Long-term decline and local extinction of Clethrionomys rufocanus in boreal Sweden

Over the past three decades in boreal Sweden, there has been a long-term decline of cyclic sympatric voles, leading to local extinctions of the most affected species, the grey-sided vole (Clethrionomys rufocanus). We monitored this decline by snap-trapping on 58 permanent plots spread over 100 km² in spring and fall from fall 1971–2003. The reason for the decline is largely unknown, although a common major factor is likely to be involved in the decline of C. rufocanus and of the coexisting voles. However, here we deal with the reasonability of one complementary hypothesis, the habitat fragmentation hypothesis, which assumes that part of the decline of C. rufocanus is caused by habitat (forest) destruction. There was considerable local variation in the decline among the 58 1-ha sampling plots, with respect to both density and timing of the decline; however, all declines ended up with local extinction almost without exception. Local declines were not associated with habitat destruction by clear-cutting within sampling-plots, as declines started about equally often before as after clear-cutting, which suggested that habitat destruction outside sampling plots could be involved. In a multiple regression analysis, local habitat preference (LHP; expressed as a ratio of observed to expected number of voles trapped per habitat) together with two habitat variables in the surrounding (2.5×2.5 km²) landscape matrix explained 56% of the variation among local cumulated densities of C. rufocanus and hence of local time-series. LHP was positively correlated and explained 31% of the variation, while connectivity among clear-cuts was negatively correlated and proximity among xeric-mesic mires was positively correlated and explained additional 16% and 9%, respectively. Even if the overall decline cannot be connected to local clear-cutting on sampling-plots, clear-cutting and hence habitat fragmentation/destruction in the surrounding landscapes potentially influenced grey-sided vole numbers negatively.     

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

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

Landscape structure and the long-term decline of cyclic grey-sided voles in Fennoscandia

Changes in forest landscape structure have been suggested as a likely contributing factor behind the long-term decline in the numbers of cyclic grey-sided voles (Clethrionomys rufocanus) in northern Fennoscandian lowland regions in contrast to mountain regions due to the absence of forest management in the mountains. This study, for the first time, formally explored landscape structure in 29 lowland (LF) and 14 mountain forest (MF) landscapes (each 2.5 × 2.5 km) in northern Sweden, and related the results to the cumulated spring trapping index of the grey-sided vole in 2002–2006. The grey-sided vole showed striking contrasts in dynamics close in space and time. The MF landscapes were characterized by larger patches and less fragmentation of preferred forest types. The grey-sided vole was trapped in all of 14 analyzed MF landscapes but only in three out of 29 of the LF landscapes. MF and LF landscapes with grey-sided vole occurrence were characterized by similar focal forest patch size (mean 357 ha, minimum 82 ha and mean 360 ha, minimum 79 ha, respectively). In contrast, these MF compared to the LF landscapes were characterized by larger patches of preferred forest types and less fragmented preferred forest types and by a lower proportion of clear-cut areas. The present results suggest that landscape structure is important for the abundance of grey-sided voles in both regions. However, in the mountains the change from more or less seasonal dynamics to high-amplitude cycles between the mid 1990s and 2000s cannot be explained by changes in landscape structure.     

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.     

Effects of habitat fragmentation on meadow vole (Microtus pennsylvanicus) population dynamics in experiment landscape patches

This study examined the effects of habitat fragmentation on meadow vole (Microtus pennsylvanicus) population dynamics in experimental landscape patches. The study was conducted from May–November 1993 at the Miami University Ecology Research Center. Eight 0.1-ha small mammal enclosures were used. Four enclosures contained a 160 m² nonfragmented patch and four enclosures contained four 40 m² fragmented patches. Thus, each treatment was replicated 4 times in a systematic research design. The patches in both treatments contained high-quality habitat surrounded by low-quality matrix. Six pairs of adult meadow voles were released in each enclosure on 27 May 1993. Populations were monitored by live-trapping and radio-telemetry methods. Significantly greater densities of female voles were found during October in the fragmented treatment compared to the nonfragmented treatment. Also, significantly more females than males were found in the fragmented treatment compared to the nonfragmented treatment for the total study period. Significantly more subadult and juvenile males were found in the matrix versus the patch of the nonfragmented treatment compared to the fragmented treatment. Males in the fragmented treatment had significantly greater mean home range size than males or females in the nonfragmented treatment. There appears to exist a relationship between patch fragmentation and the social structure of meadow vole populations; this relationship appears to function as a population regulatory mechanism.     

The Effects of Land-use and Landscape Structure on Barn Owl (<Emphasis Type="Italic">Tyto alba</Emphasis>) Breeding Success in Southern England,U.K.

To aid effective conservation and management there is a need to understand the effect of landscape on species ecology. The aim of this research was to assess the effect of landscape parameters on breeding success of barn owls throughout the Rother and Arun River catchments, Sussex, UK. We used a Geographic Information System to describe the habitat mosaic and landscape structure within an estimated home range area of 3 km² around 85 artificial nest box sites. Results showed that land cover was less heterogeneous at successful sites, with home ranges dominated by a few habitat types of regular patch shapes. Unsuccessful nesting sites had significantly more improved grassland, suburban land and wetlands than successful sites. Cluster analysis and Principle Components Analysis was used to assess the similarity of the habitat mosaic within these areas and pellet analysis was undertaken to assess barn owl diet and prey availability. Ten prey species were recovered from pellets, field vole (Microtus agrestis), common shrews (Sorex araneus) and house mice (Mus musculus) making up nearly 90% of recoveries. However box sites varied in relative proportions of small mammal, and hence prey availability. Results indicated that land use and landscape structure can affect breeding success in barn owls. Higher levels of poor quality small mammal habitat were associated with unsuccessful sites. However, at a landscape scale, the habitat mosaic across the study area lacked variation, limiting analysis and clear correlations between habitat type and positive breeding success, suggesting that a finer scale was needed in future studies utilising this approach.     

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.     

Influence of patch and landscape characteristics on the distribution of the subterranean rodent <Emphasis Type="Italic">Ctenomys porteousi</Emphasis>

The understanding and prediction of the responses of animal populations to habitat fragmentation is a central issue in applied ecology. The identification of habitat variables associated to patch occupancy is particularly important when habitat quality is affected by human activities. Here, we analyze the influence of patch and landscape characteristics on patch occupancy by the subterranean herbivorous rodent Ctenomys porteousi. Patch occupancy was monitored in a network of 63 habitat patches identified by satellite imagery analysis which extends along almost the whole distributional range for C. porteousi. Suitable habitat for the occurrence of C. porteousi is highly fragmented and represents <10% of the total area in its distributional range. The distribution of C. porteousi in the patch network is affected not only by characteristics of the habitat patches, but also by those of the surrounding landscape matrix. Significant differences between occupied and empty patches were found in several environmental variables. Overall, occupied patches were larger, less vegetated, more connected, and had larger neighbor patches than empty patches. A stepwise procedure on a generalized linear model selected four habitat variables that explain patch occupancy in C. porteousi; it included the effects of habitat quality in the matrix surrounding the patch, average vegetation cover in the patch, minimum vegetation cover in the matrix surrounding the patch, and the area of the nearest neighbor patch. These results indicate that patch occupancy in C. porteousi is strongly influenced by the availability and quality of habitat both in the patch and in the surrounding landscape matrix.     

Factors Associated with Grassland Bird Species Richness: The Relative Roles of Grassland Area, Landscape Structure, and Prey

The factors responsible for widespread declines of grassland birds in the United States are not well understood. This study, conducted in the short-grass prairie of eastern Wyoming, was designed to investigate the relationship between variation in habitat amount, landscape heterogeneity, prey resources, and spatial variation in grassland bird species richness. We estimated bird richness over a 5-year period (1994–1998) from 29 Breeding Bird Survey locations. Estimated bird richness was modeled as a function of landscape structure surrounding survey routes using satellite-based imagery (1996) and grasshopper density and richness, a potentially important prey of grassland birds. Model specification progressed from simple to complex explanations for spatial variation in bird richness. An information-theoretic approach was used to rank and select candidate models. Our best model included measurements of habitat amount, habitat arrangement, landscape matrix, and prey diversity. Grassland bird richness was positively associated with grassland habitat; was negatively associated with habitat dispersion; positively associated with edge habitats; negatively associated with landscape matrix attributes that may restrict movement of grassland bird; and positively related to grasshopper richness. Collectively, 62% of the spatial variation in grassland bird richness was accounted for by the model (adj-R² = 0.514). These results suggest that the distribution of grassland bird species is influenced by a complex mixture of factors that include habitat area affects, landscape pattern and composition, and the availability of prey.     

Importance of Wetland Landscape Structure to Shorebirds Wintering in an Agricultural Valley

Only recently has the influence of landscape structure on habitat use been a research focus in wetland systems. During non-breeding periods when food can be locally limited, wetland spatial pattern across a landscape may be of great importance in determining wetland use. We studied the influence of landscape structure on abundances of wintering Dunlin (Calidris alpina) and Killdeer (Charadrius vociferus) observed on wetlands in the agricultural Willamette Valley of Oregon, USA, during two winters (1999–2000, 2000–2001) of differing rainfall. We examined (1) shorebird use within a sample of 100 km² regions differing in landscape structure (hectares of shorebird habitat [wet, unvegetated]) and (2) use of sites differing in landscape context (area of shorebird habitat within a species-defined radius). For use of sites, we also assessed the influence of two local characteristics: percent of soil exposed and area of wet habitat. We analyzed data using linear regression and information-theoretic modeling. During the dry winter (2000–2001), Dunlin were attracted to regions with more wetland habitat and their abundances at sites increased with greater area of shorebird habitat within both the site and the surrounding landscape. In contrast, Dunlin abundances at sites were related to availability of habitat at only a local scale during the wet winter (1999–2000). Regional habitat availability was of little importance in predicting Killdeer distributions, and Killdeer site use appeared unrelated to habitat distributions at both landscape and local scales. Results suggest prioritizing sites for conservation that are located in areas with high wetland coverage.     

Modelling multi-species response to landscape dynamics: mosaic cycles support urban biodiversity

The importance of the spatial as well as the temporal structure of habitat patches for urban biodiversity has been recognised, but rarely quantified. In dynamic environments the rate of habitat destruction and recreation (i.e. the landscape turnover rate), the minimum amount of potential habitat, its spatial configuration as well as the environmental conditions determining habitat quality are crucial factors for species occurrence. We analysed species responses to environmental parameters and to the spatio-temporal configuration of urban brownfield habitats in a multi-species approach (37 plant and 43 insect species). Species presence/absence data and soil parameters, site age, vegetation structure and landscape context were recorded by random stratified sampling at 133 study plots in industrial areas in the city of Bremen (Germany). Based on the field data, we predicted species occurrences by species distribution models using a multi-model inference approach. Predicted species communities were driven by successional age both at the scale of a single building lot and at the landscape scale. Minimum average succession time of brownfield habitats required to support all and especially regionally rare species depended on the proportion of available open space; the larger the potential habitat area the faster the acceptable turnover. Most plant, grasshopper, and leafhopper species modelled could be maintained at an intermediate turnover rate (mean age of 10–15 years) and a proportion of open sites of at least 40%. Our modelling approach provides the opportunity of inferring optimal spatio-temporal landscape configurations for urban conservation management from patch scale species-environment relationships. The results indicate that urban planning should incorporate land use dynamics into the management of urban biodiversity. 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.     

Role of habitat and landscape in structuring small mammal assemblages in hedgerow networks of contrasted farming landscapes in Brittany,France

In this study, we investigated the environmental factors driving small mammal (rodents and shrews) assemblages in permanent habitat patches in response to a gradient of agricultural intensification. Small mammals were sampled using a trapping standard method in the hedgerow networks of three contrasted landscapes differing by their level of land-use intensity and hedgerow network density (BOC1: slightly intensified; BOC2: moderately intensified and POL: highly intensified). We hypothesized that habitat and landscape characteristics have to be considered to understand the structure of local community. In that way, we carried out a multi-scale study using environmental variables ranging from local habitat (structure and composition of the hedgerows) to hedgerows neighbourhoods in a radius of 300 m (land cover and connectivity around hedges) and to landscape units (three sites). During 1 year, 24 hedgerows were sampled seven times, representing a total of 1,379 captures (86% of rodents and 14% of shrews) and eight species, dominated by the wood mouse (Apodemus sylvaticus) and the bank vole (Clethrionomys glareolus). Inter-site variability was significant and accounted for 18% of total variation in small mammal species abundances. But intra-site variability was also highlighted: species abundance profiles may differ greatly among hedgerows within a site. The more explanatory variables were identified at the different scales of the study: the landscape unit POL was shown to be an important factor in structuring the community, but the predominant factors explaining differences of abundances among hedgerows were about local habitat. In fact, the width of hedges and the tree species richness appeared to be significant and explaining the greatest part of the total variation of the small mammal community composition.     

Area utilization of gulls in a coastal farmland landscape: habitat mosaic supports niche segregation of opportunistic species

The intensively farmed coastal lowland landscape of Germany, adjacent to the North Sea, provides important foraging opportunities for Black-headed, Common, Herring and Lesser Black-backed gull (Larus ridibundus, L. canus, L. argentatus and L. fuscus). We expected that spatial and temporal utilization of the landscape mosaic as well as behavioural traits and utilization of food resources would differ between these closely related species, facilitating niche segregation. We recorded habitat types and their utilization by the four species over a whole year. Furthermore, we related species abundance to several abiotic parameters. Black-headed and Common gulls were the most numerous species in the study area throughout the year. In general, the former species preferred bare fields with recently prepared soils and was often associated with tractors in the fields, whereas the latter species was most often found on pastures. Black-headed gulls seem to have a higher ability to exploit ephemeral, food sources associated with human activities whereas common gulls prefer habitats with low human activity and with naturally distributed prey. The most prominent abiotic parameter influencing gull abundance was presence of tractors. Black-headed gulls have most likely benefited from recent changes in agricultural practice, particularly the increase in cropped land, while Common gulls may have suffered from a decline in pastures. At present, utilization of the farmland habitat mosaic leads to niche segregation and supports coexistence, as two of the four gull species mainly forage in the marine environment, while there is significant habitat partitioning between the other two temporally, spatially and behaviourally.     

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.     

Influence of habitat amount,arrangement, and use on population trend estimates of male Kirtland’s warblers

Kirtland’s warblers (Dendroica kirtlandii) persist in a naturally patchy environment of young, regenerating jack pine forests (i.e., 5–23 years old) created after wildfires and human logging activities. We examined how changing landscape structure from 26 years of forest management and wildfire disturbances influenced population size and spatial dispersion of male Kirtland’s warblers within their restricted breeding range in northern Lower Michigan, USA. The male Kirtland’s warbler population was six times larger in 2004 (1,322) compared to 1979 (205); the change was nonlinear with 1987 and 1994 identified as significant points of change. In 1987, the population trend began increasing after a slowly declining trend prior to 1987, and the rate of increase appeared to slow after 1994. Total amount of suitable habitat and the relative area of wildfire-regenerated habitat were the most important factors explaining population trend. Suitable habitat increased 149% primarily due to increasing plantations from forest management. The relative amount and location of wildfire-regenerated habitat modified the distribution of males among various habitat types, and the spatial variation in their abundance across the primary breeding range. These findings indicate that the Kirtland’s warbler male population shifted its use of habitat types temporally and spatially as the population increased and as the relative availability of habitats changed through time. We demonstrate that researchers and managers need to consider not only habitat quality, but the temporal and the spatial context of habitat availability and population levels when making habitat restoration decisions.     

How do local habitat management and landscape structure at different spatial scales affect fritillary butterfly distribution on fragmented wetlands?

Habitat fragmentation, patch quality and landscape structure are important predictors for species richness. However, conservation strategies targeting single species mainly focus on habitat patches and neglect possible effects of the surrounding landscape. This project assesses the impact of management, habitat fragmentation and landscape structure at different spatial scales on the distribution of three endangered butterfly species, Boloria selene, Boloria titania and Brenthis ino. We selected 36 study sites in the Swiss Alps differing in (1) the proportion of suitable habitat (i.e., wetlands); (2) the proportion of potential dispersal barriers (forest) in the surrounding landscape; (3) altitude; (4) habitat area and (5) management (mowing versus grazing). Three surveys per study site were conducted during the adult flight period to estimate occurrence and density of each species. For the best disperser B. selene the probability of occurrence was positively related to increasing proportion of wetland on a large spatial scale (radius: 4,000 m), for the medium disperser B. ino on an intermediate spatial scale (2,000 m) and for the poorest disperser B. titania on a small spatial scale (1,000 m). Nearby forest did not negatively affect butterfly species distribution but instead enhanced the probability of occurrence and the population density of B. titania. The fen-specialist B. selene had a higher probability of occurrence and higher population densities on grazed compared to mown fens. The altitude of the habitat patches affected the occurrence of the three species and increasing habitat area enhanced the probability of occurrence of B. selene and B. ino. We conclude that, the surrounding landscape is of relevance for species distribution, but management and habitat fragmentation are often more important. We suggest that butterfly conservation should not focus only on a patch scale, but also on a landscape scale, taking into account species-specific dispersal abilities.     

Multi-scale predictive habitat suitability modeling based on hierarchically delineated patches: an example for yellow-billed cuckoos nesting in riparian forests,California, USA

The discipline of landscape ecology recognizes the importance of measuring habitat suitability variables at spatial scales relevant to specific organisms. This paper uses a novel multi-scale hierarchical patch delineation method, PatchMorph, to measure landscape patch characteristics at two distinct spatial scales and statistically relate them to the presence of state-listed endangered yellow-billed cuckoos (Coccyzus americanus occidentalis) nesting in forest patches along the Sacramento River, California, USA. The landscape patch characteristics calculated were: patch thickness, area of cottonwood forest, area of riparian scrub, area of other mixed riparian forest, and total patch area. A third, regional spatial variable, delineating the north and south portions of study area was also analyzed for the effect of regional processes. Using field surveys, the landscape characteristics were related to patch occupancy by yellow-billed cuckoos. The area of cottonwood forest measured at the finest spatial scale of patches was found to be the most important factor determining yellow-billed cuckoo presence in the forest patches, while no patch characteristics at the larger scale of habitat patches were important. The regional spatial variable was important in two of the three analysis techniques. Model validation using an independent data set of surveys (conducted 1987–1990) found 76–82% model accuracy for all the statistical techniques used. Our results show that the spatial scale at which habitat characteristics are measured influences the suitability of forest patches. This multi-scale patch and model selection approach to habitat suitability analysis can readily be generalized for use with other organisms and systems.     

Quantifying determinants contributing to plant species richness in mosaic landscapes: a single- and multi-patch perspective

Despite good theoretical knowledge about determinants of plant species richness in mosaic landscapes, validations based on complete surveys are scarce. We conducted a case study in a highly fragmented, traditional agricultural landscape. In 199 patches of 20 representative multi-patch-plots (MPPs, 1 ha) we recorded a total of 371 plant species. In addition to an additive partitioning of species diversity at the (a) patch- and (b) MPP-scale, we adopted the recently proposed ‘specificity’ measure to quantify the contribution of a spatial subunit to landscape species richness (subunit-to-landscape-contribution, SLC). SLC-values were calculated at both scales with respect to various spatial extents. General regression models were used to quantify the relative importance of hypothesis-driven determinants for species richness and SLC-values. At the patch scale, habitat type was the main determinant of species richness, followed by area and elongated shape. For SLC-values, area was more important than habitat type, and its relevance increased with the extent of the considered landscape. Influences of elongated shape and vegetation context were minor. Differences between habitat types were pronounced for species richness and also partly scale-dependent for SLC-values. Relevant predictors at the MPP-scale were nonlinear habitat richness, the gradient from anthropogenic to seminatural vegetation, and the proportions of natural vegetation and rare habitats. Linear elements and habitat configuration did not contribute to species richness and SLC. Results at the MPP-scale were in complete accordance with the predictions of the mosaic concept. Hence, our study represents its first empirical validation for plant species diversity in mosaic landscapes.     

Cultural landscapes of Germany are patch-corridor-matrix mosaics for an invasive megaforb

Predicting the vulnerability of landscapes to both the initial colonisation and the subsequent spread of invasive species remains a major challenge. The aim of this study was to assess the relative importance of sub-patch level factors and landscape factors for the invasion of the megaforb Heracleum mantegazzianum. In particular, we tested which factors affect the presence in suitable habitat patches and the cover-percentage within invaded patches. For this purpose, we used standard (logistic) regression modelling techniques. The regression analyses were based on inventories of suitable habitat patches in 20 study areas (each 1 km²) in cultural landscapes of Germany. The cover percentage in invaded patches was independent from landscape factors, except for patch shape, and even unsatisfactorily explained by sub-patch level factors included in the analysis (R ² = 0.19). In contrast, presence of H. mantegazzianum was affected by both local and landscape factors. Woody habitat structure decreased the occurrence probability, whereas vicinity to transport corridors (rivers, roads), high habitat connectivity, patch size and perimeter-area ratio of habitat patches had positive effects. The significance of corridors and habitat connectivity shows that dispersal of H. mantegazzianum through the landscape matrix is limited. We conclude that cultural landscapes of Germany function as patch-corridor-matrix mosaics for the spread of H. mantegazzianum. Our results highlight the importance of landscape structure and habitat configuration for invasive spread. Furthermore, this study shows that both local and landscape factors should be incorporated into spatially explicit models to predict spatiotemporal dynamics and equilibrium stages of plant invasions.