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.     

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.     

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.     

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.     

The relative impact of forest patch and landscape attributes on black howler monkey populations in the fragmented Lacandona rainforest, Mexico

Land-use change is forcing many animal populations to inhabit forest patches in which different processes can threaten their survival. Some threatening processes are mainly related to forest patch characteristics, but others depend principally on the landscape spatial context. Thus, the impact of both patch and landscape spatial attributes needs to be assessed to have a better understanding of the habitat spatial attributes that constraint the maintenance of populations in fragmented landscapes. Here, we evaluated the relative effect of three patch-scale (i.e., patch size, shape, and isolation) and five landscape-scale metrics (i.e., forest cover, fragmentation, edge density, mean inter-patch isolation distance, and matrix permeability) on population composition and structure of black howler monkeys (Alouatta pigra) in the Lacandona rainforest, Mexico. We measured the landscape-scale metrics at two spatial scales: within 100 and 500 ha landscapes. Our findings revealed that howler monkeys were more strongly affected by local-scale metrics. Smaller and more isolated forest patches showed a lower number of individuals but at higher densities. Population density also tended to be positively associated to matrices with higher proportion of secondary forests and arboreal crops (i.e. with greater permeability), most probably because these matrices can offer supplementary foods. The immature-to-female ratio also increased with matrix permeability, shape complexity, and edge density; habitat characteristics that can increase landscape connectivity and sources availability. The prevention of habitat loss and isolation, and the increment of matrix permeability are therefore needed for the conservation of this endangered Neotropical mammal.     

How to define a patch: a spatial model for hierarchically delineating organism-specific habitat patches

Landscape analysis and delineation of habitat patches should take into account organism-specific behavioral and perceptual responses to landscape structure because different organisms perceive and respond to landscape features over different ranges of spatial scales. The commonly used methods for delineating habitat based on rules of contiguity do not account for organism-specific responses to landscape patch structure and have undesirable properties, such as being dependent on the scale of base map used for analysis. This paper presents an improved patch delineation algorithm, “PatchMorph,” which can delineate patches across a range of spatial scales based on three organism-specific thresholds: (1) land cover density threshold, (2) habitat gap maximum thickness (gap threshold), and (3) habitat patch minimum thickness (spur threshold). This algorithm was tested on an “idealized” landscape with landscape gaps and spurs of known size, and delineated patches as expected. It was then applied to delineate patches from a neutral random fractal landscape, which showed that as the input gap and spur thickness thresholds were increased, the number of patches decreased from 59 (low thresholds) patches to 1 (high thresholds). The algorithm was then applied to model western yellow-billed cuckoo (Coccyzus americanus occidentalis) nesting habitat patches based on spur and gap thresholds specific to this organism. Both these analyses showed that fewer patches were delineated by PatchMorph than by rules of contiguity, and those patches were larger, had smoother edges, and had fewer gaps within the patches. This algorithm has many applications beyond those presented in this paper, including habitat suitability analysis, spatially explicit population modeling, and habitat connectivity analysis.     

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.     

Using patch isolation metrics to predict animal movement in binary landscapes

Habitat isolation can affect the distribution and abundance of wildlife, but it is an ambiguous attribute to measure. Presumably, isolation is a characteristic of a habitat patch that reflects how spatially inaccessible it is to dispersing organisms. We identified four isolation metrics (nearest-neighbor distance, Voronoi polygons, proximity index, and habitat buffers) that were representative of the different families of metrics that are commonly used in the literature to measure patch isolation. Using simulated data, we evaluated the ability of each isolation metric to predict animal dispersal. We examined the simulated movement of organisms in two types of landscapes: an artificially-generated point-pattern landscapes where patch size and shape were consistent and only the arrangement of patches varied, and realistic landscapes derived from a geographic information system (GIS) of forest-vegetation maps where patch size, shape, and isolation were variable. We tested the performance of the four isolation metrics by examining the strength of the correlation between observed immigration rate in the simulations and each patch isolation metric. We also evaluated whether each isolation metric would perform consistently under varying conditions of patch size/shape, total amount of habitat in the landscape, and proximity of the patch to the landscape edge. The results indicate that a commonly-used distance-based metric, nearest-neighbor distance, did not adequately predict immigration rate when patch size and shape were variable. Area-informed isolation metrics, such as the amount of available habitat within a given radius of a patch, were most successful at predicting immigration. Overall, the use of area-informed metrics is advocated despite the limitation that these metrics require parameterization to reflect the movement capacity of the organism studied.This revised version was published online in May 2005 with corrections to the Cover Date.     

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.     

The role of landscape structure on the abundance of a disease vector planthopper: a quantitative approach

Studies of patchily distributed insect populations have made clear the importance of host patch size and degree of isolation in determining the distribution of these populations. For such populations, patch connectivity will have an effect on patterns of patch occupancy and regional dynamics. In the present study we performed a series of observations to estimate the effect of landscape structure on the abundance of Delphacodes kuscheli (Homoptera: Delphacidae), vector of “Mal de Río Cuarto” disease to maize. Actively dispersing D. kuscheli individuals were collected in 19 sampling sites during the spring of 2004, using sticky traps placed at 2 m above ground level. Land use and landscape pattern were quantified, using Landsat 5 TM images for the area where each sampling site was placed. Four land use categories were considered in the analysis; winter pastures, winter cereals, perennial pasture and stubble. The spatial pattern analysis program FRAGSTATS was employed to estimate the patch area, patch proximity index, Total Class Area and the Mean Proximity Index for each of the land use categories in those sites where insect samples were taken. Partial Least Squares Regression analysis techniques were employed to relate the mean abundance of D. kuscheli and the landscape measures. Eighty percent of the variation of the mean insect abundance was explained by two first PLSR components. The proximity index of the local host patches, the amount of area left to stubble, local host patch area and total area of winter pastures were the most important variables affecting the abundance of dispersing D. kuscheli individuals. We found that the abundance of the dispersive fraction of the population of D. kuscheli is affected mostly by the surrounding landscape, particularly by the proximity of other host patches, and the permeability of the matrix represented by the stubble.     

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.     

Comparing the landscape level perceptual abilities of forest sciurids in fragmented agricultural landscapes*

Perceptual range is the maximum distance from which an animal can perceive the presence of remote landscape elements such as patches of habitat. Such perceptual abilities are of interest because they influence the probability that an animal will successfully disperse to a new patch in a landscape. Furthermore, understanding how perceptual range differs between species may help to explain differential species sensitivity to patch isolation. The objective of this research was to assess the perceptual range of eastern chipmunks (Tamias striatus), gray squirrels (Sciurus carolinensis), and fox squirrels (Sciurus niger) in fragmented agricultural landscapes. Animals were captured in remote woodlots and translocated to unfamiliar agricultural fields. There they were released at different distances from a woodlot and their movements towards or away from the woodlot were used to assess their ability to perceive forested habitat. Observed perceptual ranges of approximately 120 m for chipmunks, 300 m for gray squirrels, and 400 m for fox squirrels, suggest that differences in landscape-level perceptual abilities may influence the occurrence of these species in isolated habitat patches.     

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.     

Matrix effects on plant-frugivore and plant-predator interactions in forest fragments

Structural features of both habitat remnants and surrounding matrix can be important for explaining plant population dynamics and ecosystem functions in human-impacted landscapes. However, little is known about how the structural features of the adjacent matrix affect biotic interactions and whether such context effects are subject to temporal variations. Using the hawthorn Crataegus monogyna in northern Spain, we studied matrix effects on two sequential plant-animal interactions, frugivory by birds and postdispersal seed predation by rodents. Using Hierarchical Linear Models, we compared the magnitude of both interactions on trees located in two patch types that strongly differed in structural features of the adjacent matrix habitat: patches totally surrounded by a degraded, structurally contrasted pastures (unconnected patches), and trees growing in patches adjacent to a lowly degraded, structurally similar mature forests (connected patches). We compared outcomes for 2005 and 2006, which were years with strong differences in community-wide fruit and seed abundance. Frugivory rate did not differ between patch types in either year, likely related to high mobility of birds. Seed predation rates were higher in unconnected patches than in connected ones, but only in 2005. We conclude that strong interannual fluctuations in resource availability are not rare in temperate systems and that recruitment rates could be frequently reduced within unconnected patches, thus collapsing plant regeneration processes of hawthorn populations. Overall, our results suggest that generalizations about potential effects of the matrix on plant-animal interactions within remnant patches must consider: (1) species-specific habitat responses of the organisms, (2) suitability of neighbouring habitats in terms of food supply, and (3) temporal variations in plant-resource availability for interacting animals.     

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.     

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.     

Aspen patch and migratory bird relationships in the northern Yellowstone ecosystem

We evaluated the effects of aspen patch area and orientation (relative to North and an elevational gradient) on the early breeding season abundance and species richness of migratory and resident birds in the northern ungulate winter range of the Yellowstone ecosystem, USA. Using an information-theoretic model selection approach, we found patch area and basal area of aspen to be the most important covariates for long distance migrants, and patch orientation relative to elevational gradient the most important covariate for residents/short-distance migrants. Basal area of live aspen and aspen snags was marginally important for both migratory strategies, likely because aspen snags are an important habitat for most cavity-nesting species. Landscape ecological theory postulates passive interception of dispersing or migrating organisms by patches of suitable habitat. Our results suggest that residents/short-distance migrants are intercepted by patches that are oriented perpendicular to the elevational gradient of our study region resulting in greater abundances and species richness in those patches. However, long-distance migrants appear to use aspen patches without regard to orientation, but rather to patch area.     

The distribution of plant species in urban vegetation fragments

(1) The presence and absence of 22 plant species of various growth forms and habitat associations were analysed in 423 habitat fragments totalling 10.4 km² in a 268 km² urban and suburban region, in Birmingham, UK. (2) Multivariate logistic regressions were used to assess the effects of patch geometry and quality on the species distributions. Measures of geometry were area, shape (S-factor), distance from open countryside and various measures of isolation from other patches. Potential habitat for each species was determined quantitatively, and the distribution of each species was considered within a subset of patches containing potentially suitable habitat types. There was found to be a significant positive correlation between the density of patches available to a species and the proportion of these patches which were occupied. (3) Logistic analyses and incidence functions revealed that, for many of the species, occupancy increased with site age, area, habitat number and similarity of adjacent habitats, while increasing distance to the nearest recorded population of the same species decreased the likelihood that a species would be found in a patch. (4) Patterns of occupancy are consistent with increased extinction from small sites, and colonisation of nearby habitats, coupled with an important role for site history. We conclude that spatial dynamics at the scale of the landscape are of importance to the long-term persistence of many plant species in fragmented landscapes, and must be seriously considered in conservation planning and management. These results have direct implications for the siting and connectivity of urban habitat reserves.     

Spatio-temporal variation in the demographic attributes of a generalist mesopredator

Human land-use practices have dramatically altered the composition and configuration of native habitats throughout many ecosystems. Within heterogeneous landscapes generalist predators often thrive, causing cascading effects on local biological communities, yet there are few data to suggest how attributes of fragmentation influence local population dynamics of these species. We monitored 25 raccoon (Procyon lotor) populations from 2004 to 2009 in a fragmented agricultural landscape to evaluate the influence of local and landscape habitat attributes on spatial and temporal variation in demography. Our results indicate that agricultural ecosystems support increased densities of raccoons relative to many other rural landscapes, but that spatial and temporal variation in demography exists that is driven by non-agricultural habitat attributes rather than the availability of crops. At the landscape scale, both density and population stability were positively associated with the size and contiguity of forest patches, while at the local scale density was positively correlated with plant diversity and the density of tree cavities. In addition, populations occupying forest patches with greater levels of plant diversity and stable water resources exhibited less temporal variability than populations with limited plant species complexity or water availability. The proportion of populations comprised of females was most strongly influenced by the availability of tree cavities and soft mast. Despite the abundance of mesopredators in heterogeneous landscapes, our results indicate that all patches do not contribute equally to the regional abundance and persistence of these species. Thus, a clear understanding of how landscape attributes contribute to variation in demography is critical to the optimization of management strategies.     

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

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