Gypsy moth response to landscape structure differs from neutral model predictions: implications for invasion monitoring

Simulations of dispersal across computer-generated neutral landscapes have generated testable predictions about the relationship between dispersal success and landscape structure. Models predict a threshold response in dispersal success with increasing habitat fragmentation. A threshold is defined as an abrupt, disproportionate decline in dispersal success at a certain proportion of habitat in the landscape. To identify potential empirical threshold responses in invasion success to landscape structure, we quantified the relationship between progression of the gypsy moth (Lymantria dispar) invasion wavefront across Michigan (1985–1996) and the structure of the Michigan landscape using two indices of invasion success and six landscape metrics. We also examined the effect of scale of analysis and choice of land cover characterization on our results by repeating our analysis at three scales using two different land cover maps. Contrary to simulation model predictions, thresholds in invasion success did not correspond closely with thresholds in landscape structure metrics. Increased variation in invasion success indices at smaller scales of analysis also suggested that invasion success should be studied at larger spatial extents (≥75 km²) than would be appropriate for characterizing individual dispersal events. The predictions of individual dispersal models across neutral landscapes may have limited applications for the monitoring and management of vagile species with excellent dispersal capabilities such as the gypsy moth.     

Effect of Landscape Structure on Species Diversity

The effects of habitat fragmentation and their implications for biodiversity is a central issue in conservation biology which still lacks an overall comprehension. There is not yet a clear consensus on how to quantify fragmentation even though it is quite common to couple the effects of habitat loss with habitat fragmentation on biodiversity. Here we address the spatial patterns of species distribution in fragmented landscapes, assuming a neutral community model. To build up the fragmented landscapes, we employ the fractional Brownian motion approach, which in turn permits us to tune the amount of habitat loss and degree of clumping of the landscape independently. The coupling between the neutral community model, here simulated by means of the coalescent method, and fractal neutral landscape models enables us to address how the species–area relationship changes as the spatial patterns of a landscape is varied. The species–area relationship is one of the most fundamental laws in ecology, considered as a central tool in conservation biology, and is used to predict species loss following habitat disturbances. Our simulation results indicate that the level of clumping has a major role in shaping the species–area relationship. For instance, more compact landscapes are more sensitive to the effects of habitat loss and speciation rate. Besides, the level of clumping determines the existence and extension of the power-law regime which is expected to hold at intermediate scales. The distributions of species abundance are strongly influenced by the degree of fragmentation. We also show that the first and second commonest species have approximately self-similar spatial distributions across scales, with the fractal dimensions of the support of the first and second commonest species being very robust to changes in the spatial patterns of the landscape.     

The scale-dependent importance of habitat factors and dispersal limitation in structuring Great Lakes shoreline plant communities

Niche-based and neutral models of community structure posit distinct mechanisms underlying patterns in community structure; correlation between species’ distributions and habitat factors points to niche assembly while spatial pattern independent of habitat suggests neutral assembly via dispersal limitation. The challenge is to disentangle the relative contributions when both processes are operating, and to determine the scales at which each is important. We sampled shoreline plant communities on an island in Lake Michigan, varying the extent and the grain of sampling, and used both distance-based correlation methods and variance partitioning to quantify the proportion of the variation in plant species composition that was attributable to habitat factors and to spatial configuration independent of habitat. Our results were highly scale dependent. We found no distance decay of plant community similarity at the island scale (1−33 km). All of the explained variation (32%) in species composition among samples at this scale was attributed to habitat factors. However, at a site intensively sampled at a smaller scale (5−1,200 m), similarity of species composition did decay with distance. Using a coarse sampling grain (transects), habitat factors explained 40% of the variation, but the purely spatial component explained a comparable 22%. Analyzing plots within transects revealed variation in species composition that was still jointly determined by habitat and spatial factors (18 and 11% of the variance, respectively). For both grain sizes, most of the habitat component was spatially structured, reflecting an abrupt alongshore transition from sandy dunes to cobble beach. Space per se explained more variation in species composition at a second site where the habitat transition was more gradual; here, habitat acted as a less selective filter, allowing the signal of dispersal limitation to be detected more readily. We conclude that both adaptation to specific habitat factors and habitat-independent spatial position indicative of dispersal limitation determine plant species composition in this system. Our results support the prediction that dispersal limitation—a potentially, but not necessarily, neutral driver—is relatively more important at smaller scales.     

Linking landscape history and dispersal traits in grassland plant communities

Dispersal limitation and long-term persistence are known to delay plant species’ responses to habitat fragmentation, but it is still unclear to what extent landscape history may explain the distribution of dispersal traits in present-day plant communities. We used quantitative data on long-distance seed dispersal potential by wind and grazing cattle (epi- and endozoochory), and on persistence (adult plant longevity and seed bank persistence) to quantify the linkages between dispersal and persistence traits in grassland plant communities and current and past landscape configurations. The long-distance dispersal potential of present-day communities was positively associated with the amounts of grassland in the historical (1835, 1938) landscape, and with a long continuity of grazing management—but was not associated with the properties of the current landscape. The study emphasises the role of history as a determinant of the dispersal potential of present-day grassland plant communities. The importance of long-distance dispersal processes has declined in the increasingly fragmented modern landscape, and long-term persistent species are expected to play a more dominant role in grassland communities in the future. However, even within highly fragmented landscapes, long-distance dispersed species may persist locally—delaying the repayment of the extinction debt.     

Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes

Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km²) that differ in the amount and heterogeneity of semi‐natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi‐natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi‐natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi‐natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non‐crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.     

Short Rotation Coppice (SRC) Plantations Provide Additional Habitats for Vascular Plant Species in Agricultural Mosaic Landscapes

Increasing loss of biodiversity in agricultural landscapes is often debated in the bioenergy context, especially with respect to non-traditional crops that can be grown for energy production in the future. As promising renewable energy source and additional landscape element, the potential role of short rotation coppice (SRC) plantations to biodiversity is of great interest. We studied plant species richness in eight landscapes (225 km²) containing willow and poplar SRC plantations (1,600 m²) in Sweden and Germany, and the related SRC α-diversity to species richness in the landscapes (γ-diversity). Using matrix variables, spatial analyses of SRC plantations and landscapes were performed to explain the contribution of SRC α-diversity to γ-diversity. In accordance with the mosaic concept, multiple regression analyses revealed number of habitat types as a significant predictor for species richness: the higher the habitat type number, the higher the γ-diversity and the lower the proportion of SRC plantation α-diversity to γ-diversity. SRC plantation α-diversity was 6.9 % (±1.7 % SD) of species richness on the landscape scale. The contribution of SRC plantations increased with decreasing γ-diversity. SRC plantations were dominated more by species adapted to frequent disturbances and anthropo-zoogenic impacts than surrounding landscapes. We conclude that by providing habitats for plants with different requirements, SRC α-diversity has a significant share on γ-diversity in rural areas and can promote diversity in landscapes with low habitat heterogeneity and low species pools. However, plant diversity enrichment is mainly due to additional species typically present in disturbed and anthropogenic environments.     

Scale-specific correlations between habitat heterogeneity and soil fauna diversity along a landscape structure gradient

Habitat heterogeneity contributes to the maintenance of diversity, but the extent that landscape-scale rather than local-scale heterogeneity influences the diversity of soil invertebrates—species with small range sizes—is less clear. Using a Scottish habitat heterogeneity gradient we correlated Collembola and lumbricid worm species richness and abundance with different elements (forest cover, habitat richness and patchiness) and qualities (plant species richness, soil variables) of habitat heterogeneity, at landscape (1 km²) and local (up to 200 m²) scales. Soil fauna assemblages showed considerable turnover in species composition along this habitat heterogeneity gradient. Soil fauna species richness and turnover was greatest in landscapes that were a mosaic of habitats. Soil fauna diversity was hump-shaped along a gradient of forest cover, peaking where there was a mixture of forest and open habitats in the landscape. Landscape-scale habitat richness was positively correlated with lumbricid diversity, while Collembola and lumbricid abundances were negatively and positively related to landscape spatial patchiness. Furthermore, soil fauna diversity was positively correlated with plant diversity, which in turn peaked in the sites that were a mosaic of forest and open habitat patches. There was less evidence that local-scale habitat variables (habitat richness, tree cover, plant species richness, litter cover, soil pH, depth of organic horizon) affected soil fauna diversity: Collembola diversity was independent of all these measures, while lumbricid diversity positively and negatively correlated with vascular plant species richness and tree canopy density. Landscape-scale habitat heterogeneity affects soil diversity regardless of taxon, while the influence of habitat heterogeneity at local scales is dependent on taxon identity, and hence ecological traits, e.g. body size. Landscape-scale habitat heterogeneity by providing different niches and refuges, together with passive dispersal and population patch dynamics, positively contributes to soil faunal diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Relative importance of landscape and species characteristics on extinction debt,immigration credit and relaxation time after habitat turnover

Habitat turnover concomitantly causes destruction and creation of habitat patches. Following such a perturbation, metapopulations harbor either an extinction debt or an immigration credit, that is the future decrease or increase in population numbers due to this disturbance. Extinction debt and immigration credit are rarely considered simultaneously and disentangled from the relaxation time (time to new equilibrium). In this contribution, we test the relative importance of two potential drivers of time-delayed metapopulation dynamics: the spatial configuration of the habitat turnover and species dispersal ability. We provide a simulation-based investigation projecting metapopulation dynamics following habitat turnover in virtual landscapes. We consider two virtual species (a short-distance and a long-distance disperser) and five scenarios of habitat turnover depending on net habitat loss or gain and habitat aggregation. Our analyses reveal that (a) the main determinant of the magnitude of the extinction debt or immigration credit is the net change in total habitat area, followed by species dispersal distance and finally by the post-turnover habitat aggregation; (b) relaxation time weakly depends on the magnitude of the immigration credit or of the extinction debt; (c) the main determinant of relaxation time is dispersal distance followed by the net change in total habitat area and finally by the post-turnover habitat aggregation. These results shed light on the relative importance of dispersal ability and habitat turnover spatial structure on the components of time-delayed metapopulation dynamics.     

Competing populations on fragmented landscapes with spatially structured heterogeneities: improved landscape generation and mixed dispersal strategies

Interactions between two species competing for space were studied using stochastic spatially explicit lattice-based simulations as well as pair approximations. The two species differed only in their dispersal strategies, which were characterized by the proportion of reproductive effort allocated to long-distance (far) dispersal versus short-distance (near) dispersal to adjacent sites. All population dynamics took place on landscapes with spatially clustered distributions of suitable habitat, described by two parameters specifying the amount and the local spatial autocorrelation of suitable habitat. Whereas previous results indicated that coexistence between pure near and far dispersers was very rare, taking place over only a very small region of the landscape parameter space, when mixed strategies are allowed, multiple strategies can coexist over a much wider variety of landscapes. On such spatially structured landscapes, the populations can partition the habitat according to local conditions, with one species using pure near dispersal to exploit large contiguous patches of suitable habitat, and another species using mixed dispersal to colonize isolated smaller patches (via far dispersal) and then rapidly exploit those patches (via near dispersal). An improved mean-field approximation which incorporates the spatially clustered habitat distribution is developed for modeling a single species on these landscapes, along with an improved Monte Carlo algorithm for generating spatially clustered habitat distributions.      

Frugivory and spatial patterns of seed deposition by carnivorous mammals in anthropogenic landscapes: a multi-scale approach

Background

Knowledge about how frugivory and seed deposition are spatially distributed is valuable to understand the role of dispersers on the structure and dynamics of plant populations. This may be particularly important within anthropogenic areas, where either the patchy distribution of wild plants or the presence of cultivated fleshy-fruits may influence plant-disperser interactions.

Methodology/Principal Findings

We investigated frugivory and spatial patterns of seed deposition by carnivorous mammals in anthropogenic landscapes considering two spatial scales: ‘landscape’ (∼10 km²) and ‘habitat type’ (∼1–2 km²). We sampled carnivore faeces and plant abundance at three contrasting habitats (chestnut woods, mosaics and scrublands), each replicated within three different landscapes. Sixty-five percent of faeces collected (n = 1077) contained seeds, among which wild and cultivated seeds appeared in similar proportions (58% and 53%) despite that cultivated fruiting plants were much less abundant. Seed deposition was spatially structured among both spatial scales being different between fruit types. Whereas the most important source of spatial variation in deposition of wild seeds was the landscape scale, it was the habitat scale for cultivated seeds. At the habitat scale, seeds of wild species were mostly deposited within mosaics while seeds of cultivated species were within chestnut woods and scrublands. Spatial concordance between seed deposition and plant abundance was found only for wild species.

Conclusions/Significance

Spatial patterns of seed deposition by carnivores differed between fruit types and seemed to be modulated by the fleshy-fruited plant assemblages and the behaviour of dispersers. Our results suggest that a strong preference for cultivated fruits by carnivores may influence their spatial foraging behaviour and lower their dispersal services to wild species. However, the high amount of seeds removed within and between habitats suggests that carnivores must play an important role – often overlooked – as ‘restorers’ and ‘habitat shapers’ in anthropogenic areas.     

Landscape size affects the relative importance of habitat amount,habitat fragmentation,and matrix quality on forest birds

It is important to understand the relative effects of landscape habitat loss, habitat fragmentation, and matrix quality on biodiversity, so that potential management options can be appropriately ranked. However, their effects and relative importance may change with the size of the landscape considered because the multiple (and potentially conflicting) ecological processes that are influenced by landscape structure occur at different spatial scales (e.g. dispersal, predation, foraging). We estimated the relative effects of habitat loss, habitat fragmentation, and matrix quality (measured as the amount of forest, the proportion of forest area contained in large core forests, and the density of roads respectively) on fragmentation‐sensitive forest birds in southern Ontario, Canada using a range of landscape sizes (0.8–310 km²). We used three complementary statistical approaches to estimate relative effects of these correlated landscape factors – 1) multiple regression, 2) information theoretic (AIC) estimates of the most parsimonious model, and 3) multi‐model inference to average effects across all supported models. We controlled for spatial autocorrelation, local habitat, roadside sampling bias, time of day, season, habitat heterogeneity, and the interaction between the effects of habitat amount and fragmentation. We found that relative effects of habitat amount and fragmentation were scale dependent; habitat amount had a consistently positive effect that was consistent over more than two orders of magnitude in landscape area (~1–300 km²). In contrast, the effects of habitat fragmentation depended on the size of the landscape considered. Indeed, for veery Catharus fuscescens, habitat fragmentation had positive effects at one scale and negative effects at another. The effects of matrix quality were generally weak and changed little with scale. For the number of fragmentation sensitive species and the presence of veery, habitat amount was most important in large landscapes and habitat fragmentation in small landscapes but for the presence of ovenbird Seiurus aurocapilla, habitat amount was most important at all scales.     

Niche evolution in Australian terrestrial mammals? Clarifying scale-dependencies in phylogenetic and functional drivers of co-occurrence

Interactive forces between competition and habitat filtering drive many biogeographic patterns over evolutionary time scales. However, the responsiveness of assemblages to these two forces is highly influenced by spatial scale, forming complex patterns of niche separation. We explored these spatial dependencies by quantifying the influence of phylogeny and functional traits in shaping present day native terrestrial mammal assemblages at multiple scales, principally by identifying the spatial scales at which niche evolution operates. We modelled the distribution of 53 native terrestrial mammal species across New South Wales, Australia. Using predicted distributions, we estimated the range overlap between each pair of species at increasing grain sizes (~0.8, 5.1, 20, 81, 506, 2,025, 8,100 km²). We employed a decision tree to identify how interactions among functional traits and phylogenetic relatedness translated to levels of sympatry at increasing spatial scales. We found that Australian terrestrial mammals displayed phylogenetic over-dispersion that was inversely related to spatial scale, suggesting that ecological processes were more influential than biogeographic sympatry patterns in defining assemblages of species. While the contribution of phylogenetic relatedness to patterns of co-occurrence decreased as spatial scale increased, the reverse was true for habitat preferences. At the same time, functional traits also operated at different scales, as dietary preferences dominated at local spatial scales (<10 km²) while body mass has a stronger effect at larger spatial scales. Our findings show that ecological and evolutionary processes operate at different scales and that Australian terrestrial mammals diverged slower along their micro-scale niche compared to their macro-scale niche. By combining phylogenetic and niche methods through the modelling of species distributions, we assessed whether specific traits were related to a particular niche. More importantly, conducting multi-scale spatial analysis avoids categorical assignment of traits-to-niches, providing a clearer relationship between traits and a species ecological niche and a more precise scaling for the axes of niche evolution.     

Grazing effects on plant functional group diversity in Mediterranean shrublands

Grazing is one of the prevalent human activities that even today are taking place inside protected areas with direct or indirect effects on ecosystems. In this study we analyzed the effects of grazing on plant species diversity, plant functional group (PFG) diversity and community composition of shrublands. We analyzed plant diversity data from 582 sampling plots located in 66 protected areas of the Greek Natura 2000 network, containing in total 1102 plant species and subspecies. We also classified a priori all plant species in seven PFGs: annual forbs, annual grasses/sedges, legumes, perennial forbs, perennial grasses/sedges, small shrubs and tall shrubs. For each site, grazing intensity was estimated in four classes (no grazing, low, medium and high grazing intensity). We found that, at the spatial and temporal scale of this study, as grazing intensity increased, so did total species richness. However, each PFG displayed a different response to grazing. Short-lived species (annual grasses or forbs and legumes) benefited from grazing and their species richness and proportion in the community increased with grazing. Perennial grasses and forbs species richness increased with grazing intensity, but their dominance decreased, since their proportion in the community declined. Short shrub species richness remained unaffected by grazing, while tall shrub diversity decreased. Finally, in sites without grazing the spatial pattern of species richness of the different PFGs was not congruent with each other, while in grazed sites they were significantly positively correlated (with the exception of tall shrubs). This finding may imply that grazing is a selective pressure organizing the community structure, and imposing a certain contribution of each PFG. So, in Mediterranean shrublands in protected areas with a long historical record of grazing, it seems that grazing promotes species diversity and its continuation on a portion of the landscape may be a necessary part of an effective management plan.     

Using Multiscale Spatial Models to Assess Potential Surrogate Habitat for an Imperiled Reptile

In evaluating conservation and management options for species, practitioners might consider surrogate habitats at multiple scales when estimating available habitat or modeling species’ potential distributions based on suitable habitats, especially when native environments are rare. Species’ dependence on surrogates likely increases as optimal habitat is degraded and lost due to anthropogenic landscape change, and thus surrogate habitats may be vital for an imperiled species’ survival in highly modified landscapes. We used spatial habitat models to examine a potential surrogate habitat for an imperiled ambush predator (eastern diamondback rattlesnake, Crotalus adamanteus; EDB) at two scales. The EDB is an apex predator indigenous to imperiled longleaf pine ecosystems (Pinus palustris) of the southeastern United States. Loss of native open-canopy pine savannas and woodlands has been suggested as the principal cause of the species’ extensive decline. We examined EDB habitat selection in the Coastal Plain tidewater region to evaluate the role of marsh as a potential surrogate habitat and to further quantify the species’ habitat requirements at two scales: home range (HR) and within the home range (WHR). We studied EDBs using radiotelemetry and employed an information-theoretic approach and logistic regression to model habitat selection as use vs. availability. We failed to detect a positive association with marsh as a surrogate habitat at the HR scale; rather, EDBs exhibited significantly negative associations with all landscape patches except pine savanna. Within home range selection was characterized by a negative association with forest and a positive association with ground cover, which suggests that EDBs may use surrogate habitats of similar structure, including marsh, within their home ranges. While our HR analysis did not support tidal marsh as a surrogate habitat, marsh may still provide resources for EDBs at smaller scales.     

Additive Partitioning of Diversity Reveals No Scale-dependent Impacts of Large Ungulates on the Structure of Tundra Plant Communities

Large herbivores can change ecosystem functioning by impacting plant diversity. However, although such impacts are expected to be scale-dependent in ecosystems with wide-roaming ungulates, scaling issues rarely enter empirical assessments. We here test the hypothesis that the impact of increased reindeer abundance on plant diversity in alpine tundra is scale-dependent. Based on potentially high productivity of the focal habitat units and hence the possibility of positive grazer impacts on plant diversity we predicted higher α and β diversity at the habitat scale where reindeer densities are high. We also explored whether there were differences in diversity patterns at larger scales, including the scale of reindeer management districts. We estimated grazing disturbance as high versus low reindeer density in selected districts (a total extent of 7421 km²) of Northern Norway where reindeer-induced vegetation shifts are debated. We focus on dominance patterns because they can quantify the vegetation state and thus performed additive partitioning of Simpson diversity on multiple scales assessing also species’ contributions to diversity. Contrary to our predictions, we found only weak scale-dependent effects of reindeer grazing on plant diversity. Under high reindeer densities there was evidence for a landscape-scale homogenization of the vegetation, but the predicted α and β diversity increases at the habitat scale were not found. Consistently through all scales considered, four shrub species contributed the most to plant diversity. These results contradict the idea that reindeer at high stocking densities induce shifts in plant species dominance in productive habitats. We conclude that context-dependencies such as spatial scales of management units and habitat types need to be explicitly considered in evaluations of the impacts of large ungulates on plant diversity.     

Persistence of Dispersal-limited Species in Structured Dynamic Landscapes

Dynamic landscape models have generally assumed random distributions of habitat although real landscapes show spatial organization at many scales. To explore the role of spatial structure in determining the frequency of dispersal-limited forest species, we used a cellular landscape model divided into two zones. Zones were distributed in a random, clustered, or regular spatial pattern. Within each zone habitat cells were randomly destroyed and regenerated, and habitat density and turnover rate were systematically varied. A hypothetical habitat-limited species dispersed between adjacent habitat cells. All trials showed a reduced species frequency relative to a static landscape. Reduction was greater at low habitat density (P = 0.30) than at high density (0.90) suggesting the importance of habitat connectivity in controlling species frequency. The greatest reduction occurred when habitat was concentrated in a small, regularly distributed zone at low habitat density reflecting the enforced isolation of individual habitat cells. Very little reduction was observed when habitat cells were packed into a small clustered zone, a situation promoting connectivity between cells. Moderate–severe frequency reduction occurred when habitat turnover was concentrated in a clustered zone at high habitat density, but little was observed when turnover was widely distributed in a regular or random pattern. These results can be interpreted in terms of a source-sink function in which spatial pattern controlled the degree of contact between landscape zones and determined opportunities for dispersal between habitat cells. We conclude that clustering of forest habitat has the potential to maintain herb species frequency in sparsely forested landscapes. Conversely, clustering of forest disturbance in heavily forested regions, or regular distribution of forest stands (as often occurs in agricultural regions) creates areas which are difficult to colonize, and should be avoided.     

Distribution and abundance of dung beetles in fragmented landscapes

Related species utilising similar resources are often assumed to show similar spatial population structures and dynamics. This paper reports substantial ecological variation within a set of Aphodius dung beetles occurring in the same patchily distributed resource, livestock dung in pastures. We show how variation in habitat and resource selectivity, in the rate of movements between pastures, and in the distribution of local population sizes all contribute to interspecific differences in spatial population structures. Local dung beetle assemblages are compared between two landscapes with different densities of pastures. In one of the landscapes, we contrast the abundances and regional distributions of Aphodius before and after 15 years of rapid habitat loss. Different species show very dissimilar responses to changes in the structure of the landscape. Our results suggest that generalist Aphodius species, and specialist species with high dispersal powers, occur as large "patchy" populations in the landscape. In contrast, a strict pasture specialist species with limited dispersal powers (A. pusillus) forms classical metapopulations. At the community level, interspecific differences in spatial population structures make the local community composition a function of the structure of the surrounding landscape.     

Spatial movements of grey partridge Perdix perdix : male-biased spring dispersal and effect of habitat quality

The Grey Partridge Perdix perdix is a sedentary species with a surplus of males in spring populations. It experienced a dramatic decline in Europe primarily due to habitat loss that may have forced the species to modify its dispersal behaviour, a less transparent source of populations threats. We analysed the dispersal patterns of Grey Partridges during pre- and post-breeding periods in two landscapes with different habitat quality in the Czech Republic. Grey Partridges tended to pair off quickly, with most pre-breeding movements executed in pairs. The birds usually dispersed over a few hundred metres indicating that the habitat conditions in a landscape with a carrying capacity of 2–5 pairs/1 km² was not below the threshold initiating strong dispersal, as observed in extreme habitats and at the northern limits of the species′ distribution range. A few single males that had remained unpaired until April also finally bred. However, the high level of sedentariness of males in a good landscape suggested that males prefer to stay near wintering sites unless habitat sources become limited. Unlike the males, the females dispersed in a wide range of distances with an extensive overlap in both landscapes. Autumn movements did not differ between the sexes, but were longer than the pre-breeding movements and increased in a poor landscape. We conclude that both reduced habitat availability and reduced mating opportunities initialise primarily male-biased breeding dispersal in Grey Partridge populations.     

The relationship between multi‐scale habitat heterogeneity and farmland bird abundance

It has recently been proposed that losses in farmland habitat heterogeneity may have been a primary driver of the profound declines exhibited by many farmland bird species in recent decades. However, it has yet to be demonstrated which facets of heterogeneity and what spatial scales are most important for birds. Here we analyse the relationship between abundance and features of landscape heterogeneity at three spatial scales (1, 9 and 25 km²) for 32 bird species commonly associated with farmland. Heterogeneity was quantified using three contrasting indices reflecting 1) the spatial mixing of land uses, 2) variation in field sizes and 3) the density of field boundaries. The spatial mixing of land‐uses explained, on average, the most variation in, and was most likely to be positively associated with, abundance at all spatial scales. The majority of species (66–75%, depending on the spatial scale) were more common in heterogeneous landscapes overall; however, migrants, those under a high level of conservation concern and farmland specialist species tended to be less abundant in more heterogeneous landscapes at all scales. Ground‐nesting species were also more likely to be found in more homogeneous habitats than non‐ground‐nesters, but only at the finest spatial scale. Relationships between abundance and heterogeneity were generally consistent across spatial scales; however, species of high conservation concern had more variable associations compared with other species. These results highlight a potential role for farmland habitat heterogeneity in determining the abundance of many farmland species but suggest that population responses to an increase in heterogeneity would not be unanimously positive and would probably have negative impacts on some species, notably those that are already threatened.     

Spatial Heterogeneity in Habitat Quality and Cross-Scale Interactions in Metapopulations

Abstract Integration of habitat heterogeneity into spatially realistic metapopulation approaches reveals the potential for key cross-scale interactions. Broad-scale environmental gradients and land-use practices can create autocorrelation of habitat quality of suitable patches at intermediate spatial scales. Patch occupancy then depends not only on habitat quality at the patch scale but also on feedbacks from surrounding neighborhoods of autocorrelated patches. Metapopulation dynamics emerge from how demographic and dispersal processes interact with relevant habitat heterogeneity. We provide an empirical example from a metapopulation of round-tailed muskrats (Neofiber alleni) in which habitat quality of suitable patches was spatially autocorrelated most strongly within 1,000 m, which was within the expected dispersal range of the species. After controlling for factors typically considered in metapopulation studies—patch size, local patch quality, patch connectivity—we use a cross-variogram analysis to demonstrate that patch occupancy by muskrats was correlated with habitat quality across scales ≤1,171 m. We also discuss general consequences of spatial heterogeneity of habitat quality for metapopulations related to potential cross-scale interactions. We focus on spatially correlated extinctions and metapopulation persistence, hierarchical scaling of source–sink dynamics, and dispersal decisions by individuals in relation to information constraints.