Metapopulation theory and habitat fragmentation: a review of holarctic breeding bird studies

Metapopulations are conceived as spatially structured populations consisting of distinct units (subpopulations), separated by space or barriers, and connected by dispersal movements. Metapopulations characteristically demonstrate a turnover of local populations going extinct and becoming re-established, resulting in a distribution pattern that shifts over time. Metapopulation theory is used to analyse the effects of habitat fragmentation on birds in the temperate zone, integrating various explanations for the paucity of species in isolated ecotopes.There is some evidence that turnover of local populations occurs in fragmented systems. A few studies based on time series demonstrate the local extinction rate to be related to the size of the habitat fragment, whereas the recolonization rate depends on the degree of isolation. Most evidence comes from short-term pattern studies in which the probability of occurrence was found to depend on the size of habitat fragments, on their relative position in the landscape and on the density of corridors lowering the landscape resistance. These data are consistent with predictions from metapopulation theory. However, almost all investigations consider wood fragmentation in agricultural landscapes, and there is a great need for studies in naturally fragmented landscapes as well as for studies focussing on other, less predictable, habitat types.     

The importance of integrating landscape ecology in habitat models: isolation-driven occurrence of north island robins in a fragmented landscape

Although the role of habitat fragmentation in species declines is well recognised, the effect of habitat quality on species distributions is often studied using presence–absence models that ignore metapopulation dynamics. We compared three approaches to model the presence–absence of North Island robins in 400 sites among 74 fragments of native forest in a 15,000-ha agricultural landscape in New Zealand. The first approach only considered local habitat characteristics, the second approach only considered metapopulation factors (patch size and isolation), and the third approach combined these two types of factors. The distribution of North Island robins was best predicted by patch isolation, as their probability of occurrence was negatively correlated with isolation from neighbouring patches and from the closest major forests, which probably acted as a source of immigrants. The inclusion of habitat factors gave only a slight increase in predictive power and indicated that robins were more likely to occur in areas with tall canopy, tall understory and low density of young trees. We modelled the effect of isolation using an index of functional patch connectivity based on dispersal behaviour of radio-tracked juveniles, and this functional index greatly improved the models in comparison to classical indices relying on Euclidean distances. This study highlights the need to incorporate functional indices of isolation in presence–absence models in fragmented landscapes, as species occurrence can otherwise be a misleading predictor of habitat quality and lead to wrong interpretations and management recommendations.     

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

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

Spatial dynamics of the knob-tailed gecko Nephrurus stellatus in a fragmented agricultural landscape

In fragmented landscapes, a species?? dispersal ability and response to habitat condition are key determinants of persistence. To understand the relative importance of dispersal and condition for survival of Nephrurus stellatus (Gekkonidae) in southern Australia, we surveyed 92 woodland remnants three times. This gecko favours early post-fire succession conditions so may be at risk of extinction in the long-unburnt agricultural landscape. Using N-mixture models, we compared the influence of four measures of isolation, patch area and two habitat variables on the abundance and occurrence of N. stellatus, while taking into account detection probability. Patch occupancy was high, despite the long-term absence of fire from most remnants. Distance to the nearest occupied site was the most informative measure of patch isolation, exhibiting a negative relationship with occupancy. Distance to a nearby conservation park had little influence, suggesting that mainland?Cisland metapopulation dynamics are not important. Abundance and occurrence were positively related to ?%-cover of spinifex (Triodia), indicating that niche-related factors may also contribute to spatial dynamics. Patterns of patch occupancy imply that N. stellatus has a sequence of spatial dynamics across an isolation gradient, with patchy populations and source-sink dynamics when patches are within 300?m, metapopulations at intermediate isolation, and declining populations when patches are separated by >1?C2?km. Considering the conservation needs of the community, habitat condition and connectivity may need to be improved before fire can be reintroduced to the landscape. We speculate that fire may interact with habitat degradation and isolation, increasing the risk of local extinctions.     

Relating tradable credits for biodiversity to sustainability criteria in a dynamic landscape

Tradable biodiversity credit systems provide flexible means to resolve conflicts between development and conservation land-use options for habitats occupied by threatened or endangered species. We describe an approach to incorporate the influence of habitat fragmentation into the conservation value of tradable credits. Habitat fragmentation decreases gene flow, increases rates of genetic drift and inbreeding, and increases probabilities of patch extinction. Importantly, tradable credit systems will change the level of fragmentation over time for small and/or declining populations. We apply landscape equivalency analysis (LEA), a generalizable, landscape-scale accounting system that assigns conservation value to habitat patches based on patch contributions to abundance and genetic variance at landscape scales. By evaluating habitat trades using two models that vary the relationship between dispersal behaviors and landscape patterns, we show that LEA provides a novel method for limiting access to habitat at the landscape-scale, recognizing that the appropriate amount of migration needed to supplement patch recruitment and to offset drift and inbreeding will vary as landscape pattern changes over time. We also found that decisions based on probabilities of persistence alone would ignore changes in migration, genetic drift, and patch extinction that result from habitat trades. The general principle of LEA is that habitat patches traded should make at least equivalent contributions to rates of recruitment and migration estimated at a landscape scale. Traditional approaches for assessing the “take” and “jeopardy” standards under the Endangered Species Act based on changes in abundance and probability of persistence may be inadequate to prevent trades that increase fragmentation.     

Interactions between plant life span, seed dispersal capacity and fecundity determine metapopulation viability in a dynamic landscape

Classical metapopulation models do not account for temporal changes in the suitability of habitat patches. In reality, however, the carrying capacity of most habitat types is not constant in time due to natural succession processes. In this study, we modeled plant metapopulation persistence in a successional landscape with disappearing and emerging habitat patches, based on a realistic dune slack landscape at the Belgian–French coast. We focused on the effects of the variation of different plant traits on metapopulation persistence in this changing landscape. Therefore, we used a stage based stochastic metapopulation model implemented in RAMAS/Metapop, simulating a large variation in plant traits but keeping landscape characteristics such as patch turnover rate and patch lifespan constant. The results confirm the conclusions of earlier modeling work that seed dispersal distance and seed emigration rate both have an important effect on metapopulation persistence. We also found that high population growth rate or high recruitment considerably decreased the extinction risk of the metapopulation. Additionally, a long plant life span had a strong positive effect on metapopulation persistence, irrespective of the plant's dispersal capacity and population growth rate. Plant species that invest in life span require less investment in offspring and dispersal capacity to avoid extinction, even in dynamic landscapes with deterministic changes in habitat quality. Moreover, metapopulations of long-lived plant species were found to be much less sensitive to high levels of environmental stochasticity than short-lived species.     

Effects of Habitat Suitability and Landscape Patterns on Tick (Acarina) Metapopulation Processes

Tick density and population dynamics are important factors in the ecological processes involved in pathogen circulation in a habitat. These characteristics of tick populations are closely linked to habitat suitability, which reflects the limiting ecological factors and landscape features affecting tick populations; however, little work has been done on the regional assessment of habitat suitability. In this study, a regional model for the distribution and abundance of the tick Ixodes ricinus in central Spain is developed. An occurrence and an abundance model were constructed; climate and vegetation variables were found to be the main predictors of both occurrence and density in a relatively homogeneous matrix of habitat patches, whereas topographical variables were found to have small contributions and were therefore discarded. The residuals of the abundance model showed good correlation with the isolation of each patch. The predictive power of the abundance model was greatly enhanced by inclusion of the traversability (a measure of the permeability of each patch to the propagules of the metapopulation) and recruitment (an index of the relative importance of each patch to the traffic through the entire habitat network). The removal from the landscape of the patches whose recruitment values were in the top 10% has a critical effect on tick density, an effect not observed when patches are removed at random. These results indicate that permanent tick populations can be sustained only in landscapes containing a minimum network of viable sites. Graph theory and measurements of patch isolation should prove to be important elements in the forecasting of tick abundance and the management of the features underlying the landscape ecology of tick populations and pathogen circulation in the field.     

The influence of habitat availability and landscape structure on the distribution of wood cricket (Nemobius sylvestris) on the Isle of Wight, UK

Little information is available regarding the landscape ecology of woodland invertebrate species with limited dispersal ability. An investigation was therefore conducted within woodland fragments in an agricultural landscape for the flightless wood cricket (Nemobius sylvestris) on the Isle of Wight, UK. The current pattern of distribution of the species, established during a field survey, was related to measures of habitat availability and habitat isolation/fragmentation. Results revealed that wood cricket populations were patchily distributed and mainly found in relatively large mature woodland fragments situated closely (<50 m) to another occupied site. Although the occurrence of wood cricket was related to fragment area, isolation, habitat availability and woodland age, a logistic regression model revealed that presence of the species was most accurately predicted by fragment isolation and area alone. These results highlight the vulnerability of relatively immobile woodland invertebrate species, such as wood cricket, to the impacts of habitat loss and fragmentation.     

Evaluation of patch isolation metrics in mosaic landscapes for specialist vs. generalist dispersers

We examined the effects of matrix structure and movement responses of organisms on the relationships between 7 patch isolation metrics and patch immigration. Our analysis was based on simulating movement behaviour of two generic disperser types (specialist and generalist) across mosaic landscapes containing three landcover types: habitat, hospitable matrix and inhospitable matrix. Movement, mortality and boundary crossing probabilities of simulated individuals were linked to the landcover and boundary types in the landscapes. The results indicated that area-based isolation metrics generally predict patch immigration more reliably than distance-based isolation metrics. Relationships between patch isolation metrics and patch immigration varied between the two generic disperser types and were affected by matrix composition or matrix fragmentation. Patch immigration was always affected by matrix composition but not by matrix fragmentation. Our results do not encourage the generic use of patch isolation metrics as a substitute for patch immigration, in particular in metapopulation models where generic use may result in wrong projections of the survival probability of metapopulations. However, our examination of the factors affecting the predictive potential of patch isolation metrics should facilitate interpretation and comparison of existing patch isolation studies. Future patch isolation studies should include information about landscape structure and the dispersal distance and dispersal behaviour of the organism of interest.This revised version was published online in May 2005 with corrections to the Cover Date.     

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.

     

Incidence of competitors and landscape structure as predictors of woodland-dependent birds

Globally, modification of landscapes for agriculture has had a strong influence on the distribution and abundance of biota. In particular, woodland-dependent birds are under threat across agricultural landscapes in Britain, North America and Australia, with their decline and extirpation attributed to the loss and fragmentation of habitat. Other native species have become over-abundant in response to anthropogenic landscape change and have strong interactive effects on avian assemblage structure. In eastern Australia, the hyper-aggressive noisy miner (Manorina melanocephala) often dominates woodlands in agricultural landscapes through interspecific competition, resulting in declines of species richness of woodland-dependent birds. We aimed to determine the relative influence and importance of interspecific competition, in situ habitat structure and landscape structure for woodland-dependent bird species at the landscape level. We recorded species-specific landscape incidence of woodland-dependent birds in 24 agricultural-woodland mosaics (25 km²) in southern Queensland, Australia. We selected extensively cleared landscapes (10–23 % woodland cover) where fragmentation effects are expected to be greatest. We applied generalised linear models and hierarchical partitioning to quantify the relative importance of the landscape-level incidence of the noisy miner, mistletoe abundance, shrub cover, woodland extent, woodland subdivision and land-use intensity for the incidence of 46 species of woodland birds at the landscape-scale. The landscape-level incidence of the noisy miner was the most important explanatory variable across the assemblage. Both in situ habitat structure and landscape structure were of secondary importance to interspecific aggression, although previous research suggests that the increasing incidence of the noisy miner in fragmented agricultural landscapes is itself a consequence of anthropogenic changes to landscape structure. Species’ responses to fragmentation varied from positive to negative, but complex habitat structure had a consistently positive effect, suggesting in situ restoration of degraded habitats could be a conservation priority. Landscape wide conservation of woodland-dependent bird populations in agricultural landscapes may be more effective if direct management of noisy miner populations is employed, given the strong negative influence of this species on the incidence of woodland-dependent birds among landscapes.     

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.     

Intermediate landscape disturbance maximizes metapopulation density

The viability of metapopulations in fragmented landscapes has become a central theme in conservation biology. Landscape fragmentation is increasingly recognized as a dynamical process: in many situations, the quality of local habitats must be expected to undergo continual changes. Here we assess the implications of such recurrent local disturbances for the equilibrium density of metapopulations. Using a spatially explicit lattice model in which the considered metapopulation as well as the underlying landscape pattern change dynamically, we show that equilibrium metapopulation density is maximized at intermediate frequencies of local landscape disturbance. On both sides around this maximum, the metapopulation may go extinct. We show how the position and shape of the intermediate viability maximum is responding to changes in the landscape’s overall habitat quality and the population’s propensity for local extinction. We interpret our findings in terms of a dual effect of intensified landscape disturbances, which on the one hand exterminate local populations and on the other hand enhance a metapopulation’s capacity for spreading between habitat clusters.     

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.     

Multi-scale landscape and seascape patterns associated with marbled murrelet nesting areas on the U.S. west coast

Habitat for wide-ranging species should be addressed at multiple scales to fully understand factors that limit populations. The marbled murrelet (Brachyramphus marmoratus), a threatened seabird, forages on the ocean and nests inland in large trees. We developed statistical relationships between murrelet use (occupancy and abundance) and habitat variables quantified across many spatial scales (statewide to local) and two time periods in California and southern Oregon, USA. We also addressed (1) if old-growth forest fragmentation was negatively associated with murrelet use, and (2) if some nesting areas are more important than others due to their proximity to high quality marine habitat. Most landscapes used for nesting were restricted to low elevation areas with frequent fog. Birds were most abundant in unfragmented old-growth forests located within a matrix of mature second-growth forest. Murrelets were less likely to occupy old-growth habitat if it was isolated (> 5 km) from other nesting murrelets. We found a time lag in response to fragmentation, where at least a few years were required before birds abandoned fragmented forests. Compared to landscapes with little tono murrelet use, landscapes with many murrelets were closer to the ocean's bays, river mouths, sandy shores, submarine canyons, and marine waters with consistently high primary productivity. Within local landscapes (≤ 800ha), inland factors limited bird abundance, but at the broadest landscape scale studied (3200 ha), proximity to marine habitat was most limiting. Management should focus on protecting or creating large, contiguous old-growth forest stands, especially in low-elevation areas near productive marine habitat. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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

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

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

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

Thresholds in landscape structure for three common deforestation patterns in the Brazilian Amazon

Although abrupt changes (i.e. thresholds) have been precisely defined in simulated landscapes, such changes in the structure of real landscapes are not well understood. We tested for threshold occurrence in three common deforestation patterns in the Brazilian Amazon: small properties regularly distributed along roads (fishbone), irregularly distributed small properties (independent settlements), and large properties. We analyzed differences between real deforestation patterns, and tested the capacity of simulated landscape with different aggregation degrees to predict threshold occurrence. Three 8×8 km sites (replicates) with more than 90% of forest in 1984 and less than 30% in 1998 were selected/simulated for each deforestation pattern. Thresholds were observed for fishbone and large property patterns, especially when considering the connectivity index, although threshold incidences were more frequently observed in simulated landscapes. The capacity of simulated landscapes to predict the exact threshold point in real landscapes was limited, even when considering highly aggregate simulations. However, the general trend in landscape structural changes was similar in real and simulated landscapes. Thresholds occurred at the beginning of the deforestation for mean patch size and at an intermediate stage, corresponding to the percolation threshold, for connectivity, isolation and fragmentation. Threshold behavior for connectivity index might suggest that the survival of strictly forest species will sharply decrease when the proportion of forest reach values <0.60, indicating that conservation efforts should be done to maintain forest cover above this limit. Significant differences observed among the real deforestation patterns, especially in patch isolation and number of fragments, can have significant consequences for conservation. The independent settlement pattern is, without a doubt, the least favorable of them, resulting in a higher level of fragmentation, whereas the large property and fishbone patterns may be less detrimental if connectivity among the remnant forest patches is preserved.     

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.