Loss and fragmentation of mature woodland reduce the habitat niche breadth of forest birds

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.

     

Matrix is important for mammals in landscapes with small amounts of native forest habitat

Acknowledgment that the matrix matters in conserving wildlife in human-modified landscapes is increasing. However, the complex interactions of habitat loss, habitat fragmentation, habitat condition and land use have confounded attempts to disentangle the relative importance of properties of the landscape mosaic, including the matrix. To this end, we controlled for the amount of remnant forest habitat and the level of fragmentation to examine mammal species richness in human-modified landscapes of varying levels of matrix development intensity and patch attributes. We postulated seven alternative models of various patch habitat, landscape and matrix influences on mammal species richness and then tested these models using generalized linear mixed-effects models within an information theoretic framework. Matrix attributes were the most important determinants of terrestrial mammal species richness; matrix development intensity had a strong negative effect and vegetation structural complexity of the matrix had a strong positive effect. Distance to the nearest remnant forest habitat was relatively unimportant. Matrix habitat attributes are potentially a more important indicator of isolation of remnant forest patches than measures of distance to the nearest patch. We conclude that a structurally complex matrix within a human-modified landscape can provide supplementary habitat resources and increase the probability of movement across the landscape, thereby increasing mammal species richness in modified landscapes.     

Effects of habitat fragmentation on Eurasian red squirrel (Sciurus vulgaris) in a forest landscape

We studied the effects of habitat fragmentation, measured as forest stand size and isolation, on the distribution of Eurasian red squirrels (Sciurus vulgaris). Squirrel density was surveyed during four years in 46 forest stands (0.1–500 ha) in a forest landscape in south-central Sweden. The only factor that significantly influenced a density index was the proportion of spruce within a habitat fragment. Neither fragment size nor degree of isolation were significant. Furthermore, none of the interactions with year were significant, suggesting the same pattern in all four years. Thus, the effect of habitat fragmentation in this study seems to be only pure habitat loss, i.e. halving the proportion of preferred habitat in the landscape should result in a halving of the red squirrel population. Therefore, the landscape can be viewed as functionally continuous for the squirrels, although the preferred habitat was divided into fragments. The most likely explanation for the difference between this study and other studies on squirrels that found effects due to habitat fragmentation is a combination of shorter distances and less hostile surroundings in our study area. To identify landscape effects requires multiple studies because single studies usually consider only one landscape.     

How should we measure landscape connectivity?

The methods for measuring landscape connectivity have never been compared or tested for their responses to habitat fragmentation. We simulated movement, mortality and boundary reactions across a wide range of landscape structures to analyze the response of landscape connectivity measures to habitat fragmentation. Landscape connectivity was measured as either dispersal success or search time, based on immigration into all habitat patches in the landscape. Both measures indicated higher connectivity in more fragmented landscapes, a potential for problematic conclusions for conservation plans. We introduce cell immigration as a new measure for landscape connectivity. Cell immigration is the rate of immigration into equal-sized habitat cells in the landscape. It includes both within- and between-patch movement, and shows a negative response to habitat fragmentation. This complies with intuition and existing theoretical work. This method for measuring connectivity is highly robust to reductions in sample size (i.e., number of habitat cells included in the estimate), and we hypothesize that it therefore should be amenable to use in empirical studies. The connectivity measures were weakly correlated to each other and are therefore generally not comparable. We also tested immigration into a single patch as an index of connectivity by comparing it to cell immigration over the landscape. This is essentially a comparison between patch-scale and landscape-scale measurement, and revealed some potential for patch immigration to predict connectivity at the landscape scale. However, this relationship depends on the size of the single patch, the dispersal characteristics of the species, and the amount of habitat in the landscape. We conclude that the response of connectivity measures to habitat fragmentation should be understood before deriving conclusions for conservation management.     

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

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

Linking movement behavior and fine-scale genetic structure to model landscape connectivity for bobcats (Lynx rufus)

Spatial heterogeneity can constrain the movement of individuals and consequently genes across a landscape, influencing demographic and genetic processes. In this study, we linked information on landscape composition, movement behavior, and genetic differentiation to gain a mechanistic understanding of how spatial heterogeneity may influence movement and gene flow of bobcats in the agricultural landscape of Iowa (USA). We analyzed movement paths of 23 animals to parameterize landscape resistance surfaces, applied least cost path analysis to generate measures of effective geographic distance between DNA collection locations of 625 bobcats, and tested the correlation between genetic distance and the different models of geographic distance. We found that bobcats showed a strong preference for forest over any other habitat type, and that incorporating information on habitat composition both along the path and in the surrounding landscape provided the best model of movement. Measures of effective geographic distance were significantly correlated with genetic distance, but not once the effects of Euclidean distance were accounted for. Thus, despite the impact of habitat composition on movement behavior, we did not detect a signature of a landscape effect in genetic structure. Our results are consistent with the issue of limiting factors: the high uniformity of forest fragmentation across southern Iowa, the primary study area, results in a landscape resistance pattern virtually indistinguishable from the isolation-by-distance pattern. The northern portion of the state, however, is predicted to pose a high level of resistance to bobcat movement, which may impede the regional genetic connectivity of populations across the Midwest.     

The sensitivity of least-cost habitat graphs to relative cost surface values

Maintaining and restoring connectivity among high-quality habitat patches is recognized as an important goal for the conservation of animal populations. To provide an efficient measure of potential connectivity pathways in heterogeneous landscapes, least-cost route analysis has been combined with graph-theoretical techniques. In this study we use spatially explicit least-cost habitat graphs to examine how matrix quality and spatial configuration influence assessments of habitat connectivity. We generated artificial landscapes comprised of three landcover types ranked consistently from low to high quality: inhospitable matrix, hospitable matrix, and habitat. We controlled the area and degree of fragmentation of each landcover in a factorial experiment for a total of 20 combinations replicated 100 times. In each landscape we compared eight sets of relative landcover qualities (cost values of 1 for habitat, between 1.5 and 150 for hospitable matrix, and 3–10,000 for inhospitable matrix). We found that the spatial location of least-cost routes was sensitive to differences in relative cost values assigned to landcover types and that the degree of sensitivity depended on the spatial structure of the landscape. Highest sensitivity was found in landscapes with fragmented habitat and between 20 and 50% hospitable matrix; sensitivity decreased as habitat fragmentation decreased and the amount of hospitable matrix increased. As a means of coping with this sensitivity, we propose identifying multiple low-cost routes between pairs of habitat patches that collectively delineate probable movement zones. These probable movement zones account for uncertainty in least-cost routes and may be more robust to variation in landcover cost values.     

Can we face different types of storms under the same umbrella? Efficiency and consistency of connectivity umbrellas across different patchy landscape patterns

Context

The umbrella approach applied to landscape connectivity is based on the principle that the conservation or restoration of the dispersal habitats for some species also can facilitate the movement of others. Species traits alone do not seem to be enough to identify good connectivity umbrella species, showing the need to investigate the influence of additional factors on this property.

Objectives

We test whether the potential of a species as a connectivity umbrella can be influenced by landscape composition and configuration.

Methods

We simulated movement routes for eight hypothetical species in artificial patchy landscapes with different levels of fragmentation, habitat amount and matrix permeability. We determined the effectiveness of the connectivity umbrella of the virtual species using pairwise intersections of important habitats for their movements in all landscapes.

Results

The connectivity umbrella performance of all species was affected by the interaction of fragmentation level and habitat amount. In general, species performance increased with decreasing fragmentation and increasing habitat amount. In most landscapes and considering the same dispersal threshold, species able to move more easily through the matrix showed higher umbrella performance than those for which the matrix offered greater resistance.

Conclusions

The connectivity umbrella is not a static feature that depends only on the species traits, but rather a dynamic property that also varies according to the landscape attributes. Therefore, we do not recommend spatial transferability of the connectivity umbrella species identified in a landscape to others that have divergent levels of fragmentation and habitat quantity.

     

Circuit theory emphasizes the importance of edge-crossing decisions in dispersal-scale movements of a forest passerine

Measuring landscape connectivity in ways that reflect an animal’s propensity or reluctance to move across a given landscape is key for planning effective conservation strategies. Resistance distance, based on circuit theory, is one such measure relevant for modeling how broad-scale animal movements over long time periods may lead to gene flow across the landscape. Despite the success of circuit theory in landscape genetic studies, its applicability to model finer-scale processes such as the movement patterns of individual animals within their breeding grounds (e.g., while prospecting for territories) has yet to be tested. Here, we applied both circuit models and least-cost models to understand the relationship between landscape connectivity and return time of Ovenbirds (Seiurus aurocapilla) that had been translocated at least 20 km from their home territory near Québec City, Canada. Using an iterative optimization process, we derived resistance values for three cover types (forest, edge, and open) that resulted in resistance distance values that best explained Ovenbird return times. We also identified the cover-type resistance values that yielded length of least-cost path estimates that best explained return times of the translocated birds. The circuit theory and least-cost path methods were equally supported by the data despite being based on different sets of resistance values. The optimal resistance values for calculating resistance distance indicated that for Ovenbirds, traversing a given distance of edge habitat presented a substantially greater resistance than that of open areas. On the other hand, optimized resistances of edge and open were very similar for calculating length of least-cost path. The circuit theory approach suggested that for an Ovenbird moving through fragmented habitat, the number of forest-open transitions (i.e., edge-crossings) that an individual must make is critical to understanding return times after translocation. The least-cost path approach, on the other hand, suggested that the birds strongly avoid all open areas, regardless of size. Circuit theory offers an important new approach for understanding landscapes from the perspective of individuals moving within their breeding range, at finer spatial scales and shorter time scales than have been previously considered.     

The importance of functional connectivity in the conservation of a ground-dwelling mammal in an urban Australian landscape

The distribution of the northern brown bandicoot (Isoodon macrourus), a medium-sized ground-dwelling marsupial, was examined in habitat fragments within the urban landscape of the city of Brisbane, Australia. From surveys conducted in 68 fragments, bandicoots were found to be present in 33 (49%) despite widespread habitat loss and fragmentation. Logistic regression analysis revealed that of 13 measured independent variables, functional connectivity was the only factor that significantly predicted the presence of bandicoots within fragments, with connectivity positively correlated with the likelihood of occupation. Functional connectivity was equated to the likelihood of bandicoot immigration into the focal fragment from the nearest occupied fragment, based on the estimated resistance to movement offered by the intervening matrix. Within Brisbane, riparian habitat fragments typically have a relatively high level of functional connectivity, as thin strips of vegetation fringing waterways serve as corridors between larger riparian areas and facilitate the movement of bandicoots between patches. Analyses based on the Akaike Information Criterion revealed that the optimal model based on landscape context variables was convincingly better supported by the data than the optimal model produced from fragment characteristics. However, it is important to examine both internal attributes of habitat fragments and external features of the surrounding landscape when modelling the distribution of ground-dwelling fauna in urban environments, or other landscapes with a highly variable matrix. As urban centres throughout the world expand, it is crucial that the ecology of local wildlife be considered to ensure functional connection is maintained between habitat patches, especially for the conservation of species that are highly susceptible to fragmentation.     

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.     

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

Context

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

Objectives

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

Methods

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

Results

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

Conclusions

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

     

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.     

Dispersal success on fractal landscapes: a consequence of lacunarity thresholds

Habitat fragmentation is expected to disrupt dispersal, and thus we explored how patch metrics of landscape structure, such as percolation thresholds used to define landscape connectivity, corresponded with dispersal success on neutral landscapes. We simulated dispersal as either a purely random process (random direction and random step lengths) or as an area-limited random walk (random direction, but movement limited to an adjacent cell at each dispersal step) and quantified dispersal success for 1000 individuals on random and fractal landscape maps across a range of habitat abundance and fragmentation. Dispersal success increased with the number of cells a disperser could search (m), but poor dispersers (m<5) searching via area-limited dispersal on fractal landscapes were more successful at locating suitable habitat than random dispersers on either random or fractal landscapes. Dispersal success was enhanced on fractal landscapes relative to random ones because of the greater spatial contagion of habitat. Dispersal success decreased proportionate to habitat loss for poor dispersers (m=1) on random landscapes, but exhibited an abrupt threshold at low levels of habitat abundance (p<0.1) for area-limited dispersers (m<10) on fractal landscapes. Conventional metrics of patch structure, including percolation, did not exhibit threshold behavior in the region of the dispersal threshold. A lacunarity analysis of the gap structure of landscape patterns, however, revealed a strong threshold in the variability of gap sizes at low levels of habitat abundance (p<0.1) in fractal landscapes, the same region in which abrupt declines in dispersal success were observed. The interpatch distances or gaps across which dispersers must move in search of suitable habitat should influence dispersal success, and our results suggest that there is a critical gap-size structure to fractal landscapes that interferes with the ability of dispersers to locate suitable habitat when habitat is rare. We suggest that the gap structure of landscapes is a more important determinant of dispersal than patch structure, although both are ultimately required to predict the ecological consequences of habitat fragmentation.     

Maintenance of flying squirrel habitat and timber harvest: a site-specific spatial model in forest planning calculations

Spatial and temporal continuity of resources often benefits both ecological and economic goals in landscape management. Consideration of multiple and conflicting goals is also needed to view the future production possibilities of forests in successful forest management. Our aim was to estimate the production potential of a planning area in Finland by examining different forest management strategies from ecological and economic perspectives using long-term forest planning calculations. Economic objectives referred to timber production, whereas ecological objectives were based on suitable habitats for arboreal Siberian flying squirrel (Pteromys volans). Suitable habitats were defined using an empirical site-specific model, which includes a spatial variable reflecting the availability of habitat within an individual’s activity area. Five alternative forest plans were worked out with different objectives for flying squirrel habitat and timber production. The alternative plans were compared with respect to values of objective variables at the end of the planning period of 60 years and against a production possibility frontier among net present value and flying squirrel habitat. Varying objective values in our analyses resulted from different utilization of production possibilities, and the changes were in line with the objectives used. The formation of flying squirrel habitat clusters in the landscape was enhanced, and it did not always incur severe reductions in harvestable timber volume. Possibilities to combine ecological and economic goals, both spatial and aspatial, in the planning process seems to be an encouraging alternative for the long-term forest management in the future.     

Quantifying landscape connectivity through the use of connectivity response curves

Habitat connectivity is an essential component of biodiversity conservation. Simulated landscapes were manipulated to quantify the impact of changes to the amount, fragmentation and dispersion of habitat on a widely applied landscape connectivity metric, the probability of connectivity index. Index results for different landscape scenarios were plotted against the dispersal distances used for their calculation to create connectivity response curves for each scenario. Understanding index response to controlled changes in landscape structure at a range of spatial scales can be used to give context to comparison of alternative landscape management scenarios. Increased amounts of habitat, decreased fragmentation and decreased inter-patch distances resulted in increased connectivity index values. Connectivity response curves demonstrated increases in assessed connectivity for scenarios with continuous corridors or “stepping stone” connectors. The sensitivity of connectivity response curves to controlled changes in landscape structure indicate that this approach is able to detect and distinguish between different types of landscape changes, but that delineation of habitat and method of quantifying dispersal probability incorporate assumptions that must be recognized when interpreting results to guide landscape management. Representing landscape connectivity in this manner allows for the impacts of alternative landscape management strategies to be compared visually through comparative plots, or statistically through the parameters that describe connectivity response curves.     

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.     

Spatial patterns of bird community similarity: bird responses to landscape composition and configuration in the Atlantic forest

Studies dealing with community similarity are necessary to understand large scale ecological processes causing biodiversity loss and to improve landscape and regional planning. Here, we study landscape variables influencing patterns of community similarity in fragmented and continuous forest landscapes in the Atlantic forest of South America, isolating the effects of forest loss, fragmentation and patterns of land use. Using a grid design, we surveyed birds in 41 square cells of 100 km² using the point count method. We used multivariate, regression analyses and lagged predictor autoregressive models to examine the relative influence of landscape variables on community similarity. Forest cover was the primary variable explaining patterns of bird community similarity. Similarity showed a sudden decline between 20 and 40% of forest cover. Patterns of land use had a second order effect; native bird communities were less affected by forest loss in landscapes dominated by tree plantations (the most suitable habitat for native species) than in landscapes dominated by annual crops or cattle pastures. The effects of fragmentation were inconclusive. The trade-off between local extinctions and the invasion of extra-regional species using recently created habitats is probably the mechanism generating the observed patterns of community similarity. Limiting forest loss to 30–40% of the landscape cover and improving the suitability of human-modified habitats will contribute to maintain the structure and composition of the native forest bird community in the Atlantic forest.     

Effects of historical and present fragmentation on plant species diversity in semi-natural grasslands in Swedish rural landscapes

Habitat loss and fragmentation of natural and semi-natural habitats are considered as major threats to plant species richness. Recently several studies have pinpointed the need to analyse past landscape patterns to understand effects of fragmentation, as the response to landscape change may be slow in many organisms, plants in particular. We compared species richness in continuously grazed and abandoned grasslands in different commonplace rural landscapes in Sweden, and analysed effects of isolation and area in three time-steps (100 and 50 years ago and today). Old cadastral maps and aerial photographs were used to analyse past and present landscape patterns in 25 sites. Two plant diversity measures were investigated; total species richness and species density. During the last 100 years grassland area and connectivity have been reduced by about 90%. Present-day habitat area was positively related to total species richness in both habitats. There was also a relationship to habitat area 50 years ago for continuously grazed grasslands. Only present management was related to species density: continuously grazed grasslands had the highest species density. There were no relationships between grassland connectivity, present or past, and any diversity measure. We conclude that landscape history is not directly important for present-day plant diversity patterns in ordinary landscapes, although past grassland management is a prerequisite for the grassland habitats that can be found there today. It is important that studies are conducted, not only in very diverse landscapes, but also in managed landscapes in order to assess the effects of fragmentation on species.     

The Implications of Metalandscape Connectivity for Population Viabilityin Migratory Songbirds

Landscape connectivity is considered important for species persistence, but linkages among landscape populations (metalandscape connectivity) may be necessary to ensure the long-term viability of some migratory songbirds at a broader regional scale. Because of regional source-sink dynamics, these species can maintain steady populations within extensively fragmented landscapes (landscape sinks) owing to high levels of immigration from source landscapes. We undertook a modeling study to identify the conditions under which immigration, an index of metalandscape connectivity, could rescue declining populations of songbirds in heavily disturbed landscapes. In general, low to moderate levels of immigration (m = 0–20%) were sufficient to rescue species with low edge-sensitivity in landscapes where<70% habitat had been destroyed. At the other extreme, moderate to high levels of immigration (m = 11–40%) were usually required to rescue highly edge-sensitive species in these same landscapes. Very high levels of immigration (m>40%) were required to rescue highly edge-sensitive species in extensively fragmented landscapes that had lost >50% habitat, or when any landscape lost ≥50% habitat gradually over a period of 100 or more years (r = 0.5% habitat lost/year). Paradoxically higher levels of immigration were thus necessary to offset population declines when habitat was lost gradually than when it was lost quickly, where population response lagged behind landscape change. This implies that the importance of metalandscape connectivity for population viability may not be fully appreciated in landscapes undergoing rapid rates of change. Natural immigration rates for migratory songbirds match the very high levels (>40%) we found necessary to sustain populations in heavily disturbed landscapes, which underscores the importance of metalandscape connectivity for the continued persistence of many migratory songbirds in the face of widespread habitat loss and fragmentation.