Current velocity and habitat patchiness shape stream herbivore movement

Animal movements are influenced by the structure and arrangement of patches in a landscape. Most movement studies occur in terrestrial landscapes, though aquatic landscapes are equally heterogeneous and feature patches that differ in resistance to animal movements. Furthermore, the variable and highly directional flow of water over streambed landscapes is a unique environmental element, yet its constraint on animal movement is poorly understood. This study examines how habitat availability in a streambed landscape interacts with current velocity to affect movement patterns of two benthic grazers: glossosomatid caddisfly larvae (Agapetus boulderensis) and pulmonate snails (Physa sp.). Using experimental streambed landscapes, we found that Agapetus traveled farther as availability of smooth habitat (composed of low diatom turfs) increased compared to tall, structured filamentous stands, but only did so in slow current velocities. Swifter flows caused restricted movement of Agapetus and more upstream‐oriented paths, but only in smooth landscapes where the potential for flow refugia from filamentous stands was minimal. Similarly, increasing proportions of smooth habitat facilitated greater net displacement of Physa using more upstream‐oriented paths. Higher current velocities caused Physa to move faster, a pattern demonstrated only in smooth landscapes. Our results illustrate a strong interaction between benthic habitat structure and current velocity in shaping patterns of grazer movements in a streambed landscape. Our study also suggests that the flow of water be considered not only a strong environmental gradient in streams, but also an interactive landscape feature that can combine with streambed structure to determine the permeability of patches to the movement of benthic organisms. Landscape ecology has mainly focused on terrestrial environments, and this study offers insight into some of the unique processes that may shape animal movement in aquatic environments.     

道路对林地景观连接度的影响——以巩义市为例

基于景观连接度原理,借用景观连接度指数,在地理信息系统支持下,探讨了巩义市山区林地景观在不同距离阈值下连接度的变化,定量分析了道路对林地景观连接度的影响。结果显示,随着景观距离阈值的增大,无论是否有道路,林地景观整体可能连通性指数值都表现为逐渐增大;对林地景观连接度起"非常高"和"高"作用的林地斑块数量都比较少,但占林地总面积比例较大,面积大的林地斑块在提高景观连接度中起的作用较大;道路的分割使得林地斑块重要值降低,就单一斑块而言,随着景观距离阈值的增大,分割成的小斑块的重要值降低程度在逐渐减小。     

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.     

Biodiversity extinction thresholds are modulated by matrix type

Biodiversity extinction thresholds are abrupt declines in biological diversity that occur with habitat loss, associated with a decline in habitat connectivity. Matrix quality should influence the location of thresholds along habitat loss gradients through its effects on connectivity; however these relationships have seldom been explored empirically. Using field data from 23 independent 1254 ha landscapes in the Brazilian Atlantic Forest, we evaluated how tropical avian biodiversity responds to native forest loss within habitat patches embedded either in homogeneous pasture matrix context (with a high proportion of cattle pastures), and heterogeneous coffee matrix context (with high abundance of sun coffee plantations). We considered taxonomic, functional, and phylogenetic diversity, and tested if matrix type and choice of diversity metric influenced the location of biodiversity thresholds along the forest cover gradient. We found that matrix type postponed the abrupt loss of taxonomic diversity, from a threshold of 35% of forest cover in homogeneous pasture matrix to 19% in heterogeneous coffee matrix. Phylogenetic diversity responded similarly, with thresholds at 30 and 24% in homogeneous‐pasture and heterogeneous‐coffee matrices, respectively, but no relationship with forest cover was detected when corrected for richness correlation. Despite the absence of a threshold for functional diversity in either matrix types, a strong decline below 20% of habitat amount was detected. Finally, below 20% native habitat loss, all diversity indices demonstrated abrupt declines, indicating that even higher‐quality matrices cannot postpone diversity loss below this critical threshold. These results highlight that taxonomic diversity is a more sensitive index of biodiversity loss in fragmented landscapes, which may be used as a benchmark to prevent subsequent functional and phylogenetic losses. Furthermore, increasing matrix quality appears an efficient conservation strategy to maintain higher biodiversity levels in fragmented landscapes over a larger range of habitat loss.     

基于生境斑块的滇金丝猴景观连接度分析

基于生境斑块,结合最小费用距离并运用图论法对滇金丝猴分布区进行栖息地连接度分析,研究利用猴群的现实分布结合Logistic回归模型确定了景观功能连接的最佳距离阈值,对于功能畅通的组分,以景观指数BC定量识别出作为"踏脚石"的优先保护区域;对于功能不连接的组分,绘制出最小费用路径,确定了该路径中优先恢复区域。结果表明:最佳的最小费用距离阈值为1400,该阈值下猴群主要存在于5个组分中,所有组分中猴群间的连接度优劣排序为组分3组分1组分5组分4,龙马山猴群(G15)没有"踏脚石"斑块使其与同一组分内的其他猴群相连接,应考虑优先恢复该区域的植被,研究成果对于该物种的保护和其他濒危物种的类似研究具有较强的参考价值和借鉴意义。     

Landscape geometry determines community response to disturbance

Ecological communities are impacted by anthropogenic changes in both habitat geometry (i.e. amount, shape, fragmentation and connectivity of habitat) and disturbance regime. Although the effect of each of these drivers on diversity is well-documented, few studies have considered how habitat geometry and disturbance interact to affect diversity. We used a miniature landscape of moss patches to experimentally manipulate both habitat geometry and disturbance frequency on microarthropod communities. Species richness and abundance in local patches declined linearly with disturbance rate in all experimental landscapes, but the speed of this decline (a measure of ecological resilience) depended on the size and connectivity of the surrounding region. Reductions in region size had little effect on community resilience to disturbance until habitat loss resulted in complete loss of connectivity between patches, suggesting a threshold in community response to habitat loss. Beyond this threshold, repeated disturbance resulted in rapid declines in patch species richness and abundance and substantial changes in community composition. These effects of habitat geometry and disturbance on diversity were scale-dependent. Gamma (regional-scale) diversity was unaffected by habitat geometry, suggesting experimental reductions in alpha (local-scale) diversity were offset by increases in beta diversity. There was no effect of body size, abundance, or trophic position in determining species response to disturbance. Taxonomic grouping had a weak effect, with oribatids less affected by drought. We conclude that, in this system, dispersal from the surrounding metacommunity is vital in allowing recovery of local communities from disturbance. When habitat loss and fragmentation disrupt this process, extinctions result. Studies that examine separately the effects of habitat alterations and disturbance on diversity may therefore underestimate their combined effects.     

The taste of nectar – a neglected area of pollination ecology

We investigated the impact of landscape structure on landscape connectivity using a combination of simulation and empirical experiments. In a previous study we documented the movement behaviour of a specialized goldenrod beetle ( Trirhabda borealis Blake) in three kinds of patches: habitat (goldenrod) patches and two types of matrix patch (cut vegetation and cut vegetation containing camouflage netting as an impediment to movement). In the current study, we used this information to construct simulation and experimental landscapes consisting of mosaics of these three patch types, to study the effect of landscape structure on landscape connectivity, using the T. borealis beetle as a model system. In the simulation studies, landscape connectivity was based on movements of individual beetles, and was measured in six different ways. The simulations revealed that the six measures of landscape connectivity were influenced by different aspects of landscape structure, suggesting that: (1) landscape connectivity is a poorly defined concept, and (2) the same landscape may have different landscape connectivity values when different measures of landscape connectivity are used. There were two general predictions that held over all measures of landscape connectivity: (1) increasing interpatch distance significantly decreased landscape connectivity and (2) the influence of matrix elements on landscape connectivity was small in comparison to the influence of habitat elements. Empirical mark-release-resight experiments using Trirhabda beetles in experimental landscapes supported the simulation results.     

Mapping functional connectivity

An objective and reliable assessment of wildlife movement is important in theoretical and applied ecology. The identification and mapping of landscape elements that may enhance functional connectivity is usually a subjective process based on visual interpretations of species movement patterns. New methods based on mathematical morphology provide a generic, flexible, and automated approach for the definition of indicators based on the classification and mapping of spatial patterns of connectivity from observed or simulated movement and dispersal events. The approach is illustrated with data derived from simulated movement on a map produced from satellite imagery of a structurally complex, multi-habitat landscape. The analysis reveals critical areas that facilitate the movement of dispersers among habitat patches. Mathematical morphology can be applied to any movement map providing new insights into pattern-process linkages in multi-habitat landscapes.     

Experimental beetle metapopulations respond positively to dynamic landscapes and reduced connectivity

Interactive effects of multiple environmental factors on metapopulation dynamics have received scant attention. We designed a laboratory study to test hypotheses regarding interactive effects of factors affecting the metapopulation dynamics of red flour beetle, Tribolium castaneum. Within a four-patch landscape we modified resource level (constant and diminishing), patch connectivity (high and low) and patch configuration (static and dynamic) to conduct a 2(3) factorial experiment, consisting of 8 metapopulations, each with 3 replicates. For comparison, two control populations consisting of isolated and static subpopulations were provided with resources at constant or diminishing levels. Longitudinal data from 22 tri-weekly counts of beetle abundance were analyzed using bayesian Poisson generalized linear mixed models to estimate additive and interactive effects of factors affecting abundance. Constant resource levels, low connectivity and dynamic patches yielded greater levels of adult beetle abundance. For a given resource level, frequency of colonization exceeded extinction in landscapes with dynamic patches when connectivity was low, thereby promoting greater patch occupancy. Negative density dependence of pupae on adults occurred and was stronger in landscapes with low connectivity and constant resources; these metapopulations also demonstrated greatest stability. Metapopulations in control landscapes went extinct quickly, denoting lower persistence than comparable landscapes with low connectivity. When landscape carrying capacity was constant, habitat destruction coupled with low connectivity created asynchronous local dynamics and refugia within which cannibalism of pupae was reduced. Increasing connectivity may be counter-productive and habitat destruction/recreation may be beneficial to species in some contexts.     

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.     

Differential effects of spatial network structure and scale on population size and genetic diversity of the ninespine stickleback in a remnant wetland system

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Scale of habitat connectivity and colonization in fragmented nuthatch populations

Studies of effects of landscape pattern on population dynamics should consider the spatial scale at which habitat connectivity varies relative to the spatial scale of the species' behavioral response. In this paper, I investigate the relationship between the degree of connectivity of wooded patches measured at different spatial scales and the colonization of these patches by nuthatch Sitta europaea populations. I used different threshold distances to calculate the degree of habitat connectivity. A threshold distance is the distance beyond which the probability of successfully bridging the distance by dispersing individuals is assumed to decrease rapidly. The sum of the degree of connectivity over all patches in a region provides a measure for the overall degree of connectivity. Based on this measure, I could first give an indication under what conditions species may experience effects of constrained dispersal in that region. Second, the degree of connectivity of the individual patches was related with the observed colonizations of these patches. The degree of habitat connectivity measured for threshold distances of ea 2.4-3 km best explains the colonization probability of unoccupied patches. These threshold distances give an indication of the distances covered by dispersing nuthatches that led to successful colonizations.     

Habitat quality and connectivity in agricultural landscapes: The role of land use systems at various scales in time

Connectivity is a key concept of landscape ecology as it relates to flows and movements of organisms as driven by landscape structure. More and more aspects of landscape heterogeneity are considered in measuring connectivity, as the diversity of crops in agricultural landscapes. In this paper, we explored the value of considering changes and cumulated effects of connectivity over time. As an example, we analysed connectivity among patches influenced by maize over 7 years in an agricultural landscape in Brittany, France.Clear temporal patterns appeared: maize is concentrated in certain parts of the landscape, but over the period the whole area, 70% of the landscape, used for maize was connected. Instead of discrete patches, maize may produce large clusters allowing movement from patch to patch from year to year. This reinforces the importance of understanding land use allocation rules within farms and landscapes to evaluate the ecological effects of agriculture.     

Coupling inter-patch movement models and landscape graph to assess functional connectivity

Landscape connectivity is a key process for the functioning and persistence of spatially-structured populations in fragmented landscapes. Butterflies are particularly sensitive to landscape change and are excellent model organisms to study landscape connectivity. Here, we infer functional connectivity from the assessment of the selection of different landscape elements in a highly fragmented landscape in the Île-de-France region (France). Firstly we measured the butterfly preferences of the Large White butterfly (Pieris brassicae) in different landscape elements using individual release experiments. Secondly, we used an inter-patch movement model based on butterfly choices to build the selection map of the landscape elements to moving butterflies. From this map, functional connectivity network of P. brassicae was modelled using landscape graph-based approach. In our study area, we identified nine components/groups of connected habitat patches, eight of them located in urbanized areas, whereas the last one covered the more rural areas. Eventually, we provided elements to validate the predictions of our model with independent experiments of mass release-recapture of butterflies. Our study shows (1) the efficiency of our inter-patch movement model based on species preferences in predicting complex ecological processes such as dispersal and (2) how inter-patch movement model results coupled to landscape graph can assess landscape functional connectivity at large spatial scales.     

Conspecific and heterospecific attraction in assessments of functional connectivity

Functional connectivity is known to have an important, positive influence on species persistence. Measurements of functional connectivity traditionally focus on structural attributes of landscapes such as the distance and matrix type between habitat patches as well as on how species interact with those structural attributes. However, we propose that the social behavior of a species, through conspecific and heterospecific attraction, will also impact connectivity by changing how dispersers move with respect to each other and occupied patches. We analyzed functional connectivity patterns using circuit and graph theory for golden-headed lion tamarins (Leontopithecus chrysomelas) in Brazil under three scenarios. In the first scenario, we looked at connectivity without the effects of attraction under varying maximum dispersal distance and ecological movement cost thresholds. In the second scenario, we allowed dispersers to travel over more hostile matrix than they normally would to reach an occupied patch. In the final scenario, we allowed dispersers to move only to occupied patches. We found that, according to the first scenario, range-wide functional landscape connectivity for golden-headed lion tamarins is low at realistic maximum dispersal distance and movement cost thresholds. Incorporating the effects of conspecific or heterospecific attraction would increase functional connectivity, in the case of scenario two, or decrease functional connectivity, in the case of scenario three. Because conspecific/heterospecific attraction can have an impact on movement for some species, this factor should be incorporated in assessments of functional connectivity patterns for social species and others where patch occupancy is likely to influence the movements of dispersers.     

Trait-Specific Responses of Wild Bee Communities to Landscape Composition,Configuration and Local Factors

Land-use intensification and loss of semi-natural habitats have induced a severe decline of bee diversity in agricultural landscapes. Semi-natural habitats like calcareous grasslands are among the most important bee habitats in central Europe, but they are threatened by decreasing habitat area and quality, and by homogenization of the surrounding landscape affecting both landscape composition and configuration. In this study we tested the importance of habitat area, quality and connectivity as well as landscape composition and configuration on wild bees in calcareous grasslands. We made detailed trait-specific analyses as bees with different traits might differ in their response to the tested factors. Species richness and abundance of wild bees were surveyed on 23 calcareous grassland patches in Southern Germany with independent gradients in local and landscape factors. Total wild bee richness was positively affected by complex landscape configuration, large habitat area and high habitat quality (i.e. steep slopes). Cuckoo bee richness was positively affected by complex landscape configuration and large habitat area whereas habitat specialists were only affected by the local factors habitat area and habitat quality. Small social generalists were positively influenced by habitat area whereas large social generalists (bumblebees) were positively affected by landscape composition (high percentage of semi-natural habitats). Our results emphasize a strong dependence of habitat specialists on local habitat characteristics, whereas cuckoo bees and bumblebees are more likely affected by the surrounding landscape. We conclude that a combination of large high-quality patches and heterogeneous landscapes maintains high bee species richness and communities with diverse trait composition. Such diverse communities might stabilize pollination services provided to crops and wild plants on local and landscape scales.     

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.     

基于景观连接度的森林景观恢复研究——以巩义市为例

景观连接度是研究景观促进或阻碍生物体或某种生态过程在斑块间运动的程度。基于景观连接度原理,借用景观连接度指数,在地理信息系统支持下,探讨了巩义市丘陵和低山地区森林景观在不同景观距离阈值下连接度的变化,确定了分析森林景观连接度的合适距离阈值。在此基础上分析了要恢复为森林景观的农业斑块的重要值的大小,确定了对要恢复为森林景观的每一农业景观斑块对新形成的森林景观的连接度贡献大小,并结合区域地形特征,明确了森林恢复之初的重要斑块的选取及恢复的先后次序。结果显示,在不同的距离阈值下,低山地区森林景观的整体连通性指数值(IIC)都大于丘陵地区森林景观的IIC值;森林景观的适宜距离阈值,在丘陵地区可选择750 m,在低山地区可选择500 m,或者更小尺度;通过农业景观斑块重要值(dIIC)确定的对森林景观连接度作用起"非常高"和"高"的斑块的数量非常少,但对森林景观连接度的贡献作用却比较大。提出的基于景观连接来分析在森林恢复时重要斑块的选取的方法,具有一定的可操作性与实用性,对区域生态恢复和生态建设具有重要意义。     

Biodiversity and ecological forest-cover domains in boreal landscapes

Studies of spatial forest patterns have traditionally focused on habitat remnants within a landscape, but few have explicitly accounted for natural habitat patterns inherent to those landscapes. At broader scales, all cover types are to some degree subdivided. In boreal forest landscapes, forests, peat lands and water bodies form a variety of different spatial structures depending on their proportions and arrangement. To assess the spatial arrangement of forest patches relevant to organisms and ecological processes, we systematically sampled and analysed 57 boreal forest landscapes (10,000 ha in size) in central Finland. Our results show that even though forest is spatially very subdivided into discrete patches in boreal landscapes it becomes generally well-connected if narrow non-forested gaps are ignored. The proportion of forest cover varied from 17.8 to 75.3% and the number of discrete forest patches from 37 to 213. The average percolation threshold for forest cover was 46.8%. If ≤100 m wide non-forested gaps were ignored and forest patches were joined, the percolation threshold dropped to 33.3%. There were on average 13 discrete clusters of forest patches if forest patches within 200 m distance were combined. These results suggest that if a boreal forest species is able to cross even relatively narrow non-forested gaps, it is likely to perceive these boreal forest landscapes as continuous. Even though our present analysis was based on static forest cover patterns, it is important to consider landscape pattern domains when assessing habitat fragmentation and its consequences to populations of organisms.     

The relationships between habitat topology, critical scales of connectivity and tick abundance Ixodes ricinus in a heterogeneous landscape in northern Spain

The habitat mosaic was used to quantify connectivity between patches of different tick density of the notorious tick species Ixodes ricinus in an attempt to determine the cause of variations in tick abundance among apparently homogeneous sites in northern Spain. The analysis revealed that patches with high tick abundance are "stepping-stone" territories that, when removed from the landscape, cause large changes in connectivity. Sites with medium tick abundance do not cause such a critical transition in connectivity. Patches with low tick abundance, but optimal abiotic conditions for survival, are located within the minimum cost corridors network joining the patches, while those sites where the tick has been intermittently collected are located at variable distances from this network. Sites where the tick is consistently absent, but where the habitat is predicted to be suitable (old, heterogeneous forests of Quercus spp.) for the tick, are very separated from this main network of connections. These results suggest that tick distribution in a zone is highly affected not only by abiotic variables (vegetation and weather) but also by host movements. Dispersal of the tick is a function of how the hosts perceive the habitat, and the habitat's permeability to host movement. Permanent tick populations seem to be supported by the existence of these critical, high density patches, located at significant places within the habitat network.