradation include habitat loss and fragmentation. However, a recovery of modified landscape is occurring in some areas.

Objectives

The main goal is to know how lichen and bryophyte epiphytic richness growing on Mediterranean forests is influenced not only by fragments characteristics but also by the structure of the landscape. We introduce a temporal dimension in order to evaluate if the historical landscape structure is relevant for current epiphytic communities.

Methods

40 well-preserved forest fragments were selected in a landscape with a large habitat loss over decades, but with a recovery of forest surface in the last 55 years. The most relevant fragment and landscape-scale attributes were considered. Some of the variables were measured in three different years to incorporate a temporal framework.

Results

The results showed that variables at fragment scale had a higher influence, whereas variables at the landscape scale were irrelevant. Among all the historical variables analyzed, only the shift in forest fragment size had influence on species richness.

Conclusions

Mediterranean forests had suffered fragmentation along centuries. Their epiphytic communities also suffer the hard conditions of Mediterranean climate. Our results indicate that Mediterranean epiphytic communities may be in a threshold since it they will never be similar to those communities existing previous fragmentation process even a recovery habitat occur or, they may require more time to response to this habitat recovery.

Context

Connectivity assessments typically rely on resistance surfaces derived from habitat models, assuming that higher-quality habitat facilitates movement. This assumption remains largely untested though, and it is unlikely that the same environmental factors determine both animal movements and habitat selection, potentially biasing connectivity assessments.

Objectives

We evaluated how much connectivity assessments differ when based on resistance surfaces from habitat versus movement models. In addition, we tested how sensitive connectivity assessments are with respect to the parameterization of the movement models.

Methods

We parameterized maximum entropy models to predict habitat suitability, and step selection functions to derive movement models for brown bear (Ursus arctos) in the northeastern Carpathians. We compared spatial patterns and distributions of resistance values derived from those models, and locations and characteristics of potential movement corridors.

Results

Brown bears preferred areas with high forest cover, close to forest edges, high topographic complexity, and with low human pressure in both habitat and movement models. However, resistance surfaces derived from the habitat models based on predictors measured at broad and medium scales tended to underestimate connectivity, as they predicted substantially higher resistance values for most of the study area, including corridors.

Conclusions

Our findings highlighted that connectivity assessments should be based on movement information if available, rather than generic habitat models. However, the parameterization of movement models is important, because the type of movement events considered, and the sampling method of environmental covariates can greatly affect connectivity assessments, and hence the predicted corridors.

Context

The ability to detect ecological networks in landscapes is of utmost importance for managing biodiversity and planning corridors.

Objectives

The objective of this study was to evaluate the information provided by a synthetic aperture radar (SAR) image for landscape connectivity modeling compared to aerial photographs (APs).

Methods

We present a novel method that integrates habitat suitability derived from remote sensing imagery into a connectivity model to explain species abundance. More precisely, we compared how two resistance maps constructed using landscape and/or local metrics derived from AP or SAR imagery yield different connectivity values (based on graph theory), considering hedgerow networks and forest carabid beetle species as a model.

Results

We found that resistance maps using landscape and local metrics derived from SAR imagery improve landscape connectivity measures. The SAR model is the most informative, explaining 58% of the variance in forest carabid beetle abundance. This model calculates resistance values associated with homogeneous patches within hedgerows according to their suitability (canopy cover density and landscape grain) for the model species.

Conclusions

Our approach combines two important methods in landscape ecology: the construction of resistance maps and the use of buffers around sampling points to determine the importance of landscape factors. This study was carried out through an interdisciplinary approach involving remote sensing scientists and landscape ecologists. This study is a step forward in developing landscape metrics from satellites to monitor biodiversity.

Context

Distribution and connectivity of suitable habitat for species of conservation concern is critical for effective conservation planning. Capercaillie (Tetrao urogallus), an umbrella species for biodiversity conservation, is increasingly threatened because of habitat loss and fragmentation.

Objective

We assessed the impact of drastic changes in forest management in the Carpathian Mountains, a major stronghold of capercaillie in Europe, on habitat distribution and connectivity.

Methods

We used field data surveys with a forest disturbance dataset for 1985–2010 to map habitat suitability, and we used graph theory to analyse habitat connectivity.

Results

Climate, topography, forest proportion and fragmentation, and the distance to roads and settlements best identified capercaillie presence. Suitable habitat area was 7510 km² in 1985; by 2010, clear-cutting had reduced that area by 1110 km². More suitable habitat was lost inside protected areas (571 km²) than outside (413 km²). Habitat loss of 15 % reduced functional connectivity by 33 % since 1985.

Conclusions

Forest management, particularly large-scale clear-cutting and salvage logging, have substantially diminished and fragmented suitable capercaillie habitat, regardless of the status of forest protection. Consequently, larger areas with suitable habitat are now isolated and many patches are too small to sustain viable populations. Given that protection of capercaillie habitat would benefit many other species, including old-growth specialists and large carnivores, conservation actions to halt the loss of capercaillie habitat is urgently needed. We recommend adopting policies to protect natural forests, limiting large-scale clear-cutting and salvage logging, implementing ecological forestry, and restricting road building to reduce forest fragmentation.

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.