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.     

Successional change in the insect community of a fragmented landscape

Habitat fragmentation strongly affects insect species diversity and community composition, but few studies have examined landscape effects on long term development of insect communities. As mobile consumers, insects should be sensitive to both local plant community and landscape context. We tested this prediction using sweep-net transects to sample insect communities for 8 years at an experimentally fragmented old-field site in northeastern Kansas, USA. The site included habitat patches undergoing secondary succession, surrounded by a low turf matrix. During the first 5 years, plant richness and cover were measured in patches. Insect species richness, total density, and trophic diversity increased over time on all transects. Cover of woody plants and perennial forbs increased each year, adding structural complexity to successional patches and potentially contributing to increased insect diversity. Within years, insect richness was significantly greater on transects through large successional patches (5000 m²) than on transects through fragmented arrays of 6 medium-sized (total area 1728 m²) or 15 small (480 m²) patches. However, plant cover did not differ among patch types and was uncorrelated with insect richness within years. Insect richness was strongly correlated with insect density, but trophic and α diversities did not differ among patch types, indicating that patch insect communities were subsets of a common species pool. We argue that differences in insect richness resulted from landscape effects on the size of these subsets, not patch succession rates. Greater insect richness on large patches can be explained as a community-level consequence of population responses to resource concentration.     

Large-scale experimental landscapes reveal distinctive effects of patch shape and connectivity on arthropod communities

The size, shape, and isolation of habitat patches can affect organism behavior and population dynamics, but little is known about the relative role of shape and connectivity in affecting ecological communities at large spatial scales. Using six sampling sessions from July 2001 until August 2002, we collected 33,685 arthropods throughout seven 12-ha experimental landscapes consisting of clear-cut patches surrounded by a matrix of mature pine forest. Patches were explicitly designed to manipulate connectivity (via habitat corridors) independently of area and edge effects. We found that patch shape, rather than connectivity, affected ground-dwelling arthropod richness and beta diversity (i.e. turnover of genera among patches). Arthropod communities contained fewer genera and exhibited less turnover in high-edge connected and high-edge unconnected patches relative to low-edge unconnected patches of similar area. Connectivity, rather than patch shape, affected the evenness of ground-dwelling arthropod communities; regardless of patch shape, high-edge connected patches had lower evenness than low- or high-edge unconnected patches. Among the most abundant arthropod orders, increased richness in low-edge unconnected patches was largely due to increased richness of Coleoptera, whereas Hymenoptera played an important role in the lower evenness in connected patches and patterns of turnover. These findings suggest that anthropogenic habitat alteration can have distinct effects on ground-dwelling arthropod communities that arise due to changes in shape and connectivity. Moreover, this work suggests that corridors, which are common conservation tools that change both patch shape and connectivity, can have multiple effects on arthropod communities via different mechanisms, and each effect may alter components of community structure.     

Determinants of plant species richness and patterns of nestedness in fragmented landscapes: evidence from land-bridge islands

Land-bridge islands formed by dam construction are considered to be “experimental” systems for studying the effects of habitat loss and fragmentation, offering many distinct advantages over terrestrial fragments. The Thousand Island Lake in Southeast China is one such land-bridge system with more than 1000 islands. Based on a field survey of vascular plant richness on 154 land-bridge islands during 2007–2008, we examined the effects of island and landscape attributes on plant species richness and patterns of species nestedness. We also examined the different responses of plant functional groups (classified according to growth form and shade tolerance) to fragmentation. We found that island area explained the greatest amount of variation in plant species richness. Island area and shape index positively affected species diversity and the degree of nestedness exhibited by plant communities while the perimeter to area ratio of the islands had a negative effect. Shade-tolerant plants were the most sensitive species group to habitat fragmentation. Isolation negatively affected the degree of nestedness in herb and shade-intolerant plants including species with various dispersal abilities in the fragmented landscape. Based on these results, we concluded that the effects of habitat loss and fragmentation on overall species richness depended mostly on the degree of habitat loss, but patterns of nestedness were generated from different ecological mechanisms due to species-specific responses to different characteristics of habitat patches.     

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.     

Scaling properties in landscape patterns: New Zealand experience

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

Comparing the role of site disturbance and landscape properties on understory species richness in fragmented periurban Mediterranean forests

We hypothesized that the spatial configuration and dynamics of periurban forest patches in Barcelona (NE of Spain) played a minor role in determining plant species richness and assemblage compared to site conditions, and particularly to both direct (measured at plot level) and potential (inferred from landscape metrics) human-associated site disturbance. The presence of all understory vascular plants was recorded on 252 plots of 100 m² randomly selected within forest patches ranging in size from 0.25 ha to 218 ha. Species were divided into 6 groups, according to their ecology and conservation status. Site condition was assessed at plot level and included physical attributes, human-induced disturbance and Quercus spp. tree cover. Landscape structure and dynamics were assessed from patch metrics and patch history. We also calculated a set of landscape metrics related to potential human accessibility to forests. Results of multiple linear regressions indicated that the variance explained for non-forest species groups was higher than for forest species richness. Most of the main correlates corresponded to site disturbance variables related to direct human alteration, or to landscape variables associated to indirect human effects on forests: Quercus tree cover (a proxy for successional status) was the most important correlate of non-forest species richness, which decreased when Quercus tree cover increased. Human-induced disturbance was an important correlate of synanthropic and total species richness, which were higher in recently managed and in highly frequented forests. Potential human accessibility also affected the richness of most species groups. In contrast, patch size, patch shape and connectivity played a minor role, as did patch history. We conclude that human influence on species richness in periurban forests takes place on a small scale, whereas large-scale effects attributable to landscape structure and fragmentation are comparatively less important. Implications of these results for the conservation of plant species in periurban forests are discussed.     

The landscape matrix modifies the effect of habitat fragmentation in grassland butterflies

The landscape matrix is suggested to influence the effect of habitat fragmentation on species richness, but the generality of this prediction has not been tested. Here, we used data from 10 independent studies on butterfly species richness, where the matrix surrounding grassland patches was dominated by either forest or arable land to test if matrix land use influenced the response of species richness to patch area and connectivity. To account for the possibility that some of the observed species use the matrix as their main or complementary habitat, we analysed the effects on total species richness and on the richness of grassland specialist and non-specialist (generalists and specialists on other habitat types) butterflies separately. Specialists and non-specialists were defined separately for each dataset. Total species richness and the richness of grassland specialist butterflies were positively related to patch area and forest cover in the matrix, and negatively to patch isolation. The strength of the species-area relationship was modified by matrix land use and had a slope that decreased with increasing forest cover in the matrix. Potential mechanisms for the weaker effect of grassland fragmentation in forest-dominated landscapes are (1) that the forest matrix is more heterogeneous and contains more resources, (2) that small grassland patches in a matrix dominated by arable land suffer more from negative edge effects or (3) that the arable matrix constitutes a stronger barrier to dispersal between populations. Regardless of the mechanisms, our results show that there are general effects of matrix land use across landscapes and regions, and that landscape management that increases matrix quality can be a complement to habitat restoration and re-creation in fragmented landscapes.     

Dispersal behaviour in fragmented landscapes: Deriving a practical formula for patch accessibility

Dispersal has been shown to be a key determinant of spatially structured populations. One crucial aspect is predicting patch accessibility: the probability r_ij of a certain patch j being reached by individuals starting at another patch i. Patch accessibility r_ij depends on both the landscape structure and the individuals’ dispersal behaviour. To investigate the effects of these factors on r_ij, we developed a simulation model focusing on animal dispersal. Our model analyses show that there is an important intrinsic effect of the interplay between landscape structure and dispersal behaviour on patch accessibility: the competition between patches for migrants. We derive a formula for patch accessibility. This formula is very simple because it just takes distances into account: not only the distance between start patch and target patch, but also between the start patch and all the other patches in the landscape. Despite its simplicity, the formula is able to cover effects such as the competition for migrants. The formula was found to have high predictive power for a variety of movement behaviours (random walk with various degrees of correlation, Archimedean spirals and loops) in any given landscape. The formula can be interpreted as a generic function for patch accessibility for further population dynamics analyses. It also delivers insights into the consequences of dispersal in fragmented landscapes.     

Modeling scale-dependent landscape pattern,dispersal, and connectivity from the perspective of the organism

Understanding the impacts of habitat fragmentation on dispersal is an important issue in landscape and conservation ecology. Here I examine the effects of fine- to broad-scale patterns in landscape structure on dispersal success of organisms with differing life-history traits. An individual-based model was used to simulate dispersal of amphibian-like species whose movements were driven by land cover and moisture conditions. To systematically control spatial pattern, a landscape model was created by merging simulated land cover maps with synthetic topographic surfaces. Landscapes varied in topographic roughness and spatial contagion in agriculture and urban land cover. Simulations included three different species types that varied in their maximum potential dispersal distances by 1-, 2-, or 4-fold. Two sets of simulations addressed effects of varying aspects of landscape structure on dispersal success. In the first set of simulations, which incorporated variable distances between breeding patches, dispersal success was lowest for all species types when anthropogenic cover was patchily distributed. In the second set, with interpatch distances held constant as landscape composition varied, dispersal success decreased as anthropogenic cover became spatially contagious. Both sets revealed strong main effects of species characteristics, interpatch distances and landscape composition on dispersal success; furthermore, scale-dependent patterns in land cover and moisture gradients had a stronger effect on longer- than shorter-ranging species types. Taken together, these simulations suggest that heuristic conservation strategies could potentially be developed based on important but limited life history information.     

Assessing habitat fragmentation on marine epifaunal macroinvertebrate communities: an experimental approach

Habitat fragmentation is considered a major cause of biodiversity loss, both on terrestrial and marine environments. Understanding the effects of habitat fragmentation on the structure and dynamics of natural communities is extremely important to support management actions for biodiversity conservation. However, the effects of habitat fragmentation on marine communities are still poorly understood. Here we evaluated whether habitat fragmentation affects the structure of epifaunal communities in the sublittoral zone, in the northern coast of São Paulo state, Brazil. Five experimental landscapes were constructed, each one forming a large continuous patch. After 4 weeks, each landscape was cut on three patches of different sizes. Epifaunal macroinvertebrate communities were sampled at the edge and interior of experimental landscapes before manipulation to evaluate edge effects. After four more weeks, communities from the three patch sizes were also sampled to evaluate patch size effects. We compared the diversity of communities at different levels of fragmentation by total abundance, rarefied taxon richness, Shannon–Wiener diversity index, Simpson’s dominance index, and abundance of dominant taxa. Higher taxon richness and gastropod abundance were recorded in the patch edges, but no significant differences were found among patch sizes. We found a significant effect of habitat fragmentation, with lower abundances of Gammaridea (the dominant taxon), Ophyuroidea, and Pycnogonida after the experimental fragmentation. Lower abundances of dominant taxa resulted in higher diversity and lower dominance in fragmented landscapes when compared to integral, pre-manipulation landscapes. Our results suggest that fragmentation of landscapes in the system studied can reduce dominance, and that even small patch sizes can be important for the conservation of macroinvertebrate diversity.     

Confounding of patch quality and matrix effects in herbivore movement studies

Although the landscape matrix is increasingly incorporated into spatial-ecological population studies, little consideration has been given to the likely possibility that patch quality is confounded with the composition of the matrix surrounding each patch. For example, the nutritional quality of host-plant patches to an herbivore may be highly correlated with matrix composition, consequently obfuscating the importance of the matrix itself to interpatch dispersal. From a literature survey of the effects of the matrix on herbivore movement among host-plant patches, we found that 55% of the studies (6/11) failed to experimentally or statistically isolate the effects of the matrix from potential patch-quality effects on dispersal. Most studies consisted of mark-recapture experiments in natural landscapes where patch equality was not controlled or manipulated. Of the few studies that evaluated the relationship between matrix composition and patch quality, all of them (3/3) found that these two landscape factors covaried. These data suggest that in most matrix studies, effects of the matrix on dispersal may be wholly, or in part, due to underlying differences in patch quality. This revised version was published online in May 2005 with corrections to the Cover Date. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of corridor width and presence on the population dynamics of the meadow vole (Microtus pennsylvanicus)

We tested the effects of increased landscape corridor width and corridor presence on the population dynamics and home range use of the meadow vole (Microtus pennsylvanicus) within a small-scale fragmented landscape. Our objective was to observe how populations behaved in patchy landscapes where the animals home range exceeded or equaled patch size. We used a small-scale replicated experiment consisting of three sets of two patches each, unconnected or interconnected by 1-m or 5-m wide-corridors, established in an old-field community (S.W. Ohio). Control (0-m) treatments supported significantly lower vole densities than either corridor treatment. Females were the dominant resident sex establishing smaller home ranges (<150m²) than males (>450m²). Significantly more male voles dispersed between patches with corridors than between patches without corridors. However, no difference was observed regarding the number of male voles dispersing between patches connected by corridors when compared to the number dispersing across treatments. Dispersal between connected patches was restricted to corridors based on tracking tube data. Corridor presence was more important than corridor width regarding the movement of male voles within their home range.     

Sacrificing patches for linear habitat elements enhances metapopulation performance of woodland birds in fragmented landscapes

It is generally assumed that large patches of natural habitat are better for the survival of species than the same amount of habitat in smaller fragments or linear elements like hedges and tree rows. We use a spatially explicit individual-based model of a woodland bird to explore this hypothesis. We specifically ask whether mixtures of large, small and linear habitat elements are better for population performance than landscapes that consist of only large elements. With equal carrying capacity, metapopulations perform equally or better in heterogeneous landscape types that are a mix of linear, large and small habitat elements. We call this increased metapopulation performance of large and small elements “synergy”. These mixed conditions are superior because the small linear elements facilitate dispersal while patches secure the population in the long run because they have a lower extinction risk. The linear elements are able to catch and guide dispersing animals which results in higher connectivity between patches leading to higher metapopulation survival. Our results suggest that landscape designers should not always seek to conserve and create larger units but might better strive for more variable landscapes with mixtures of patch sizes and shapes. This is especially important when smaller units play a key role in connecting patches and dispersal through the matrix is poor.     

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

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

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.     

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

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

Environmental factors at different spatial scales governing soil fauna community patterns in fragmented forests

Spatial and temporal changes in community structure of soil organisms may result from a myriad of processes operating at a hierarchy of spatial scales, from small-scale habitat conditions to species movements among patches and large-sale landscape features. To disentangle the relative importance of spatial and environmental factors at different scales (plot, patch and landscape), we analyzed changes in Collembola community structure along a gradient of forest fragmentation, testing predictions of the Hierarchical Patch Dynamics Paradigm (HPDP) in different European biogeographic regions (Boreal, Continental, Atlantic, Mediterranean, Alpine). Using variance partitioning methods, based on partial CCAs, we observed that the independent effect of environmental processes was significantly explaining Collembola community variance in all regions, while the relative effect of spatial variables was not significant, due to the observed high levels of landscape heterogeneity along the gradient. Environmental factors at the patch and plot scales were generally significant and explained the larger part of community changes. Landscape variables were not significant across all study sites. Yet, at the landscape level, an increase in forest habitat and proximity of forest patches were showed to have an indirect influence on local community changes, by influencing microhabitat heterogeneity at lower spatial scales in all studied regions. In line with HPDP, large-scale landscape features influenced spatio-temporal changes in soil fauna communities by constraining small-scale environmental processes. In turn, these provided mechanistic understanding for diversity patterns operating at the patch scale, via shifts in community weighted mean of Collembola life-forms occurring in local communities along the fragmentation gradient.     

Landscape connectivity and animal behavior: functional grain as a key determinant for dispersal

Landscape connectivity can be viewed from two perspectives that could be considered as extremes of a gradient: functional connectivity (refers to how the behavior of a dispersing organism is affected by landscape structure and elements) and structural connectivity (depends on the spatial configuration of habitat patches in the landscape like vicinity or presence of barriers). Here we argue that dispersal behavior changes with landscape configuration stressing the evolutionary dimension that has often been ignored in landscape ecology. Our working hypothesis is that the functional grain of resource patches in the landscape is a crucial factor shaping individual movements, and therefore influencing landscape connectivity. Such changes are likely to occur on the short-term (some generations). We review empirical studies comparing dispersal behavior in landscapes differing in their fragmentation level, i.e., with variable resource grain. We show that behavioral variation affecting each of the three stages of the dispersal process (emigration, displacement or transfer in the matrix, and immigration) is indeed likely to occur according to selective pressures resulting from changes in the grain of the landscape (mortality or deferred costs). Accordingly, landscape connectivity results from the interaction between the dispersal behavior of individuals and the grain of each particular landscape. The existence of this interaction requires that connectivity estimates (being based on individual-based models, least cost distance algorithms, and structural connectivity metrics or even Euclidian distance) should be carefully evaluated for their applicability with respect to the required level of precision in species-specific and landscape information.     

Streambed landscapes: evidence that stream invertebrates respond to the type and spatial arrangement of patches

The availability and spatial arrangement of habitat patches are known to strongly influence fauna in terrestrial ecosystems. The importance of patch arrangement is not well-studied within running-water systems where flow-induced movements of patches and of fauna could decouple habitat characteristics and faunal habitat preferences. Using small, stream-dwelling invertebrates, we asked if fauna in such systems can distinguish among patch types and if patch arrangement at their `landscape scale' (i.e., within a streambed across which they move and forage) can be linked to faunal abundance. We quantified the spatial distribution of sand and leaf patches at multiple sites on a streambed at regular intervals over a 1 yr period, estimated faunal abundance in the two patch types, and experimentally determined if faunal colonization varied among leaf patches that were similar structurally but differed in their potential microbial food resources. We show that despite their small size and limited swimming abilities, these stream invertebrates did respond to patch type, that specific characteristics of an individual patch influenced faunal colonization, and that the spatial arrangement of patches on the streambed was linked to field abundances. Larval chironomids and adult copepods were more abundant in leaves than in sand and preferentially colonized leaf patches made with rapidly decomposing leaves that harbored higher microbial (bacteria and fungi) abundances over leaf patches with more refractory leaves and lower microbial abundances. Further, statistical models that included spatially-explicit data on patch arrangement (e.g., patch contagion, distance between patches) explained significantly more variation in faunal abundance, than models that included only nonspatial information (e.g., date, time since last flood). Despite the fact that these fauna live in a highly dynamic environment with variable flow rates during the year, unstable patch configurations, and seasonal changes in total abundance, our findings suggest a need for aquatic ecologists to test the hypothesis that small-scale landscape attributes within streams (e.g., leaf patch aggregation) may be important to faunal dynamics. If patch aggregation has negative consequences for stream biota, streambed `landscapes' may be fundamentally different from many terrestrial landscapes due to the inherent connectivity provided by the water and the over-riding importance of patch edges. Regardless of these differences, our findings suggest that the spatial configuration of patches in a landscape may have consequences for fauna even in highly dynamic systems, in which patches move and fauna periodically experience high levels of passive dispersal.