Landscape heterogeneity and fire behavior: scale-dependent feedback between fire and grazing processes

Fire and grazing are ecological processes that frequently interact to modify landscape patterns of vegetation. There is empirical and theoretical evidence that response of herbivores to heterogeneity is scale-dependent however the relationship between fire and scale of heterogeneity is not well defined. We examined the relationship between fire behavior and spatial scale (i.e., patch grain) of fuel heterogeneity. We created four heterogeneous landscapes modeled after those created by a fire–grazing interaction that differed in grain size of fuel patches. Fire spread was simulated through each model landscape from 80 independent, randomly located ignition points. Burn area, burn shape complexity and the proportion of area burnt by different fire types (headfire, backfire and flankfire) were all affected by the grain of fuel patch. The area fires burned in heterogeneous landscapes interacted with the fuel load present in the patch where ignition occurred. Burn complexity was greater in landscapes with small patch grain than in landscapes with large patch grain. The proportion of each fire type (backfire, flankfire and headfire) was similar among all landscapes regardless of patch grain but the variance of burned area within each of the three fire types differed among treatments of patch grain. Our landscape fire simulation supports the supposition that feedbacks between landscape patterns and ecological processes are scale-dependent, in this case spatial scale of fuel loading altering fire spread through the landscape.     

Spatial variation in forage nutrient concentrations and the distribution of Serengeti grazing ungulates

Resident grazing ungulates in the Serengeti National Park, Tanzania, are conspicuously patchy in their distribution among regions of the Park. Linear programming models that maximize nitrogen (N) consumption by foraging ungulates in Serengeti regions having high and low resident animal densities were compared using forage ingestion rate and twelve nutritional requirements as simultaneously imposed constraints on forage choice. Model results indicate that (1) growing season N or crude protein is not limiting in either region although greater N ingestion is possible within the eastern corridor under other nutritional constraints, (2) grazing ungulates in the eastern corridor region occur in greater density and are capable of balancing dietary requirements solely from forage while simultaneously consuming more protein than ungulates in the northeast region, and (3) rarer landscape elements are most capable of providing ungulate dietary requirements in both the northeast and eastern corridor. These results provide a nutritional basis to understand patchy spatial distributions of grazers within Serengeti regions and landscapes, and provide a partial test of the hypothesis that large generalist herbivores should graze rare forages more frequently. The ability of uncommon landscape elements to support ungulate grazing over the growing season is supported by previous ecosystem studies that demonstrate the capability of grass forages for compensatory growth and the ability of grazing to stimulate rapid nutrient recycling.     

Small-scale patch structure in North American and South African grasslands responds differently to fire and grazing

Fire and grazing significantly impact small-scale patch structure and dynamics in savanna grasslands. We assessed small-scale grass-forb associations in long-term fire and grazing experiments in North America (NA) and Southern Africa (SA). Transects of 128 0.25 m² contiguous quadrats were sampled in areas with different combinations of grazing (ungrazed, single grazer, or multiple grazers) and fire frequency (unburned or annually burned). We predicted that (1) the patch structure of each of the dominant grasses in NA and SA would respond similarly to fire and grazing, and (2) that forb richness would be correlated to grass patch structure. Semi-variance analysis was used to determine patch structure of dominant grasses and forb cover. Community structure responded similarly in NA and SA to fire, grazing, and fire-grazing interactions. Species richness, diversity, and community heterogeneity were significantly higher in unburned-grazed sites. Grazing significantly increased forb cover and decreased cover of the dominant grasses, and the effects of fire on community structure depended on the grazing regime. Contrary to our prediction, we found that small-scale patch structure of the dominant grass species in NA and SA responded differently to grazing and fire. We found strong grass patch structure in unburned-ungrazed grasslands in both sites; however, grazing and fire reduced patch structure in NA but not SA, and in no instance did grass patch structure influence forb community structure. We conclude that fire and grazing have larger impacts on small-scale patch structure in NA than they do in SA even though overall community structure responded similarly on both continents.     

Modelling the effects of landscape pattern and grazing regimes on the persistence of plant species with high conservation value in grasslands in south-eastern Sweden

Semi-natural grasslands in Sweden are threatened by land-use change and lack of management with attendant risk to their biodiversity. We present a model to explore the effects of grazing frequency and intensity on plant species persistence, and the relative effects of grassland size and pattern. We used a landscape modelling platform, LAMOS (LAndscape MOdelling Shell), to design a landscape model of vegetation dynamics incorporating the effects of local succession, dispersal and grazing disturbance. Five plant functional groups (PFG), representing various combinations of persistence and dispersal character, light requirements and disturbance responses, were defined to model species dynamics. Based on old cadastral maps three different landscapes were designed representing specific time-layers, i.e., a historical (17th to 18th century), a pre-modern (1940s) and a present-day landscape. Simulations showed that a threshold was crossed when grasslands decreased in area to about 10–30% of the modelled area, and as a consequence the biomass of grassland-specific PFGs was strongly reduced. These competition sensitive groups did not persist in the model even with intense grazing in the present-day landscape, where grasslands occupy 11% of the total area. However, all grassland species would have been able to persist in the historical landscape, where grasslands occupied 59% of the total area, even without grazing. Our results suggest that continuous but low-intensity grazing is more positive for grassland PFGs than discontinuous but highly intensive grazing. This effect was particularly strong when the frequency and/or intensity of grazing dropped below a threshold of 20%. Simulations using three landscape maps designed to explore effects of further fragmentation and habitat loss showed that the spatial pattern of remaining grasslands is important for the persistence of grassland-specific PFG. The model presented here is an advance towards more realistic grazing models to explore the effects of prescribed grazing and landscape fragmentation on the persistence species or plant functional groups.This revised version was published online in May 2005 with corrections to the Cover Date.     

Surface metrics: an alternative to patch metrics for the quantification of landscape structure

Modern landscape ecology is based on the patch mosaic paradigm, in which landscapes are conceptualized and analyzed as mosaics of discrete patches. While this model has been widely successful, there are many situations where it is more meaningful to model landscape structure based on continuous rather than discrete spatial heterogeneity. The growing field of surface metrology offers a variety of surface metrics for quantifying landscape gradients, yet these metrics are largely unknown and/or unused by landscape ecologists. In this paper, we describe a suite of surface metrics with potential for landscape ecological application. We assessed the redundancy among metrics and sought to find groups of similarly behaved metrics by examining metric performance across 264 sample landscapes in western Turkey. For comparative purposes and to evaluate the robustness of the observed patterns, we examined 16 different patch mosaic models and 18 different landscape gradient models of landscape structure. Surface metrics were highly redundant, but less so than patch metrics, and consistently aggregated into four cohesive clusters of similarly behaved metrics representing surface roughness, shape of the surface height distribution, and angular and radial surface texture. While the surface roughness metrics have strong analogs among the patch metrics, the other surface components are largely unique to landscape gradients. We contend that the surface properties we identified are nearly universal and have potential to offer new insights into landscape pattern–process relationships. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Interaction of food resources and landscape structure in determining the probability of patch use by carnivores in fragmented landscapes

Studies on the distribution of mammalian carnivores in fragmented landscapes have focused mainly on structural aspects such as patch and landscape features; similarly, habitat connectivity is usually associated with landscape structure. The influence of food resources on carnivore patch use and the important effect on habitat connectivity have been overlooked. The aim of this study is to evaluate the relative importance of food resources on patch use patterns and to test if food availability can overcome structural constraints on patch use. We carried out a patch-use survey of two carnivores: the beech marten (Martes foina) and the badger (Meles meles) in a sample of 39 woodland patches in a fragmented landscape in central Italy. We used the logistic model to investigate the relative effects on carnivore distribution of patch, patch neighbourhood and landscape scale variables as well as the relative abundance of food resources. Our results show how carnivore movements in fragmented landscapes are determined not only by patch/landscape structure but also by the relative abundance of food resources. The important take-home message of our research is that, within certain structural limits (e.g. within certain limits of patch isolation), by modifying the relative amount of resources and their distribution, it is possible to increase suitability in smaller/relatively isolated patches. Conversely, however, there are certain thresholds above which an increase in resources will not achieve high probability of presence. Our findings have important and generalizable consequences for highly fragmented landscapes in areas where it may not be possible to increase patch sizes and/or reduce isolation so, for instance, forest regimes that will increase resource availability could be implemented. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Scaling properties in landscape patterns: New Zealand experience

In this paper we present a case study of spatial structure in landscape patterns for the North and South Islands of New Zealand. The aim was to characterise quantitatively landscape heterogeneity and investigate its possible scaling properties. The study examines spatial heterogeneity, in particular patchiness, at a range of spatial scales, to help build understanding on the effects of landscape heterogeneity on water movement in particular, and landscape ecology in general.We used spatial information on various landscape properties (soils, hydrogeology, vegetation, topography) generated from the New Zealand Land Resource Inventory. To analyse this data set we applied various methods of fractal analyses following the hypothesis that patchiness in selected landscape properties demonstrates fractal scaling behaviour at two structural levels: (1) individual patches; and (2) mosaics (sets) of patches.Individual patches revealed scaling behaviour for both patch shape and boundary. We found self-affinity in patch shape with Hurst exponent H from 0.75 to 0.95. We also showed that patch boundaries in most cases were self-similar and in a few cases of large patches were self-affine. The degree of self-affinity was lower for finer patches. Similarly, when patch scale decreases the orientation of patches tends to be uniformly distributed, though patch orientation on average is clearly correlated with broad scale geological structures. These results reflect a tendency to isotropic behaviour of individual patches from broad to finer scales. Mosaics of patches also revealed fractal scaling in the total patch boundaries, patch centers of mass, and in patch area distribution. All these reflect a special organisation in patchiness represented in fractal patch clustering. General relationships which interconnect fractal scaling exponents were derived and tested. These relationships show how scaling properties of individual patches affect those for mosaics of patches and vice-versa. To explain similarity in scaling behaviour in patchiness of different types we suggest that the Self-Organised Criticality concept should be used. Also, potential applications of our results in landscape ecology are discussed, especially in relation to improved neutral landscape models.     

Bush cover and range condition assessments in relation to landscape and grazing in southern Ethiopia

Progressive growth of bush cover in dry savannahs is responsible for declines in range conditions. In southern Ethiopia, the Booran pastoralists assisted our understanding of spatial patterns of bush cover and range conditions in 54 landscape patch types grouped into six landscape units within an area of 30000 km². The size of landscape patches sampled was 625 m². We assessed the relationships between bush cover, grass cover and bare soil and grazing pressure and soil erosion and changes in range condition. Externally, political conflicts and internally, break down of land use, and official bans on the use of fire promoted bush cover and the decline in range conditions. Bush cover was negatively correlated with grass cover, and positively correlated with bare soil. Grass cover was negatively correlated with bare soil and grazing pressure in most landscape patch types. Grazing pressure was not significantly correlated with bush cover or bare soil, while soil erosion was directly related to bare soil. Soil erosion was absent in 64% of the landscape patch types, and seemingly not a threat to the rangelands. The relationship between bush cover, grass cover, bare soil and soil erosion is complex and related to climate, landscape geology, and patterns of land use. Main threats to range conditions are bush climax, loss of grass cover and unpalatable forbs. Currently, >70% of the landscape patch types are in poor to fair range conditions. Decline in range conditions, unless reversed, will jeopardise the pastoral production system in southern Ethiopia.     

Metapopulation dynamics and a landscape approach to conservation of lowland water voles (<Emphasis Type="Italic">Arvicola amphibius</Emphasis>)

Effective conservation management for species that function as metapopulations requires an understanding of population dynamics at the landscape scale. The water vole, Arvicola amphibius, is one such species. Water voles have recently undergone a significant decline in the UK, as a result of habitat loss and predation from the introduced American mink, Neovison vison. Large reed bed and grazing marsh sites can provide refuge habitats for water voles from mink predation, in which case populations within these sites could sustain metapopulations in the surrounding landscape where conditions are less favourable. We carried out a study using a stochastic patch occupancy model to determine the long term viability of water vole metapopulations in the wider landscape around a series of extensive reed bed and grazing marsh sites designated as National Key Sites for water voles. The results of our model simulations show that a large protected core site, or mainland, is essential in maintaining the long term viability of these systems. Our results also show how these metapopulations could be enhanced by increasing patch numbers through habitat creation and/or restoration and suggest what the minimum effective size of created or restored patches should be. The study shows how population modelling can provide insight into some effective practical ways of enhancing the viability of water vole metapopulations at the landscape scale. Furthermore it demonstrates that extensive wetlands are an appropriate focus for water vole conservation measures.     

Landscape context matters: local habitat and landscape effects on the abundance and patch occupancy of collared lizards in managed grasslands

The distribution and abundance of a species may be simultaneously influenced by both local-scale habitat features and the broader patch and landscape contexts in which these populations occur. Different factors may influence patch occupancy (presence–absence) versus local abundance (number of individuals within patches), and at different scales, and thus ideally both occupancy and abundance should be investigated, especially in studies that seek to understand the consequences of land management on species persistence. Our study evaluated the relative influences of variables associated with the local habitat patch, hillside (patch context), and landscape context on patch occupancy and abundance of the collared lizard (Crotaphytus collaris) within tallgrass prairie managed under different fire and grazing regimes in the northern Flint Hills of Kansas, USA. Using a multi-model information-theoretic approach that accounted for detection bias, we found that collared lizard abundance and occupancy was influenced by factors measured at both the local habitat and landscape scales. At a local scale, collared lizard abundance was greatest on large rock ledges that had lots of crevices, high vegetation complexity, and were located higher up on the hillslope. At the landscape scale, collared lizard abundance and occupancy were both higher in watersheds that were burned frequently (1–2 year intervals). Interestingly, grazing only had a significant effect on occupancy and abundance within less frequently burned (4-year burn interval) watersheds. Our results suggest that, in addition to the obvious habitat needs of this species (availability of suitable rock habitat), land-management practices have the potential to influence collared lizard presence and abundance in the grasslands of the Flint Hills. Thus, mapping the availability of suitable habitat is unlikely to be sufficient for evaluating species distributions and persistence in such cases without consideration of landscape management and disturbance history.     

Characterizing the importance of habitat patches and corridors in maintaining the landscape connectivity of a Pholidoptera transsylvanica (Orthoptera) metapopulation

Since the fragmentation of natural habitats is one of the most serious problems for many endangered species, it is highly interesting to study the properties of fragmented landscapes. As a basic property, landscape connectivity and its effects on various ecological processes are frequently in focus. First, we discuss the relevance of some graph properties in quantifying connectivity. Then, we propose a method how to quantify the relative importance of habitat patches and corridors in maintaining landscape connectivity. Our combined index explicitly considers pure topological properties and topographical measures, like the quality of both patches (local population size) and corridors (permeability). Finally, for illustration, we analyze the landscape graph of the endangered, brachypterous bush-cricket Pholidoptera transsylvanica. The landscape contains 11 patches and 13 corridors and is situated on the Aggtelek Karst, NE-Hungary. We characterize the importance of each node and link of the graph by local and global network indices. We show how different measures of connectivity may suggest different conservation preferences. We conclude, accordingly to our present index, by identifying one specific habitat patch and one specific corridor being in the most critical positions in maintaining connectivity.This revised version was published online in May 2005 with corrections to the Cover Date.     

Estimates of dryland degradation in Argentina with Fourier signatures from low-altitude monochromatic images with high spatial resolution

Most world drylands are used as graziny lands and undergo degradation of their vegetation cover. The plant cover is typically structured in patchy arrangements, inducing fertility islands critical to maintenance of ecosystem properties. The characteristics of patch structure (size of patches, connectivity-continuity of patch units, etc.) are indicators of the degree of dryland deterioration. We characterized changes in patch structure induced by sheep grazing at a landscape scale using monochromatic low-altitude imagery digitized to a spatial resolution of about 1 m with standard techniques of harmonic analysis applied to develop Fourier signatures. The signatures developed on image line transects were tested with ground samples and mathematical models of plant cover in several dryland fields where spatial deterioration gradients existed. The sensitivity and errors associated to long-wave noise introduced by the geometry of the camera-field-sun spatial arrangement and to high frequency noise introduced by the digitizing process were evaluated by applying suitable filters in the frequency domain. Fourier signatures developed on monochromatic low-altitude imagery proved to be indicative of changes in the patching arrangements of plant cover. We concluded that adequately filtered, high spatial resolution monochromatic images can be used to evaluate the degree of deterioration of dryland landscapes through the computation of selected Fourier signatures in their frequency domain. At comparable cost, aerial photography allows inspecting the landscape at higher spatial resolutions than those attainable with satellite imagery. Also, aerial photos of many areas are available for earlier dates than images from remote sensors, which would allow better inspection of long-term ecosystem changes.This revised version was published online in May 2005 with corrections to the Cover Date.     

The effect of fire spatial scale on <Emphasis Type="Italic">Bison</Emphasis> grazing intensity

To determine whether fire spatial and temporal scales affect foraging behavior and grazing intensity by Bison (Bison bison), we burned three different patch sizes (225, 900, and 3600 m²) across an otherwise homogeneous grassland landscape. We then monitored grazing intensity for the succeeding 14 months. During the first 5 months after the burn (August–January), the Bison grazing intensity pattern was affected by whether a plot was burned and only marginally affected by plot size. During the next 5 months (January–June), grazing intensity was unaffected by plot size, but was greatest in the unburned 225 and 3600-m² plots. The final 4 months (June–October), grazing intensity was unaffected by treatments other than being higher in the unburned 3600-m² plots. By the final sampling date, biomass was significantly greater in the burned plots and grazing intensity appeared to be responding to the amount of biomass present and the total amount of N present. The pattern displayed within the first 5 months after the burn is congruent with the expectations of optimal foraging theory with overmatching in the smallest plot size of 225 m² (BioScience 37 (1987) 789–799). The next two sampling periods displayed a matching aggregate response relative to biomass availability (Oecologia 100 (1999) 107–117) and total nitrogen mass (g m⁻²). The temporal shift that we found in Bison response to burn patch size is, to our knowledge, the first such examination of both spatial and temporal responses by Bison to landscape heterogeneity. We now have quantitative evidence of how native herbivores can alter their foraging responses to changes in landscape structure over time.     

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.     

Scale-dependent importance of environment, land use and landscape structure for species richness and composition of SE Norwegian modern agricultural landscapes

Knowledge of variation in vascular plant species richness and species composition in modern agricultural landscapes is important for appropriate biodiversity management. From species lists for 2201 land-type patches in 16 1-km² plots five data sets differing in sampling-unit size from patch to plot were prepared. Variation in each data set was partitioned into seven sources: patch geometry, patch type, geographic location, plot affiliation, habitat diversity, ecological factors, and land-use intensity. Patch species richness was highly predictable (75% of variance explained) by patch area, within-patch heterogeneity and patch type. Plot species richness was, however, not predictable by any explanatory variable, most likely because all studied landscapes contained all main patch types – ploughed land, woodland, grassland and other open land – and hence had a large core of common species. Patch species composition was explained by variation along major environmental complex gradients but appeared nested to lower degrees in modern than in traditional agricultural landscapes because species-poor parts of the landscape do not contain well-defined subsets of the species pool of species-rich parts. Variation in species composition was scale dependent because the relative importance of specific complex gradients changed with increasing sampling-unit size, and because the amount of randomness in data sets decreased with increasing sampling-unit size. Our results indicate that broad landscape structural changes will have consequences for landscape-scale species richness that are hard or impossible to predict by simple surrogate variables.     

When relationships estimated in the past cannot be used to predict the future: using mechanistic models to predict landscape ecological dynamics in a changing world

Researchers and natural resource managers need predictions of how multiple global changes (e.g., climate change, rising levels of air pollutants, exotic invasions) will affect landscape composition and ecosystem function. Ecological predictive models used for this purpose are constructed using either a mechanistic (process-based) or a phenomenological (empirical) approach, or combination. Given the accelerating pace of global changes, it is becoming increasingly difficult to trust future projections made by phenomenological models estimated under past conditions. Using forest landscape models as an example, I review current modeling approaches and propose principles for developing the next generation of landscape models. First, modelers should increase the use of mechanistic components based on appropriately scaled “first principles” even though such an approach is not without cost and limitations. Second, the interaction of processes within a model should be designed to minimize a priori constraints on process interactions and mimic how interactions play out in real life. Third, when a model is expected to make accurate projections of future system states it must include all of the major ecological processes that structure the system. A completely mechanistic approach to the molecular level is not tractable or desirable at landscape scales. I submit that the best solution is to blend mechanistic and phenomenological approaches in a way that maximizes the use of mechanisms where novel driver conditions are expected while keeping the model tractable. There may be other ways. I challenge landscape ecosystem modelers to seek new ways to make their models more robust to the multiple global changes occurring today.     

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.     

Quantifying habitat specificity to assess the contribution of a patch to species richness at a landscape scale

Assessing and predicting the species richness of a complex landscape remains a problem because there is no simple scaling function of species richness in a heterogeneous environment. Furthermore, the potential value of an area for biodiversity conservation may depend on which, rather than how many, species the area contains. This paper shows how we can objectively evaluate the contribution of an area, e.g., a habitat patch, to larger-scale plant species richness, e.g., a landscape composed of patches of several habitat types, and how we can test hypotheses that attempt to explain this contribution. We quantified the concept of habitat specificity to assess the proportion of each observed plant population that is concentrated within a given spatial element. A case study of a biodiversity-monitoring program in the Swiss Canton of Aargau showed that the relative contribution of the three main types of land use to the overall species richness differed strongly between higher taxa (vascular plants and molluscs). However, the type of data, i.e., presence-absence or abundance, was not important. Resampling of the plant data suggested that stratification provided an unbiased estimate of relative specificity, whereas unstratified sampling caused bias even for large samples. In a second case study of vascular plants in an agricultural landscape in central Switzerland, we tested whether the type, size or shape of a landscape element can predict its contribution to the species richness of the landscape. Habitat types that were less frequently disturbed contributed more per m² to landscape species richness than more frequently disturbed ones. Contrary to expectation, patch size was negatively correlated to specificity per m² for arable fields, whereas patch shape appeared to be unrelated to the specificity per m² both for arable fields and for meadows. The specificity approach provides a solution to the problem of scaling species richness and is ideally suited for testing hypotheses on the effect of landscape structure on landscape species richness. Specificity scores can easily be combined with measures of other aspects of rarity to assess the contribution of a spatial element to conservation goals formulated at regional, national or global level.     

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.