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.     

Migration rates of grassland plants along corridors in fragmented landscapes assessed with a cellular automation model

This study investigated the efficacy of linear landscape elements in fragmented landscapes as corridors for perennial grassland species with short-range seed dispersal. Corridors are assumed to be essential for the persistence of metapopulations in fragmented landscapes, but it is unclear to what extent linear landscape elements such as ditch banks and road verges can function as corridors for those species. The principal factors that determine the rate of migration through corridors include the width and habitat quality of patches within a corridor (expressed as the population growth rate λ) and the dispersal capacity of plants (expressed as the slope α of the relationship between seed number and log-distance). A cellular automation model was used to simulate the effects of the principal factors on the rate of migration. Simulations with different levels of the principal factors showed highly significant and positive main effects of dispersal capacity, habitat quality and width of corridors on the migration rate. Significant interactions existed between dispersal capacity × width and dispersal capacity × habitat quality (p<0.0001), indicating that the effects of width and habitat quality depended on the dispersal capacity. In narrow corridors most of the dispersed seeds were deposited outside the corridor, which significantly reduced migration rates, especially for species with long-range dispersal (α=−0.4). In wide corridors (up to 20 m), seed losses were much smaller and migration rates approximated those of continuous habitats. The contribution of the few long-range seeds to the rate of migration was significant when habitat quality was high (population growth rates up to 2.5). However, in all simulations migration rates were very low,i.e.<5 m/yr. It is concluded that linear landscape elements are not effective corridors in fragmented landscapes for plants with short-range seed dispersal, because migration rates are low (<5 m/yr), landscape elements vary in the percentage of high quality patches, and refugia and suitable habitat patches are frequently several kilometres apart, making a cohesive infrastructure of corridors for plants elusive. It is argued that the best way to conserve endangered plant species that encounter dispersal barriers is to harvest seeds from nearby source populations and introduce them as suitable habitats.     

The Viability of Metapopulations: Individual Dispersal Behaviour Matters

Metapopulation models are frequently used for analysing species–landscape interactions and their effect on structure and dynamic of populations in fragmented landscapes. They especially support a better understanding of the viability of metapopulations. In such models, the processes determining metapopulation viability are often modelled in a simple way. Animals’ dispersal between habitat fragments is mostly taken into account by using a simple dispersal function that assumes the underlying process of dispersal to be random movement. Species-specific dispersal behaviour such as a systematic search for habitat patches is likely to influence the viability of a metapopulation. Using a model for metapopulation viability analysis, we investigate whether such specific dispersal behaviour affects the predictions of ranking orders among alternative landscape configurations rated regarding their ability to carry viable metapopulations. To incorporate dispersal behaviour in the model, we use a submodel for the colonisation rates which allows different movement patterns to be considered (uncorrelated random walk, correlated random walk with various degrees of correlation, and loops). For each movement pattern, the landscape order is determined by comparing the resulting mean metapopulation lifetime T_m of different landscape configurations. Results show that landscape orders can change considerably between different movement patterns. We analyse whether and under what circumstances dispersal behaviour influences the ranking orders of landscapes. We find that the ‘competition between patches for migrants’ – i.e. the fact that dispersers immigrating into one patch are not longer available as colonisers for other patches – is an important factor driving the change in landscape ranks. The implications of our results for metapopulation modelling, planning and conservation are discussed.     

Plant strategies and agricultural landscapes: survival in spatially and temporally fragmented habitat

In agricultural landscapes many plant species are limited to the network of landscape elements that are not used for agricultural production. This habitat is fragmented in space and time due to anthropogenic, biotic and abiotic factors. Therefore, plant populations are spatially sub-divided and their persistence might be dependent on the spatial dynamics in the network of local populations. Dispersal characteristics and seed bank persistence are main determinants of colonization ability which in turn is a key determinant of metapopulation viability. We propose a conceptual model that relates plant population dynamics to habitat quality, configuration and dynamics. In this model, the habitat is arranged as a network of suitable and unsuitable patches,and the distribution of the patches is assumed to be dynamic in time. Based on dispersal and seed bank characteristics four plant strategies are distinguished:species having either long (> 100 m) or short (< 100m) distance dispersal and either a long (> 5 yr)or short (< 5 yr) term persistent seed bank. We expect that species with contrasting strategies have different survival probabilities in landscapes with contrasting habitat arrangement in space and time. We found few empirical studies for testing the hypotheses based on the model. Therefore the relation between landscapes and plant survival needs to be further explored,especially the quantitative aspects. We propose an iterative process of empirical and modelling research to determine this relation and to define management options for multifunctional farms in which biodiversity is one of the land use aims. This revised version was published online in July 2006 with corrections to the Cover Date.     

The elements of connectivity where corridor quality is variable

Small mammals in heterogeneous environments have been found to disperse along corridors connecting habitat patches. Corridors may have different survivability values depending on their size and the degree of cover they provide. This deterministic model tests the effects of varying corridor quality on the demographics of a metapopulation of Peromyscus leucopus. Two types of corridors are defined based on the probability of survival during a dispersal event. Results indicate that mortality during movement through corridors influences metapopulation demographics. We found that:

1. Any connection between two isolated patches is better than no connection at all in terms of persistence and population size at equilibrium.
2. Metapopulations with exclusively high quality corridors between patches have a larger population size at equilibrium than do those with one or more low quality corridors.
3. Increasing the number of high quality corridors between patches has a positive effect on the size of the metapopulation while increasing the number of low quality corridors has a negative effect.
4. The addition to a metapopulation of a patch connected by low quality corridors has a negative effect on the metapopulation size. This suggest the need for caution in planning corridors in a managed landscape.
5. There is no relationship between the number of corridors and the metapopulation size at equilibrium when the number of connected patches is held constant.
6. Geometrically isolated patches connected by low quality corridors are most vulnerable to local extinctions.

We conclude that corridor quality is an important element of connectivity. It contributes substantially to the effects of fragmentation and should be carefully considered by landscape planners.     

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.     

A Practical Map-Analysis Tool for Detecting Potential Dispersal Corridors

We describe the Pathway Analysis Through Habitat (PATH) tool, which can predict the location of potential corridors of animal movement between patches of habitat within any map. The algorithm works by launching virtual entities that we call `walkers' from each patch of habitat in the map, simulating their travel as they journey through land cover types in the intervening matrix, and finally arrive at a different habitat `island.' Each walker is imbued with a set of user-specified habitat preferences that make its walking behavior resemble a particular animal species. Because the tool operates in parallel on a supercomputer, large numbers of walkers can be efficiently simulated. The importance of each habitat patch as a source or a sink for a species is calculated, consistent with existing concepts in the metapopulation literature. The manipulation of a series of contrived artificial landscapes demonstrates that the location of potential dispersal corridors and relative source and sink importance among patches can be purposefully altered in expected ways. Finally, potential dispersal corridors are predicted among remnant woodlots within three actual landscape maps. The U.S. Government’s right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Movements of cactus bugs: Patch transfers,matrix resistance,and edge permeability

Individual movement is a key process affecting the distribution of animals in heterogeneous landscapes. For specialist species in patchy habitat, a central issue is how dispersal distances are related to landscape structure. We compared dispersal distances for cactus bugs (Chelinidea vittiger) on two naturally fragmented landscapes (≤ 4% suitable habitat) with different matrix structures (i.e., vegetation height of nonsuitable habitat between suitable patches). Using mark-release-recapture studies, we determined that most transfers between cactus patches occurred during the mating season. Dispersal distances were reduced by > 50% on the landscape that had reduced structural connectivity due to relatively high matrix structure and low patch density. An experiment with detailed movement pathways demonstrated that greater matrix structure decreased mean step lengths, reduced directionality, and thus decreased net displacement by > 60%. However, habitat edges between two matrix elements that differed substantially in resistance to movement were completely permeable. Therefore, the difference in distributions of dispersal distances between the two landscapes mainly reflected the average resistance of matrix habitat and not the level of matrix heterogeneity per se. Our study highlights the merits of combining estimates of dispersal distances with insights on mechanisms from detailed movement pathways, and emphasizes the difficulty of treating dispersal distances of species as fixed traits independent of landscape structure.     

Patterns of habitat occupancy, genetic variation and predicted movement of a flightless bush cricket, Pholidoptera griseoaptera, in an agricultural mosaic landscape

Habitat fragmentation has been generally regarded detrimental to the persistence of many species, especially those with limited dispersal abilities. Yet, when exactly habitat elements become functionally disconnected very much depends on the dispersal ability of a species in combination with the landscape’s composition in which it occurs. Surprisingly, for many small and ground-walking generalists knowledge at what spatial scale and to what extent landscape structure affects dispersal is very scarce. Because it is flightless, the bush cricket Pholidoptera griseoaptera may be regarded susceptible to fragmentation. We applied habitat occupancy surveys, population genetic analyses and movement modelling to investigate the performance of P. griseoaptera in an agricultural mosaic landscape with suitable habitat patches of varying size and isolation. Despite its presumed dispersal limitation we could show that P. griseoaptera occupied the majority of suitable habitats, including small and isolated patches, showed a very low and non-significant genetic differentiation (F _ST = 0.0072) and, in the model, managed to colonize around 73% of all suitable habitat patches within one generation under weak and strong landscape-effect scenarios. We conclude that P. griseoaptera possesses the behavioural attributes (frequent inter-patch dispersal) necessary to persist in this landscape characterized by a patchy distribution of habitat elements. Yet, sound recommendations to landscape planning and conservation require more research to determine whether this represents a general behaviour of the species or a behavioural adaptation to this particular landscape.     

Integrative approach for landscape-based graph connectivity analysis: a case study with the common frog (Rana temporaria) in human-dominated landscapes

Graph-based analysis is a promising approach for analyzing the functional and structural connectivity of landscapes. In human-shaped landscapes, species have become vulnerable to land degradation and connectivity loss between habitat patches. Movement across the landscape is a key process for species survival that needs to be further investigated for heterogeneous human-dominated landscapes. The common frog (Rana temporaria) was used as a case study to explore and provide a graph connectivity analysis framework that integrates habitat suitability and dispersal responses to landscape permeability. The main habitat patches influencing habitat availability and connectivity were highlighted by using the software Conefor Sensinode 2.2. One of the main advantages of the presented graph-theoretical approach is its ability to provide a large choice of variables to be used based on the study’s assumptions and knowledge about target species. Based on dispersal simulation modelling in potential suitable habitat corridors, three distinct patterns of nodes connections of differing importance were revealed. These patterns are locally influenced by anthropogenic barriers, landscape permeability, and habitat suitability. And they are affected by different suitability and availability gradients to maximize the best possible settlement by the common frog within a terrestrial habitat continuum. The study determined the key role of landscape-based approaches for identifying the “availability-suitability-connectivity” patterns from a local to regional approach to provide an operational tool for landscape planning.     

Joint effects of habitat configuration and temporal stochasticity on population dynamics

Habitat configuration and temporal stochasticity in the environment are recognized as important drivers of population structure, yet few studies have examined the combined influence of these factors. We developed a spatially explicit simulation model to investigate how stochasticity in survival and reproduction influenced population dynamics on landscapes that differed in habitat configuration. Landscapes ranged from completely contiguous to highly fragmented, and simulated populations varied in mean survival probability (0.2, 0.4, 0.8) and dispersal capacity (1, 3, or 5 cells). Overall, habitat configuration had a large effect on populations, accounting for >80% of the variation in population size when mean survival and dispersal capacity were held constant. Stochasticity in survival and reproduction were much less influential, accounting for <1–14% of the variation in population size, but exacerbated the negative effects of habitat fragmentation by increasing the number of local extinctions in isolated patches. Stochasticity interacted strongly with both mean survival probability and habitat configuration. For example, survival stochasticity reduced population size when survival probability was high and habitat was fragmented, but had little effect on population size under other conditions. Reproductive stochasticity reduced population size irrespective of mean survival and habitat configuration, but had the largest effect when survival probability was intermediate and habitat was well connected. Stochasticity also enhanced the variability of population size in most cases. Contrary to expectations, increasing dispersal capacity did not increase population persistence, because the probability of finding suitable habitat within the dispersal neighborhood declined more for the same level of dispersal capacity when fragmentation was high compared to when it was low. These findings suggest that greater environmental variability, as might arise due to climate change, is likely to compound population losses due to habitat fragmentation and may directly reduce population size if reproductive output is compromised. It may also increase variability in population size.     

Analyzing the effect of stepping stones on target patch colonisation in structured landscapes for Eurasian lynx

With habitat loss and fragmentation having become two of the major threats to the viability of species, the question of how to manage landscapes for species conservation has attracted much attention. In this context, the planning of stepping stones has been proposed to increase connectivity in fragmented landscapes. We present a simulation study with a neutral landscape approach to assess the effects of stepping stones on colonization success. To that end, we used a spatially explicit, calibrated population model of the European lynx (Lynx lynx) coupled with structured landscapes, in which we could control the landscape parameters of dispersal habitat coverage and contagion, as well as the number and size of stepping stones available for breeding. In general, we found that colonization success increased with increasing habitat coverage but decreased with increasing habitat contagion, while the introduction of stepping stones had significant effects in critical situations. Especially at low to medium dispersal habitat coverage and high disperser mortality, stepping stones had a positive effect on colonization success when they were large enough to produce new dispersers, but negative effects when they were small and located in a way that dispersers would be distracted from more suitable breeding habitat patches. The latter clearly constituted a shading effect and argues for a thorough consideration of the trade-offs related to stepping stone size and location when implementing stepping stones as a conservation measure, especially when the number of individuals of conservation concern is low.     

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.     

Using network theory to prioritize management in a desert bighorn sheep metapopulation

Connectivity models using empirically-derived landscape resistance maps can predict potential linkages among fragmented animal and plant populations. However, such models have rarely been used to guide systematic decision-making, such as identifying the most important habitat patches and dispersal corridors to protect or restore in order to maximize regional connectivity. Combining resistance models with network theory offers one means of prioritizing management for connectivity, and we applied this approach to a metapopulation of desert bighorn sheep (Ovis canadensis nelsoni) in the Mojave Desert of the southwestern United States. We used a genetic-based landscape resistance model to construct network models of genetic connectivity (potential for gene flow) and demographic connectivity (potential for colonization of empty habitat patches), which may differ because of sex-biased dispersal in bighorn sheep. We identified high-priority habitat patches and corridors and found that the type of connectivity and the network metric used to quantify connectivity had substantial effects on prioritization results, although some features ranked highly across all combinations. Rankings were also sensitive to our empirically-derived estimates of maximum effective dispersal distance, highlighting the importance of this often-ignored parameter. Patch-based analogs of our network metrics predicted both neutral and mitochondrial genetic diversity of 25 populations within the study area. This study demonstrates that network theory can enhance the utility of landscape resistance models as tools for conservation, but it is critical to consider the implications of sex-biased dispersal, the biological relevance of network metrics, and the uncertainty associated with dispersal range and behavior when using this approach.     

Effects of patch attributes, barriers, and distance between patches on the distribution of a rock-dwelling rodent (Lagidium viscacia)

We tested whether size of habitat patches and distance between patches are sufficient to predict the distribution of the mountain vizcacha Lagidium viscacia a large, rock-dwelling rodent of the Patagonian steppe Argentina, or whether information on other patch and landscape characteristics also is required. A logistic regression model including the distance between rock crevices and depth of crevices, distance between a patch and the nearest occupied patch, and whether or not there was a river separating it from the nearest occupied patch was a better predictor of patch occupancy by mountain vizcachas than was a model based only on patch size and distance between patches. Our results indicate that a simple metapopulation analysis based on size of habitat patches and distance between patches may not provide an accurate representation of regional population dynamics if patches vary in habitat quality independently of patch size and features in the matrix alter connectivity. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Testing assumptions of cost surface analysis—a tool for invasive species management

Applied ecology could benefit from new tools that identify potential movement pathways of invasive species, particularly where data are sparse. Cost surface analysis (CSA) estimates the permeability (friction) across a landscape and can be applied to dispersal modelling. Increasingly used in a diversity of applications, several fundamental assumptions that might influence the outputs of CSA (cost surfaces and least-cost pathways) have yet to be systematically examined. Thus, we explore two issues: the presumed relationship between habitat preferences and dispersal behaviour as well as the degree of landscape fragmentation through which an organism moves by modelling a total of 18 sensitivity and dispersal scenarios. We explored the effect of fragmentation by altering the friction values (generally assigned using expert opinion) associated with patch and linear features. We compared these sensitivity scenarios in two sites that differed in fragmentation. We also used eastern grey squirrels (Sciurus carolinensis) as an example invading species and compared diffusion models and two contrasting cost surface dispersal scenarios. The diffusion model underestimated spread because squirrels did not move randomly through the landscape. Despite contrasting assumptions regarding dispersal behaviour, the two cost surfaces were strikingly similar while the least-cost paths differed. Furthermore, while the cost surfaces were insensitive to changes in friction values for linear features, they were sensitive to assumptions made for patch features. Our results suggest that movement in fragmented landscapes may be more sensitive to assumptions regarding friction values than contiguous landscapes. Thus, the reliability of CSA may depend not only on the range of friction values used for patches but also the degree of contiguity in the landscape.     

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.     

Extinction risk of wood-living model species in forest landscapes as related to forest history and conservation strategy

Dead wood is a critical resource for biodiversity in boreal forests. We analysed the persistence of five model species inhabiting dead wood. By parameterising a metapopulation model (the incidence function model), the model species were all assigned characteristics that makes it likely that they have disappeared from some (20%) forest landscapes with a long history of forest management. In the metapopulation model, a forest stand (5 ha) was regarded as a habitat patch. The amount of habitat in each patch was obtained from models of dead wood dynamics of Norway spruce in central Sweden. Dead wood generated by altered management over the entire landscape was found to be less efficient in reducing extinction risks in comparison to the same amount of dead wood generated by protecting reserves. Because generation of dead wood by altered management is often less expensive than setting aside reserves, it is difficult to determine which conservation measure is most cost-efficient. In a landscape subjected to forestry for the first time, it was better to preserve a few large reserves than many small ones. However, in a managed, highly fragmented forest landscape it was better to set aside many small reserves. The reason for this was that small plots with high habitat quality could be selected, while large reserves originally contained habitats both of high and low quality, and the rate of habitat quality increase was low. A strategy for biodiversity conservation in a managed forest landscape should include information about the history of the landscape, the current amount and spatial distribution of forest habitats, and the potential for rapid restoration of forest habitats, both on managed and unmanaged forest land.     

The effects of the number,size and isolation of patches along a gradient of native vegetation cover: how can we increment habitat availability?

Habitat availability—or how much habitat species can reach at the landscape scale—depends primarily on the percentage of native cover. However, attributes of landscape configuration such as the number, size and isolation of habitat patches may have complementary effects on habitat availability, with implications for the management of landscapes. Here, we determined whether, and at which percentages of native cover, the number, size and isolation of patches contribute for habitat availability. We quantified habitat availability in 325 landscapes spread across the state of Rio de Janeiro, in the Atlantic Forest hotspot, with either high (>50 %), intermediate (50–30 %), low (30–10 %) or very low (<10 %) percentage of native cover, and for six hypothetical species differing in inter-patch dispersal ability. Above 50 % of native cover, the percentage of cover per se was the only determinant of habitat availability, but below 50 % the attributes of landscape configuration also contributed for habitat availability. The number of patches had a negative effect on habitat availability in landscapes with 50–10 % of native cover, whereas patch size had a positive effect in landscapes with <10 % of native cover. The different species generally responded to the same set of landscape attributes, although to different extents, potentially facilitating decision making for conservation. In landscapes with >50 % of native cover, conservation actions are probably sufficient to guarantee habitat availability, whereas in the remaining landscapes additional restoration efforts are needed, especially to reconnect and/or enlarge remaining habitat patches.