Gene flow and functional connectivity in the natterjack toad

Functional connectivity is a key factor for the persistence of many specialist species in fragmented landscapes. However, connectivity estimates have rarely been validated by the observation of dispersal movements. In this study, we estimated functional connectivity of a real landscape by modelling dispersal for the endangered natterjack toad (Bufo calamita) using cost distance. Cost distance allows the evaluation of 'effective distances', which are distances corrected for the costs involved in moving between habitat patches in spatially explicit landscapes. We parameterized cost-distance models using the results of our previous experimental investigation of natterjack's movement behaviour. These model predictions (connectivity estimates from the GIS study) were then confronted to genetic-based dispersal rates between natterjack populations in the same landscape using Mantel tests. Dispersal rates between the populations were inferred from variation at six microsatellite loci. Based on these results, we conclude that matrix structure has a strong effect on dispersal rates. Moreover, we found that cost distances generated by habitat preferences explained dispersal rates better than did the Euclidian distances, or the connectivity estimate based on patch-specific resistances (patch viscosity). This study is a clear example of how landscape genetics can validate operational functional connectivity estimates.     

Habitat quality of source patches and connectivity in fragmented landscapes

Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.     

Metacommunities of spiders in grassland habitat fragments of an agricultural landscape

Arthropod communities in fragmented agricultural landscapes depend on local processes and the interactions between communities in the habitat islands. We aimed to study metacommunity structure of spiders, a group that is known for high dispersal power, local niche partitioning and for engaging in species interactions. While living in fragmented habitats could lead to nestedness, other ecological traits of spiders might equally lead to patterns dominated either by species interactions or habitat filtering. We asked, which community pattern will prevail in a typical agricultural landscape with isolated patches of semi-natural habitats. Such a situation was studied by sampling spiders in 28 grassland locations in a Hungarian agricultural landscape. We used the elements of metacommunity structure (EMS) framework to distinguish between alternative patterns that reveal community organization. The EMS analysis indicated coherent species ranges, high turnover and boundary clumping, suggesting Clementsian community organization. The greatest variation in species composition was explained by local habitat characteristics, indicating habitat filtering. The influence of dispersal could be detected by the significant effect of landscape composition, which was strongest at 500 m. We conclude that dispersal allows spiders to respond coherently to the environment, creating similar communities in similar habitats. Consistent habitat differences, such as species rich versus species poor vegetation, lead to recognisably different, recurrent communities. These characteristics make spiders a predictable and diverse source of natural enemies in agricultural landscapes. Sensitivity to habitat composition at medium distances warns us that landscape homogenization may alter these metacommunity processes.     

Landscape matrix modifies richness of plants and insects in grassland fragments

There is an increasing awareness that not only area and isolation, but also the characteristics of the landscape surrounding habitat patches influence population persistence and species diversity in fragmented landscapes. In this study, we examine the effects of grassland fragmentation and land use in the landscape matrix (on a 2 km scale) on species richness of plants, butterflies, bees and hoverflies. These organisms were studied in replicated remnant patches of different sizes and isolation, embedded in landscapes dominated either by forest, arable land or a mix of these. We found positive effects of patch area on species richness of the three insect taxa, but not of plants. Isolation had a negative effect only on hoverflies. Matrix type had contrasting effects on the studied taxa. Species richness of plants and butterflies was lowest in patches in landscapes dominated by arable land and highest in forest‐dominated landscapes. For hoverflies, the negative effect of small patch area was strongest in forest‐dominated landscapes, and there was a similar non‐significant trend for bees. Our study shows the importance of considering matrix characteristics when studying responses to habitat fragmentation. Differences in matrix response among organism groups probably impinge on differing mechanisms. A forest matrix is likely to provide additional resources for butterflies but either constitute a barrier to dispersal or deprive resources as compared to an arable matrix for hoverflies. Enhanced plant diversity in grassland patches embedded in forested landscapes can be explained by habitat generalists more easily invading these patches, or by an unpaid extinction debt in these landscapes.     

Patch occupancy and abundance of local populations in landscapes differing in degree of habitat fragmentation: a case study of the colonial black‐headed gull,Chroicocephalus ridibundus

Aim This study investigated whether habitat fragmentation at the landscape level influences patch occupancy and abundance of the black‐headed gull, Chroicocephalus ridibundus, and whether the response of the species to environmental factors is consistent across replicated landscape plots. Location Water bodies (habitat patches) in southern Poland. Methods Surveys were conducted in two landscape types (four plots in each): (1) more‐fragmented landscape, in which habitat patches were small (mean size 2.2–6.2 ha) and far apart (mean distance 2.5–3.1 km); and (2) less‐fragmented landscape, in which habitat patches were large (mean size 9.2–16.5 ha) and separated by short distances (mean 0.9–1.4 km). Observations were performed twice in 284 potential habitat patches during the 2007 breeding season. Results Colonies were significantly more frequent and larger in the less‐fragmented landscapes than in the more‐fragmented ones. Probability of patch occupancy and number of breeding birds were positively related with patch size and these relationships were especially strong in the more‐fragmented landscapes. In the less‐fragmented landscapes, the occurrence of black‐headed gulls was negatively related to the distance to the nearest local population, but in the more‐fragmented landscapes such a relationship was not detected. As distance to the nearest habitat patch increased, the probability of the patch occupancy decreased in the more‐fragmented landscapes. Moreover, abundance was negatively influenced by distance to the nearest habitat patch, especially strongly in more‐fragmented landscapes. Proximity of corridors (rivers) positively influenced the occupation of patches regardless of landscape type. The number of islets positively influenced occupancy and abundance of local populations, and this relationship was stronger in the more‐fragmented landscapes. Main conclusions Our results are in agreement with predictions from metapopulation theory and are the first evidence that populations of black‐headed gulls may have a metapopulation structure. However, patch occupancy and abundance were differentially affected by explanatory variables in the more‐fragmented landscapes than in the less‐fragmented ones. This implies that it is impossible to derive, a priori, predictions about presence/abundance patterns based on only a single landscape.     

The genetic structure of populations of Metrioptera bicolor in a spatially structured landscape: effects of dispersal barriers and geographic distance

The stability and long-term survival of animal populations in fragmented landscapes largely depends on the colonisation of habitat patches and the exchange of individuals between patches. The degree of inter-patch dispersal, in turn, depends on the dispersal abilities of species and the landscape structure (i.e. the nature of the landscape matrix and habitat distribution). Here, we investigated the genetic structure of populations of Metrioptera bicolor, a wing-dimorphic bush cricket, in a spatially structured landscape with patches of suitable habitat distributed within a diverse matrix of different habitat types. Using six microsatellite markers, we assessed the effects of geographic distance and different matrix types on the extent of genetic differentiation among 24 sampling sites. We found that forest and a river running through the study area both impede inter-patch dispersal. The presence of these two matrix types was positively correlated with the extent of genetic differentiation between sites. In addition, we found a significant positive correlation between pairwise genetic and geographic distances for a subsample of sites which were separated only by arable land or settlements. For the complete data set, this correlation could not be found. This is most probably because the adverse effect of forest and river on gene flow dominates the effect of geographic distance in our limited set of patches. Our analyses clearly emphasize the differential resistance of different habitat types on dispersal and the importance of a more detailed view on matrix “quality” in metapopulation studies.     

Anthropogenic landscape change promotes asymmetric dispersal and limits regional patch occupancy in a spatially structured bird population

1. Local extinctions in habitat patches and asymmetric dispersal between patches are key processes structuring animal populations in heterogeneous environments. Effective landscape conservation requires an understanding of how habitat loss and fragmentation influence demographic processes within populations and movement between populations. 2. We used patch occupancy surveys and molecular data for a rainforest bird, the logrunner (Orthonyx temminckii), to determine (i) the effects of landscape change and patch structure on local extinction; (ii) the asymmetry of emigration and immigration rates; (iii) the relative influence of local and between-population landscapes on asymmetric emigration and immigration; and (iv) the relative contributions of habitat loss and habitat fragmentation to asymmetric emigration and immigration. 3. Whether or not a patch was occupied by logrunners was primarily determined by the isolation of that patch. After controlling for patch isolation, patch occupancy declined in landscapes experiencing high levels of rainforest loss over the last 100 years. Habitat loss and fragmentation over the last century was more important than the current pattern of patch isolation alone, which suggested that immigration from neighbouring patches was unable to prevent local extinction in highly modified landscapes. 4. We discovered that dispersal between logrunner populations is highly asymmetric. Emigration rates were 39% lower when local landscapes were fragmented, but emigration was not limited by the structure of the between-population landscapes. In contrast, immigration was 37% greater when local landscapes were fragmented and was lower when the between-population landscapes were fragmented. Rainforest fragmentation influenced asymmetric dispersal to a greater extent than did rainforest loss, and a 60% reduction in mean patch area was capable of switching a population from being a net exporter to a net importer of dispersing logrunners. 5. The synergistic effects of landscape change on species occurrence and asymmetric dispersal have important implications for conservation. Conservation measures that maintain large patch sizes in the landscape may promote asymmetric dispersal from intact to fragmented landscapes and allow rainforest bird populations to persist in fragmented and degraded landscapes. These sink populations could form the kernel of source populations given sufficient habitat restoration. However, the success of this rescue effect will depend on the quality of the between-population landscapes.     

Isolation determines patterns of species presence in highly fragmented landscapes

In fragmented landscapes, changes in habitat availability, patch size, shape and isolation may affect survival of local populations. Proposing efficient conservation strategies for such species relies initially on distinguishing the particular effects of those factors. To address these issues, we investigated the occurrence of 3 bird species in fragmented Brazilian Atlantic Forest landscapes. Playback techniques were used to collect presence/absence data of these species inside 80 forest patches, and incidence models were used to infer their occupancy pattern from landscape spatial structure. The relative importance of patch size, shape and surrounding forest cover and isolation was assessed using a model selection approach based on maximum likelihood estimation. The presence of all species was in general positively affected by the amount of surrounding habitat and negatively affected by inter‐patch distances. The joint effects of patch size and the surrounding landscape characteristics were important determinants of occupancy for two species. The third species was affected only by forest cover and mean patch isolation. Our results suggest that local species presence is in general more influenced by the isolation from surrounding forests than by patch size alone. We found evidence that, in highly fragmented landscapes, birds that can not find patches large enough to settle may be able to overcome short distances through the matrix and include several nearby patches within their home‐ranges to complement their resource needs. In these cases, patches must be defined as functionally connected habitat networks rather than mere continuous forest segments. Bird conservation strategies in the Atlantic forest should focus on increasing patch density and connectivity, in order to implement forest networks that reduce the functional isolation between large remnants with remaining core habitat.     

Effect of habitat fragmentation on dispersal in the butterfly Proclossiana eunomia

Comparison of dispersal rates of the bog fritillary butterfly between continuous and fragmented landscapes indicates that between patch dispersal is significantly lower in the fragmented landscape, while population densities are of the same order of magnitude. Analyses of the dynamics of the suitable habitat for the butterfly in the fragmented landscape reveal a severe, non linear increase in spatial isolation of patches over a time period of 30 years (i.e. 30 butterfly generations), but simulations of the butterfly metapopulation dynamics using a structured population model show that the lower dispersal rates in the fragmented landscape are far above the critical threshold leading to metapopulation extinction. These results indicate that changes in individual behaviour leading to the decrease of dispersal rates in the fragmented landscape were rapidly selected for when patch spatial isolation increased. The evidence of such an adaptive answer to habitat fragmentation suggests that dispersal mortality is a key factor for metapopulation persistence in fragmented landscapes. We emphasise that landscape spatial configuration and patch isolation have to be taken into account in the debate about large-scale conservation strategies.     

Landscape connectivity of the grassland biome in Mpumalanga,South Africa

The South African grassland biome is one of the most threatened biomes in South Africa. Approximately 45% of the grassland biome area is transformed, degraded or severely invaded by alien plants and the remaining natural areas are highly fragmented. In this fragmented landscape, the connectivity between habitat patches is very important to maintain viable populations. In this study we aimed to quantify connectivity of the grassland biome in Mpumalanga using graph theory in order to identify conservation priorities and to direct conservation efforts. Graph theory‐based connectivity indices have the ability to combine spatially explicit habitat data with species specific dispersal data and can quantify structural and functional connectivity over large landscapes. We used these indices to quantify the overall connectivity of the study area, to determine the influence of abandoned croplands on overall connectivity, and to identify the habitat patches and vegetation types most in need of maintaining overall connectivity. Natural areas were identified using 2008 land cover data for Mpumalanga. Connectivity within the grassland biome of Mpumalanga was analysed for grassland species with dispersal distances ranging from 50 to 1000 m. The grassland habitat patches were mostly well connected, with 99.6% of the total habitat area connected in a single component at a threshold distance of 1000 m. The inclusion of abandoned croplands resulted in a 33% increase in connectivity at a threshold distance of 500 m. The habitat patches most important for maintaining overall connectivity were the large patches of continuous habitat in the upper and lower centres of the study area and the most important vegetation types were the Wakkerstroom Montane Grassland and the Eastern Temperate Freshwater Wetlands. These results can be used to inform management decisions and reserve design to improve and maintain connectivity in this biome.     

Effects of landscape connectivity on the spatial distribution of insect diversity in agricultural mosaic landscapes

European agricultural landscapes are mosaics of intensively cultivated areas and semi-natural elements. Although comprising only a small fraction of the total area, semi-natural elements provide habitat for most of the landscape biodiversity. Agricultural intensification has increasingly fragmented semi-natural elements and species numbers are in decline. Insights into the effects of landscape structure on species’ distributions within and among semi-natural habitats are needed to conserve biodiversity in agricultural landscapes more effectively. We investigated the landscape- and habitat-specific diversity partitions of wild bees, true bugs, and carabid beetles in two differently structured agricultural landscapes in Switzerland. In each landscape, we partitioned the total species diversity (γ) into its additive components within (_P) and among patches (β_P) and among habitats (β_H). In the landscape characterized by a patchy, isolated distribution of habitat elements, among-patch diversity (β_P) explained 44% of the total species richness (γ) and was significantly higher than expected under a random distribution of samples among habitat patches; in the landscape with higher habitat connectivity, among-patch diversity (β_P) comprised 32% of the total species richness (γ) and did not differ from the random expectation. Habitat-specific within-patch contributions to species richness were similarly low across habitat types (_P=23–24%) in the patchy landscape, whereas in the more connected landscape within-patch partitions tended to be higher and differed among habitat types (_P=22–38%). Functionally different groups of bees, true bugs, and carabids also responded differently to landscape structure in a manner that was consistent with known differences in resource specialization and dispersal ability. Differences in diversity partitions among landscapes and taxa indicate the need for flexible conservation strategies. Conservation of habitat-specific diversity may require more habitat patches in landscapes that have lower habitat connectivity and low within-patch diversity (_P) than in landscapes with higher within-patch diversity (_P).     

Experimental evidence for extreme dispersal limitation in tropical forest birds

Movements of organisms between habitat remnants can affect metapopulation structure, community assembly dynamics, gene flow and conservation strategy. In the tropical landscapes that support the majority of global biodiversity and where forest fragmentation is accelerating, there is particular urgency to understand how dispersal across habitats mediates the demography, distribution and differentiation of organisms. By employing unique dispersal challenge experiments coupled with exhaustive inventories of birds in a Panamanian lacustrine archipelago, we show that the ability to fly even short distances (< 100 m) between habitat fragments varies dramatically and consistently among species of forest birds, and that this variation correlates strongly with species' extinction histories and current distributions across the archipelago. This extreme variation in flight capability indicates that species' persistence in isolated forest remnants will be differentially mediated by their respective dispersal abilities, and that corridors connecting such fragments will be essential for the maintenance of avian diversity in fragmented tropical landscapes.     

Habitat use and dispersal characteristic by <Emphasis Type="Italic">Stethophyma grossum</Emphasis>: the role of habitat isolation and stable habitat conditions towards low dispersal

Species in a highly fragmented environment, such as the intensively used agricultural landscapes of Europe, are expected to be in danger of extinction. We hypothesize according to Kisdi’s theory (Am Nat 159:579–596, 2002) that species in fragmented landscapes with isolated habitats in general tend to possess low dispersal. In order to verify this hypothesis we studied the movement patterns of Stethophyma grossum, a hygrophilous species of wetlands, by mark–release–recapture techniques in a landscape with scattered suitable habitats over 3 years. The study focused on the major population in this landscape (site #1) as dispersal behaviour was assumed to be greatest. Actually, marked individuals of S. grossum were never found in any further suitable habitats in close vicinity to site #1. Despite that the peatland meadow of study site #1 was all over covered with homogenous vegetation only 6% (1.8 ha) of the whole area (30 ha) were occupied by S. grossum. The mean recapture rate over 3 years amounted to 39% with no significant differences between males and females. Both covered little distances within their mean range size of 1.8 ha; the median distances were 36.91 m for males and 26.65 m for females. We confirm the hypothesis that sub-populations of species in longstanding naturally isolated habitats, which habitat conditions have been stable; evolved low dispersal with little movements which are routine movements to find mating partners or food.     

Condition-dependent dispersal of a patchily distributed riparian ground beetle in response to disturbance

In common with many habitat elements of riverine landscapes, exposed riverine sediments (ERS) are highly disturbed, naturally patchy and regularly distributed, whose specialists are strongly adapted to flood disturbance and loss of habitat due to succession. Investigations of dispersal in ERS habitats therefore provide an important contrast to the unnaturally fragmented, stable systems usually studied. The present investigation analysed the three interdependent stages of dispersal: (1) emigration, (2) inter-patch movement and (3) immigration of a common ERS specialised beetle, Bembidion atrocaeruleum (Stephens 1828) (Coleoptera, Carabidae), in a relatively unmodified section of river, using mark–resight methods. Dispersal was correlated with estimates of local population size and density, water level and patch quality in order to test for condition-dependent dispersal cues. Flood inundation of habitat was found to increase strongly the overall rate of dispersal, and the rate of emigration was significantly higher from patches that were heavily trampled by cattle. Strongly declining numbers of dispersers with distance suggested low dispersal rates during periods of low water level. Dispersal in response to habitat degradation by cattle trampling would likely lead to a higher overall population fitness than a random dispersal strategy. Dispersal distances were probably adapted to the underlying habitat landscape distribution, high-flow dispersal cues and ready means of long-distance dispersal through hydrochory. Species whose dispersal is adapted to the natural habitat distribution of riverine landscapes are likely to be strongly negatively affected by reduced flood frequency and intensity and habitat fragmentation through flow regulation or channelisation.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Annual Survival and Turnover Rates of an Afrotropical Robin in a Fragmented Forest

Whether or not subdivided populations persist in fragmented landscapes primarily depends on how well individuals can survive within discrete habitat patches. Using data from six capture–recapture sessions, survival probabilities of the white-starred robin were estimated in seven indigenous forest patches in the highly fragmented Taita Hills forests, SE Kenya. We found no significant differences in survival probability either among fragment-size categories (large 135 ha, medium 95 ha, and small 2–8 ha) or between adult and first-year birds. However, males had a higher probability of survival from one year to the next than females. Turnover rates of adult birds were higher for females than males, but also higher in the medium and small patches than in the large one within each sex. That survival probability was similar among fragments, but turnover rates differed denoted that different processes caused extirpation from the patches. We suggest that mortality associated with dispersal was probably a more important cause of extirpation than within-patch mortality in the largest habitat, which had the lowest turnover rates. Conversely, high within-patch mortality, for instance due to predation during incubation, could have been more important in the smaller, more disturbed habitats. These results lend support to the proposition that avian conservation efforts should be focussed both at the landscape level to improve connectivity between fragments and reduce mortality during dispersal, and at the patch level to exclude other mortality sources such as nest predation.     

Unravelling seed dispersal through fragmented landscapes: Frugivore species operate unevenly as mobile links

Seed dispersal constitutes a pivotal process in an increasingly fragmented world, promoting population connectivity, colonization and range shifts in plants. Unveiling how multiple frugivore species disperse seeds through fragmented landscapes, operating as mobile links, has remained elusive owing to methodological constraints for monitoring seed dispersal events. We combine for the first time DNA barcoding and DNA microsatellites to identify, respectively, the frugivore species and the source trees of animal‐dispersed seeds in forest and matrix of a fragmented landscape. We found a high functional complementarity among frugivores in terms of seed deposition at different habitats (forest vs. matrix), perches (isolated trees vs. electricity pylons) and matrix sectors (close vs. far from the forest edge), cross‐habitat seed fluxes, dispersal distances and canopy‐cover dependency. Seed rain at the landscape‐scale, from forest to distant matrix sectors, was characterized by turnovers in the contribution of frugivores and source‐tree habitats: open‐habitat frugivores replaced forest‐dependent frugivores, whereas matrix trees replaced forest trees. As a result of such turnovers, the magnitude of seed rain was evenly distributed between habitats and landscape sectors. We thus uncover key mechanisms behind “biodiversity–ecosystem function” relationships, in this case, the relationship between frugivore diversity and landscape‐scale seed dispersal. Our results reveal the importance of open‐habitat frugivores, isolated fruiting trees and anthropogenic perching sites (infrastructures) in generating seed dispersal events far from the remnant forest, highlighting their potential to drive regeneration dynamics through the matrix. This study helps to broaden the “mobile‐link” concept in seed dispersal studies by providing a comprehensive and integrative view of the way in which multiple frugivore species disseminate seeds through real‐world landscapes.     

Influence of Patch Factors and Connectivity on the Avifauna of Fragmented Polylepis Forest in the Ecuadorian Andes

Human‐induced alteration of habitat is a major threat to biodiversity worldwide, especially in areas of high biological diversity and endemism. Polylepis (Rosaceae) forest, a unique forest habitat in the high Andes of South America, presently occurs as small and isolated patches in grassland dominated landscapes. We examine how the avian community is likely influenced by patch characteristics (i.e., area, plant species composition) and connectivity in a landscape composed of patches of Polylepis forest surrounded by páramo grasslands in Cajas National Park in the Andes of southern Ecuador. We used generalized linear mixed models and an information‐theoretic approach to identify the most important variables probably influencing birds inhabiting 26 forest patches. Our results indicated that species richness was associated with area of a patch and floristic composition, particularly the presence of Gynoxys (Asteraceae). However, connectivity of patches probably influenced the abundance of forest and generalists species. Elsewhere, it has been proposed that effective management plans for birds using Polylepis should promote the conservation of mature Polylepis patches. Our results not only suggest this but also show that there are additional factors, such as the presence of Gynoxys plants, which will probably play a role in conservation of birds. More generally, these findings show that while easily measured attributes of the patch and landscape may provide some insights into what influences patch use by birds, knowledge of other factors, such as plant species composition, is essential for better understanding the distribution of birds in fragmented landscapes.     

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.     

Movement patterns of Rhyssomatus lineaticollis Say (Coleoptera: Curculionidae) within and among Asclepias syriaca (Asclepiadaceae) patches in a fragmented landscape

Abstract.  1. Dispersal capabilities of organisms are critical in determining the landscape population structure of species as well as their likelihood of survival in fragmented landscapes. Using mark–recapture techniques on the monophagous weevil Rhyssomatus lineaticollis Say (Curculionidae), within- and between-patch dispersal capabilities, landscape level population structure, and the role of beetle density and host patch characteristics in setting distances, amounts, and timing of dispersal were studied.
2. The data indicate that R. lineaticollis is sedentary, with 50% of recaptured beetles moving < 1 m and the maximum distance moved < 1 km. Within- and between-patch movement of beetles was unrelated to host plant patch characteristics and beetle densities.
3. Despite limited dispersal, R. lineaticollis probably functions as a patchy population in east-central Iowa, U.S.A. because dispersals between patches are common and because all host patches surveyed contained this herbivore, indicating a lack of suitable vacant patches, a prerequisite for metapopulation structure.
4. Between-patch distances are well within the dispersal capabilities of R. lineaticollis , although this may be the result of an increase in the density of patches of its host, Asclepias syriaca , in the landscape over the last 150 years as a result of human disturbance and this species' weedy habit.
5. Metapopulation structure in monophagous prairie herbivores may be most likely in species whose non-weedy host plants form highly predictable resources in space and time, but which are now widely scattered in habitat fragments.     

Landscape connectivity and the role of small habitat patches as stepping stones: an assessment of the grassland biome in South America

Connectivity losses lead to a reduction of the amount of habitat resources that can be reached and used by species, and hence to a decline in the ranges and abundance of multiple taxa. Despite the recognized important role of small habitat patches for many species inhabiting fragmented landscapes, their potential contribution as stepping stones for maintaining overall landscape connectivity has received less attention. Using connectivity metrics based on a graph-theoretic approach we (i) quantified the connectivity of grassland patches in a sector of the Pampa region in Argentina, using a range of dispersal distances (from 100 to 10,000 m) representative of the scale of dispersal of different species; (ii) identified the most relevant patches for maintaining overall connectivity; and (iii) studied the importance of small patches (defined for different area thresholds of 5, 20, and 50 ha) as connectivity providers in the landscape. Although grassland patches were in general poorly connected at all distances, some of them were critical for overall connectivity and were found to play different crucial roles in the patch network. The location of small patches in the grassland network allowed them to function as stepping stones, yielding significant connectivity gains for species that move large distances (>5000 m) for the three area thresholds considered. Thus, under the spatial pattern of the studied landscape, species that move long distances would benefit from stepping stones, while less mobile organisms would benefit from, and mostly rely on the largest patches. We recommend that future management activities should (i) aim at preserving the grassland patches with the highest potential as stepping stones to promote landscape-level connectivity; and (ii) pay more attention to the conservation of key small patches, particularly given that usually they are those more vulnerable to land clearing for agriculture.