Habitat Selection and Breeding Success in a Forest-nesting Alcid, the Marbled Murrelet, in Two Landscapes with Different Degrees of Forest Fragmentation

We studied habitat selection and breeding success in marked populations of a protected seabird (family Alcidae), the marbled murrelet (Brachyramphus marmoratus), in a relatively intact and a heavily logged old-growth forest landscape in south-western Canada. Murrelets used old-growth fragments either proportionately to their size frequency distribution (intact) or they tended to nest in disproportionately smaller fragments (logged). Multiple regression modelling showed that murrelet distribution could be explained by proximity of nests to landscape features producing biotic and abiotic edge effects. Streams, steeper slopes and lower elevations were selected in both landscapes, probably due to good nesting habitat conditions and easier access to nest sites. In the logged landscape, the murrelets nested closer to recent clearcuts than would be expected. Proximity to the ocean was favoured in the intact area. The models of habitat selection had satisfactory discriminatory ability in both landscapes. Breeding success (probability of nest survival to the middle of the chick rearing period), inferred from nest attendance patterns by radio-tagged parents, was modelled in the logged landscape. Survivorship was greater in areas with recent clearcuts and lower in areas with much regrowth, i.e. it was positively correlated with recent habitat fragmentation. We conclude that marbled murrelets can successfully breed in old-growth forests fragmented by logging.     

The need for biological realism in habitat modeling: a reinterpretation of Zharikov et al. (2006)

Zharikov et al. (2006: Landscape Ecology 21:107–120) modeled the nest-site habitat use of marbled murrelets (Brachyramphus marmoratus) in Desolation Sound (DS) and Clayoquot Sound (CS), British Columbia. They compared known nest sites, located with radio-telemetry, with randomly-located points within the same areas. Their conclusions suggest that murrelets tended to nest in disproportionately smaller fragments within the more disturbed DS landscape; streams, steeper slopes, and lower elevations were selected in both landscapes; murrelets nested closer to recent clearcuts than would be expected in the DS landscape; and survivorship of nestlings was greater in areas with recent clearcuts and was positively correlated with recent habitat fragmentation. These conclusions are contrary to current management guidelines in British Columbia, and therefore require close scrutiny. Our detailed examination reveals flaws in their use of data, application of modeling, and most seriously, interpretation of the results. Problems include: conceptual errors in the interpretation of models; inappropriate spatial resolution; confusing use and interpretation of fragmentation and patch size data; overemphasis of statistically significant but biologically trivial results; and ignoring some contradictory studies. We conclude that it would be risky to apply the results from Zharikov et al. in the selection of murrelet nesting habitat for management purposes in British Columbia. Our review identifies issues that may arise in other ecological modeling studies and stresses the need for biological realism in addition to statistical rigour.     

Interpreting habitat distribution models of an elusive species,the marbled murrelets: a response to Burger and Page

Burger and Page (this volume) evaluated our models of habitat preferences and breeding success of a threatened seabird, the marbled murrelet (Brachyramphus marmoratus), based on the largest available set of confirmed nest-sites found in coastal old-growth forest of the Pacific North-West. We believe our study documented novel and unexpected patterns of landscape-level distribution of marbled murrelets in both heavily logged and relatively intact old-growth landscapes and provided insights into how these patterns influence their reproduction, and, eventually, management. Considering the importance of the issue and to ensure appropriate and responsible use of the information we welcome discussion, detailed scrutiny and evaluation of our original results. Burger and Page claim to have identified flaws with model interpretation, data quality, statistical approaches, presentation and interpretation of our results that would invalidate our conclusions. We respond that most of their critique is irrelevant and/or misdirected with respect to our study and the interpretation of GIS data models, and that valid aspects of their claims do not critically affect our conclusions.     

Effects of landscape structure and forest reserve location on old-growth forest bird species in Northern Finland

Old-growth forest birds in Fennoscandia have sharply declined in numbers during the last decades apparently due to commercial forest harvesting and fragmentation of old-growth forests. Conservation measures have led to the establishment of a forest reserve network to assure the persistence of forest birds at a regional scale. However, little is known about the effects of landscape structure within and around the reserves on the distribution of old-growth forest birds. We used a hierarchical approach to address the questions of how landscape structure and composition within forest reserves, landscape composition of surrounding areas and reserve location affect the abundance of resident, old-growth forest birds in the Northern Finnish forest reserve network. The positive role of particular landscape features on bird distribution indicates that both the proportion of old-growth forests and the structure of boreal landscape mosaic has an important role in determining the distribution of these birds. The landscape composition surrounding the reserves proved to be only a weak predictor in species distribution models, which argues against the primary role of the surrounding matrix in determining species distribution within forest reserves. Reserves located near the Russian border showed a higher abundance of old-growth birds than more western ones. Once east-west gradients in overall landscape composition had been accounted for, however, reserves did not differ significantly in the number of species present. These results suggest that landscape gradients, rather than ecological processes such as the presence of source areas located along the border with Russia, are the main determinant of the distribution of old-growth forest birds in the Finnish reserve network. We propose that to enhance regional persistence of old-growth forest birds, conservation efforts should be primarily directed towards the protection and enhancement of forest habitat quality and natural heterogeneity of landscapes within targeted areas. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

The influence of forest harvesting on landscape spatial patterns and old-growth-forest fragmentation in southeast British Columbia

Habitat fragmentation is considered one of the major conservation issues of recent decades. We tested predictions of landscape patterns in a 352,253-ha managed forest area in southeast British Columbia. We did this by focussing on forest fragmentation concerns among old-growth, harvest, and wildfire patches in 44 delineated landscapes using patch indices as measures of landscape pattern. We found no significant association between amount of harvesting and 15 old-growth patch indices. Comparisons among patch types revealed that amounts and spatial patterns of harvest patches differed little from amounts and spatial patterns of old-growth patches in control landscapes. Variability indices revealed similar variability between harvest patches and old-growth patches, and more variability between harvest patches and wildfire patches. Little of the evidence gathered in this study supported predictions of fragmentation of old-growth spatial patterns, or predicted differences between harvest spatial patterns and more naturally occurring spatial patterns. We suggest these results could be due to the relatively small amounts of harvesting and old-growth forest in these landscapes, and therefore habitat amount may be a more important factor than spatial configuration of patches in these landscapes.     

Response of snails and slugs to fragmentation of lowland forests in NW Germany

Habitat fragmentation is a major cause for species loss, but its effect on invertebrates with low active dispersal power, like terrestrial gastropods, has rarely been studied. Such species can not cross a hostile habitat matrix, for which the predictions of island theory, such as positive relations between species richness and patch size, should apply. In order to test this prediction, we studied gastropod species diversity by assessing gastropod assemblage characteristics from 35 sites in 19 fragments of deciduous old-growth forests in the Lower Rhine Embayment, Germany. Assemblages differed between larger (≥700 ha) and smaller forests (<400 ha), those of large forests held a higher percentage of forest species. Although α-diversity was similar between the two forest size classes, small forests often comprised matrix species, resulting in a higher β-diversity. Edge effects on the species richness of matrix species were noticeable up to 250 m into the forest. Hierarchical partitioning revealed that distance to disturbances (external edge, internal edges like roads) explained most assemblage variables, whereas forest size and woodland cover within a 1 km radius from the sites explained only a few assemblage variables. Densities of two forest-associated species, Discus rotundatus and Arion fuscus, decreased with forest size. Yet, forest size was positively correlated with richness of typical forest species and densities of Limax cinereoniger. The latter species seems to need forests of >1,000 ha, i.e., well above the size of most fragments. In conclusion, the prediction is valid only for forest species. The response to fragmentation is species specific and seems to depend on habitat specialization and macroclimatic conditions. Jean-Pierre Maelfait: Deceased.     

Using landscape metrics to model source habitat for Neotropical migrants in the midwestern U.S.

Size of a forest patch is a useful predictor of density and reproductive success of Neotropical migratory birds in much of eastern North America. Within these forested landscapes, large forest tracts appear to be sources – fragments in which surpluses of offspring are produced and can potentially colonize new fragments including woodlot sinks where reproduction fails to balance adult mortality. Within agricultural landscapes of the midwestern U.S., where forests are severely fragmented, high levels of brood parasitism by brown-headed cowbirds (Molothrus ater) and intense predation on nests generally result in low reproductive success for Neotropical migrants regardless of forest size. In some midwestern U.S. landscapes, however, the variation in reproductive success among forest fragments suggests that `source' habitat could still exist for Neotropical migrants. We used vegetation, fragment and landscape metrics to develop multivariate models that attempt to explain the variation in abundance and reproductive success of Neotropical migrants nesting in an agricultural landscape in northern Indiana, USA. We produced models that reasonably described the pattern of species richness of Neotropical migrants and the abundance of wood thrushes (Hylocichla mustelina) and several other Neotropical migrant species within 14 forest fragments. In contrast, we were unable to produce useful models of the reproductive success of wood thrushes breeding in the same forest fragments. Our results suggest that (1) abundance patterns of Neotropical migrants are probably influenced by both landscape- and fragment-scale factors; (2) multivariate analyses of Neotropical migrant abundance are not useful in modeling the corresponding patterns of reproductive success; and (3) the location of any remaining `source' habitat for Neotropical migrants breeding within agricultural landscapes in North America will be difficult to predict with indirect measures such as vegetation composition or landscape context. As a result, the potential for developing conservation strategies for Neotropical migrants will be limited without labor-intensive, direct measurements of demographic parameters.     

Identification of Landscape Elements Related to Local Declines of a Boreal Grey-sided Vole Population

Several studies indicate a long-term decline in numbers of different species of voles in northern Fennoscandia. In boreal Sweden, the long-term decline is most pronounced in the grey-sided vole (Clethrionomys rufocanus). Altered forest landscape structure has been suggested as a possible cause of the decline. However, habitat responses of grey-sided voles at the landscape scale have never been studied. We analyzed such responses of this species in lowland forests in Västerbotten, northern Sweden. Cumulated spring densities representing 22 local time series from 1980–1999 were obtained by a landscape sampling design and were related to the surrounding landscape structure of 2.5×2.5 km plots centred on each of the 22 1-ha trapping plots. In accordance with general knowledge on local habitat preferences of grey-sided voles, our study supported the importance of habitat variables such as boulder fields and old-growth pine forest at the landscape scale. Densities were negatively related to clear cuts. Habitat associations were primarily those of landscape structure related to habitat fragmentation, distance between habitat patches and patch interspersion rather than habitat patch type quantity. Local densities of the grey-sided vole were positively and exponentially correlated with spatial contiguity (measured with the fragmentation index) of old-growth pine forest, indicating critical forest fragmentation thresholds. Our results indicate that altered land use might be involved in the long-term decline of the grey-sided vole in managed forest areas of Fennoscandia. We propose two further approaches to reveal and test responses of this species to changes in landscape structure.     

Siberian flying squirrel responses to high- and low-contrast forest edges

We examined responses of Siberian flying squirrels ( Pteromys volans ) to edges between nesting habitat (mature spruce forests), movement habitat (other forests, pine bogs), and open areas within their home ranges in southern Finland in 1996-2000. Radio-tracked squirrels (n=146) were generally associated to edges when they were ac tive at night. Compared to distances expected from the habitat pattern of their home range, squirrels occurred closer to high-contrast edges (of open areas) and low-contrast edges (nesting or movement forest types). Asso ciation with edges of open areas was more pronounced when squirrels were in movement habitat than in nesting habitat, possibly because of stronger channeling of movements in the former habitat. When in nesting habitat, squirrels responded more strongly to field edges than to recent clearcut edges, probably as a result of the pres ence of more deciduous trees on field edges, unlike clearcut edges. Responses to open areas were independent of spatial scale. However, responses to movement habitat from nesting habitat, and vice versa, were more pronounced over hundreds than tens of meters. Nesting cavities and dreys were generally located at random with respect to edges. We conclude that squirrel responses to edges of landscape attributes are diverse and depend both on spatial scale and edge contrast.This revised version was published online in May 2005 with corrections to the Cover Date.     

Multi-scale use of lands providing anthropogenic resources by American Crows in an urbanizing landscape

The conversion of forests and farmlands to human settlements has negative impacts on many native species, but also provides resources that some species are able to exploit. American Crows (Corvus brachyrhynchos), one such exploiter, create concern due to their impact as nest predators, disease hosts, and cultural harbingers of evil. We used various measures of crow abundance and resource use to determine crows’ response to features of anthropogenic landscapes in the Puget Sound region of the United States. We examined land cover and land use composition at three spatial scales: study sites (up to 208 ha), crow home ranges within sites (18.1 ha), and local land cover (400 m²). At the study site and within-site scales crow abundance was strongly correlated with land cover providing anthropogenic resources. In particular, crows were associated with the amount of ‘maintained forest’ cover, and were more likely to use grass and shrub cover than forest or bare soil cover. Although crows did not show a generalized response to an edge variable, they exhibited greater use of patchy habitat created by human settlements than of native forests. Radio-tagged territorial adults used resources within their home ranges relatively evenly, suggesting resource selection had occurred at a larger spatial scale. The land conversion pattern of new suburban and exurban settlements creates the mix of impervious surfaces and maintained vegetation that crows use, and in our study area crow populations are expected to continue to increase. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Estimating the consequences of habitat fragmentation on extinction risk in dynamic landscapes

Analyzing the population dynamic consequences of spatio-temporal changes in landscape structure is a formidable challenge for spatial ecology. One key population dynamic process in fragmented landscapes is the influence of isolation on colonization rate and thereby on the occurrence of species in habitat fragments, but it is not obvious how isolation should be measured in landscapes that are affected by on-going habitat loss and fragmentation. We suggest the following procedure for the measurement of spatio-temporal isolation. First, a historical record of habitat loss and fragmentation in the landscape is prepared based on snapshots of the extent of the suitable habitat for the focal species. Second, a metapopulation model is used to simulate the occurrence of the species in this landscape, assuming the empirically observed landscape change. The model-predicted pattern of habitat occupancy at a particular point in time (usually the present time) is then compared with empirical observations on the occurrence of the species. We describe a metapopulation model that has been constructed for this purpose, and we apply it to a changing landscape of boreal forests in eastern Finland. We give an example on the occurrence of four threatened polyporous fungi in 18 small fragments of old-growth forest. In none of the species does the current isolation of the fragments nor the time since their isolation explain the occurrence of the species in the study fragments, but in three species the model-predicted occupancy probability had a significant effect on the observed abundance of the species. The model-predicted occupancy probabilities were also calculated by ignoring past landscape changes, that is, by assuming that the landscape had remained in the present configuration for a long time. These probabilities had a significant effect on the abundance of only one of the four species, suggesting that the occurrence of the species tracks landscapes changes with a noticable time lag.This revised version was published online in May 2005 with corrections to the Cover Date.     

Scale-dependent proximity of wildlife habitat in a spatially-neutral Bayesian model

Organisms may be constrained by the energetic costs incurred while obtaining resources in fragmented landscapes. We used a spatially neutral model of deer wintering habitat to evaluate the effects of landscape fragmentation on the aggregation of deer habitat. The spatially neutral model used Bayesian probabilities to predict where deer wintering areas occurred. The probabilities were conditional on 12 landscape variables measured at 22,750 contiguous 0.4 ha locations. The model predicted deer habitat at each location independently, thereby enabling a comparison of habitat aggregation in observed, simulated, and random distributions of deer habitat. The predictions of the neutral model exhibited greater fragmentation than observed in nature, suggesting that suitable, yet isolated, locations were not visited by deer. The most suitable sites for deer were clumped in the neutral model, regardless of scale. The inclusion of less suitable sites preserved significant aggregation at fine scales but not at broad scales. Species operate at different scales within a landscape, so ecologists, nature reserve designers and natural resource planners may benefit from models that focus on the proximity of habitat sites as a function of both spatial scale and habitat quality.     

Landscape context and selection for forest edge by breeding Brown-headed Cowbirds

We evaluated support for four alternate hypotheses explaining the distribution of breeding Brown-headed Cowbirds (Molothrus ater) in forests at varying distances from the forest edge in three Midwestern USA landscapes with varying amounts of forest fragmentation (core forest area ranged from 5 to 70%). We focused on breeding cowbirds’ use of forest because of the risk of nest parasitism to forest-dwelling hosts and to identify factors affecting breeding cowbird habitat selection. We compared distances of cowbird locations in the forest from the forest edge (“edge distances”) to distances of random forest locations in the entire landscape or within individual cowbird home ranges. We analyzed 1322 locations of 84 cowbirds across three landscapes. We found support for the landscape context hypothesis that breeding cowbird preference for forest edge varied with landscape context. Ninety percent of cowbird locations were within 150–350 m of forest edge, despite the overall availability of forest at greater distances from edge (as far as 500–1450 m) both within cowbird home ranges and the entire forested landscape. Cowbird preference for edge varied by landscape context largely due to differences in the availability of forest edge. In a highly fragmented forest cowbirds utilized the entire forest and likely viewed it as “all edge.” In less fragmented forests, cowbirds preferred edge. We consider how variation in cowbird edge preference might relate to patterns in host abundance, host diversity, and host quality because cowbird movements indicate they are capable of using forest farther from edges.     

The influence of sampling scheme and interpolation method on the power to detect spatial effects of forest birds in Ontario (Canada)

Spatial ecology is becoming an increasingly important component of resource management, and the general monitoring of how human activities affect the distribution and abundance of wildlife. Yet most work on the reliability of sampling strategies is based on a non-spatial analysis of variance paradigm, and little work has been done assessing the power of alternative spatial methods for creating reliable maps of animal abundance. Such a map forms a critical response variable for multiple scale studies relating landscape structure to biotic function. The power to reconstruct patterns of distribution and abundance is influenced by sample placement strategy and density, the nature of spatial auto-correlation among points, and by the technique used to extrapolate points into an animal abundance map. Faced with uncertainty concerning the influence of these factors, we chose to first synthesize a model reference system of known properties and then evaluate the relative performance of alternative sampling and mapping procedures using it. We used published habitat associations of tree nesting boreal neo-tropical birds, a classified habitat map from the Manitou Lakes area of northwestern Ontario, and point count means and variances determined from field studies in boreal Canada to create 4 simulated models of avian abundance to function as reference maps. Four point sampling strategies were evaluated by 4 spatial mapping methods. We found mixed-cluster sampling to be an effective point sampling strategy, particularly when high habitat fragmentation was avoided by restricting samples to habitat patches >10 ha in size. We also found that of the 4 mapping methods, only stratified ordinary point kriging (OPK) was able to generate maps that reproduced an embedded landscape-scale spatial effect that reduced nesting bird abundance in areas of higher forest age-class fragmentation. Global OPK was effective only for detecting broader, regional-scale differences. This revised version was published online in July 2006 with corrections to the Cover Date.     

The relative impact of forest patch and landscape attributes on black howler monkey populations in the fragmented Lacandona rainforest, Mexico

Land-use change is forcing many animal populations to inhabit forest patches in which different processes can threaten their survival. Some threatening processes are mainly related to forest patch characteristics, but others depend principally on the landscape spatial context. Thus, the impact of both patch and landscape spatial attributes needs to be assessed to have a better understanding of the habitat spatial attributes that constraint the maintenance of populations in fragmented landscapes. Here, we evaluated the relative effect of three patch-scale (i.e., patch size, shape, and isolation) and five landscape-scale metrics (i.e., forest cover, fragmentation, edge density, mean inter-patch isolation distance, and matrix permeability) on population composition and structure of black howler monkeys (Alouatta pigra) in the Lacandona rainforest, Mexico. We measured the landscape-scale metrics at two spatial scales: within 100 and 500 ha landscapes. Our findings revealed that howler monkeys were more strongly affected by local-scale metrics. Smaller and more isolated forest patches showed a lower number of individuals but at higher densities. Population density also tended to be positively associated to matrices with higher proportion of secondary forests and arboreal crops (i.e. with greater permeability), most probably because these matrices can offer supplementary foods. The immature-to-female ratio also increased with matrix permeability, shape complexity, and edge density; habitat characteristics that can increase landscape connectivity and sources availability. The prevention of habitat loss and isolation, and the increment of matrix permeability are therefore needed for the conservation of this endangered Neotropical mammal.     

Landscape level effects of modern forestry on bird communities in North Swedish boreal forests

We address effects of large-scale forestry on landscape structure and the structure and composition of boreal bird communities in North Sweden. Specifically, we ask: after controlling for the effect of patch size, forest age and tree species composition, is there any residual effect attributable to the reduction in area of old forest? Pairs of landscape blocks (25 by 25 km) were selected to maximize area difference in human-induced disturbance, clear-cut as opposed to semi-natural old forest. Median distance to natural edge (wetlands, open water) from randomly selected points in forest was 250 and 200 m in high and low impact landscapes, respectively, indicating a high degree of ‘natural’ fragmentation of the pristine boreal landscape in the area. By contrast, median distance to clear-cut in uncut forest was 750 and 100 m, respectively. Clear-cuts in high impact landscapes were disproportionally more common in areas with contiguous forest land than in areas with spatially disjunct forest, implicating that forestry increases natural fragmentation of the landscape by subdividing larger forest tracts. Point counts along forestry roads showed that species richness and relative abundance of forest birds were higher in landscapes with low forestry impact. These differences can partly be explained by differences in age composition of forest and composition of tree species. After controlling for patch size, forest age and tree species composition, a significant effect of forestry impact remained for Sibirian species and the Tree pipitAnthus trivialis. Our results thus imply that this group of species and the Tree pipit may be sensitive to forest fragmentation. In contrast to previous Finnish studies, we found relatively small negative effects on relative abundance of species hypothesized to be negatively affected by large-scale clear-cutting forestry. However, our picture of the present does not contradict results from Finnish long-term population studies. Five factors may account for this: 1) clear-cut areas are not permanently transformed into other land use types, 2) planted forests are not completely inhabitable for species preferring older forest, 3) the majority of species in the regional pool are habitat generalists, 4) the region studied is still extensively covered with semi-natural forest, and 5) our study area is relatively close to contiguous boreal forest in Russia, a potential source area for taiga species.     

Modeling configuration dynamics of harvested forest landscapes in the Canadian boreal plains

Habitat configuration has important implications for the persistence of faunal and floral populations at a variety of spatial scales. Forest harvesting alters habitat configurations. However, measuring and predicting such alterations remains challenging, in part because previously developed metrics of habitat configuration are often not statistically independent of habitat amount. Thus, their ability to measure independent effects of habitat configurations and habitat amount on ecosystem components such as wildlife populations has been limited. Here, we evaluate habitat configuration based on newly developed metrics that are independent of habitat amount but do not depend on regression residuals of abundance and configuration relationships on any population of landscapes. We use these new metrics to measure and predict changes in habitat configuration following forest harvesting in the boreal forest of Alberta, Canada. Our findings clearly demonstrate changes in habitat configuration resulting from forest harvesting can be predicted precisely with information about initial habitat patch structure and harvesting patterns. Because forest harvesting has significant implications for habitat configuration, accurately predicting these changes is critical for determining if forest harvesting strategies are sustainable for ecosystem components and processes. This study provides a set of novel, robust metrics for tracking landscape-scale changes in habitat configuration in harvested boreal forests.     

Assessing habitat fragmentation on marine epifaunal macroinvertebrate communities: an experimental approach

Habitat fragmentation is considered a major cause of biodiversity loss, both on terrestrial and marine environments. Understanding the effects of habitat fragmentation on the structure and dynamics of natural communities is extremely important to support management actions for biodiversity conservation. However, the effects of habitat fragmentation on marine communities are still poorly understood. Here we evaluated whether habitat fragmentation affects the structure of epifaunal communities in the sublittoral zone, in the northern coast of São Paulo state, Brazil. Five experimental landscapes were constructed, each one forming a large continuous patch. After 4 weeks, each landscape was cut on three patches of different sizes. Epifaunal macroinvertebrate communities were sampled at the edge and interior of experimental landscapes before manipulation to evaluate edge effects. After four more weeks, communities from the three patch sizes were also sampled to evaluate patch size effects. We compared the diversity of communities at different levels of fragmentation by total abundance, rarefied taxon richness, Shannon–Wiener diversity index, Simpson’s dominance index, and abundance of dominant taxa. Higher taxon richness and gastropod abundance were recorded in the patch edges, but no significant differences were found among patch sizes. We found a significant effect of habitat fragmentation, with lower abundances of Gammaridea (the dominant taxon), Ophyuroidea, and Pycnogonida after the experimental fragmentation. Lower abundances of dominant taxa resulted in higher diversity and lower dominance in fragmented landscapes when compared to integral, pre-manipulation landscapes. Our results suggest that fragmentation of landscapes in the system studied can reduce dominance, and that even small patch sizes can be important for the conservation of macroinvertebrate diversity.     

Positive effects of forest fragmentation,independent of forest amount,on bat abundance in eastern Ontario,Canada

While studies have found that bat abundance is positively related to the amount of forest cover in a landscape, the effects of forest fragmentation (breaking apart of forest, independent of amount) are less certain, with some indirect evidence for positive effects of fragmentation. However, in most of these studies, the variables used to quantify fragmentation are confounded with forest amount, making it difficult to interpret the results. The purpose of this study was to examine how forest amount and forest fragmentation independently affect bat abundance. We conducted acoustic bat surveys at the centers of 22 landscapes throughout eastern Ontario, Canada, where landscapes were chosen to avoid a correlation between forest amount and forest fragmentation (number of patches) at multiple spatial scales, while simultaneously controlling for other variables that could affect bat activity. We found that the effects of forest amount on bat relative abundance were mixed across species (positive for Lasiurus borealis, negative for Perimyotis subflavus and Lasionycteris noctivagans). When there was evidence for an effect of forest fragmentation, independent of forest amount, on bat relative abundance, the effect was positive (Myotis septentrionalis, Myotis lucifugus and Lasiurus borealis). We suggest that the mechanism driving the positive responses to fragmentation is higher landscape complementation in more fragmented landscapes; that is, increased access to both foraging and roosting sites for these bat species. We conclude that fragmented landscapes that maximize complementation between roosting and foraging sites should support a higher diversity and abundance of bats.