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1.

Context

Urban landscapes are a mixture of built structures, human-altered vegetation, and remnant semi-natural areas. The spatial arrangement of abiotic and biotic conditions resulting from urbanization doubtless influences the establishment and spread of non-native species in a city.

Objectives

We investigated the effects of habitat structure, thermal microclimates, and species coexistence on the spread of a non-native lizard (Anolis cristatellus) in the Miami metropolitan area of South Florida (USA).

Methods

We used transect surveys to estimate lizard occurrence and abundance on trees and to measure vegetation characteristics, and we assessed forest cover and impervious surface using GIS. We sampled lizard body temperatures, habitat use, and relative abundance at multiple sites.

Results

At least one of five Anolis species occupied 79 % of the 1035 trees surveyed in primarily residential areas, and non-native A. cristatellus occupied 25 % of trees. Presence and abundance of A. cristatellus were strongly associated with forest patches, dense vegetation, and high canopy cover, which produced cooler microclimates suitable for this species. Presence of A. cristatellus was negatively associated with the ecologically similar non-native A. sagrei, resulting in reduced abundance and a shift in perch use of A. cristatellus.

Conclusions

The limited spread of A. cristatellus in Miami over 35 years is due to the patchy, low-density distribution of wooded habitat, which limits dispersal by diffusion. The presence of congeners may also limit spread. Open habitats—some parks, yards and roadsides—contain few if any A. cristatellus, and colonization of isolated forest habitat appears to depend on human-mediated dispersal.
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2.

Context

Methods for measuring restoration success that include functional connectivity between species’ populations are rare in landscape ecology and restoration practices. We developed an approach that analyzes connectivity between populations of target species and their dispersal probabilities to assess restoration success based on easily accessible input data. Applying this method to landscape development scenarios can help optimize restoration planning.

Objectives

We developed an assessment for restoration success and restoration planning based on functional connectivity between species’ populations and spatially explicit scenarios. The method was used in a case study to test its applicability.

Methods

Based on data on available habitat, species’ occurrence and dispersal ranges, connectivity metrics and dispersal probabilities for target species are calculated using the software Conefor Sensinode. The metrics are calculated for scenarios that reflect possible changes in the landscape to provide a basis for future restoration planning. We applied this approach to floodplain meadows along the Upper Rhine for four plant species and three future scenarios.

Results

In the case study, habitats of the target species were poorly connected. Peucedanum officinale and Sanguisorba officinalis were more successful in recolonizing new habitats than Iris spuria and Serratula tinctoria. The scenarios showed that restoration of species-rich grassland was beneficial for dispersal of the target species. As expected in the agriculturally dominated study area, restoration of former arable land significantly increased dispersal probabilities.

Conclusions

In the case study, the developed approach was easily applicable and provided reasonable results. Its implementation will be helpful in decision-making for future restoration planning.
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3.

Context

In the ecology of Lyme disease emergence, it remains unclear to what extent spread of the tick vector (Ixodes scapularis) and the pathogen (Borrelia burgdorferi) are dependent upon the dispersal of vertebrate hosts in spatially heterogeneous landscapes. Yet, empirical measure of these complex ecologically driven spread processes present conceptual and methodological challenges despite important public health implications.

Objectives

To examine the relationship between landscape characteristics and tick-borne disease spread, we modeled the influence of landscape connectivity for a simplified vertebrate host community (white-footed mouse—Peromyscus leucopus, American robin—Turdus migratorius, white-tailed deer—Odocoileus virginianus) on the potential spread of the tick population compared to the pathogen in a spatially-structured landscape.

Methods

We parameterized a hybrid demographic-dispersal connectivity model by combining a series of reported host dispersal and tick burden estimates with empirically-measured tick abundance and pathogen prevalence sampled from a Lyme-endemic island landscape in Thousand Islands National Park (Ontario, Canada) and simulated several tick- and pathogen-spread scenarios.

Results

The extent of tick spread by mice [amount of reachable habitat (ARH)?=?18.0%] is considerably similar to that of robins (ARH?=?18.7%), while deer support the greatest tick spread extent (ARH?=?82.0%). Infected mice carrying ticks support the highest pathogen spread (ARH?=?19.8%). Short-distance pathogen spread and long-distance tick spread were facilitated by intermediate stepping stone habitat fragments.

Conclusions

We provide evidence that host functional connectivity mediates tick spread differently than pathogen spread, and depends strongly on landscape configuration. Our study therefore emphasizes the importance of landscape spatial heterogeneity on the ecological processes that influence regional tick-borne disease spread.
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4.

Context

Various species of forest trees are commonly used for ornamental purposes and are therefore frequently found in nonforest ecosystems. They constitute an important component of the so-called trees outside forests (TOF). Not much is known, however, about the drivers of TOF spatial distribution either in urbanized or in agricultural landscapes since they are generally absent from forest inventories.

Objective

The present study focused on the spatial distribution of TOF across agricultural landscapes and their potential role in the dispersal of a forest pest insect, the pine processionary moth, Thaumetopoea pityocampa (PPM).

Methods

All the TOF belonging to the genera Pinus, Cedrus and Pseudotsuga were considered as potential hosts and inventoried within a 22 × 22 km study window. We fitted a nonstationary Poisson process to the empirical data and used the distance to the nearest building as a covariate.

Results

Both empirical and simulated data indicated that TOF associated to human artifacts/urbanized areas constituted the main source of landscape connectivity for the PPM in the open fields under study. Because they do not account for TOF, forest inventories dramatically underestimate landscape connectivity and provide an erroneous picture of the PPM habitat distribution.

Conclusions

We conclude that TOF, especially the ornamental component, must be taken into account when it comes to understanding forest insect landscape dynamics or genetics. The omnipresence of TOF also suggests a potentially huge role in pest dispersal and invasive species expansion.
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5.

Context

Dispersal is essential for species persistence and landscape genetic studies are valuable tools for identifying potential barriers to dispersal. Macaws have been studied for decades in their natural habitat, but we still have no knowledge of how natural landscape features influence their dispersal.

Objectives

We tested for correlations between landscape resistance models and the current population genetic structure of macaws in continuous rainforest to explore natural barriers to their dispersal.

Methods

We studied scarlet macaws (Ara macao) over a 13,000 km2 area of continuous primary Amazon rainforest in south-eastern Peru. Using remote sensing imagery from the Carnegie Airborne Observatory, we constructed landscape resistance surfaces in CIRCUITSCAPE based on elevation, canopy height and above-ground carbon distribution. We then used individual- and population-level genetic analyses to examine which landscape features influenced gene flow (genetic distance between individuals and populations).

Results

Across the lowland rainforest we found limited population genetic differentiation. However, a population from an intermountain valley of the Andes (Candamo) showed detectable genetic differentiation from two other populations (Tambopata) located 20–60 km away (F ST = 0.008, P = 0.001–0.003). Landscape resistance models revealed that genetic distance between individuals was significantly positively related to elevation.

Conclusions

Our landscape resistance analysis suggests that mountain ridges between Candamo and Tambopata may limit gene flow in scarlet macaws. These results serve as baseline data for continued landscape studies of parrots, and will be useful for understanding the impacts of anthropogenic dispersal barriers in the future.
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6.

Context

Considerable research has examined scale effects for patch-based metrics with the ultimate goal of predicting values at finer resolutions (i.e., downscaling), but results have been inconsistent. Surface metrics have been suggested as an alternative to patch-based metrics, although far less is known about their scaling relationships and downscaling potential. If successful, downscaling would enable integration of disparate datasets and comparison of landscapes using different resolution datasets.

Objectives

(1) Determine how surface metrics scale as resolution changes and how consistent those scaling relationships are across landscapes. (2) Test whether these scaling relationships can be accurately downscaled to predict metric values for finer resolutions.

Methods

Various scaling functions were fit to 16 surface metrics computed for multiple resolutions for a set of landscapes. Best-fitting functions were then extrapolated to test downscaling behavior (i.e., predict metric value for a finer resolution) for an independent set of validation landscapes. Relative error was assessed between the predicted and true values to determine downscaling robustness.

Results

Seven surface metrics (Sa, Sq, S10z, Sdq, Sds, Sdr, Srwi) fit consistently well (R2 > 0.99) with a 3rd order polynomial or power law. Of those, the scaling functions for Sa, Sq, and S10z were able to predict metric values at a finer resolution within 5 %. Three metrics, (Ssk, Sku, Sfd) were also notable in terms of fit and downscaling.

Conclusions

Many metrics exhibit consistent scaling relations across resolution, and several are able to accurately predict values at finer resolutions. However, prediction accuracy is likely related to the amount of information lost during aggregation.
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7.

Context

The species–area relationship (SAR) is the most ubiquitous scaling relationship in ecology, yet we still do not know how different aspects of scale affect this relationship. Scale is defined by grain, extent, and focus. Focus here pertains to whether patches or landscapes are used to derive SARs.

Objective

To explore whether altering the focal scale influences the resulting SAR. If the SAR is scale-invariant, patch-based and landscape-based SARs should be congruent.

Methods

I fit a power-law function (S = cA z) to arthropod data obtained from an experimental landscape system, in which habitat amount and configuration (clumped vs. fragmented) of red clover (Trifolium pratense) varied among plots (256 m2). The scaling coefficient (z) was compared among patch-based and landscape-based SARs for congruence.

Results

Patches gained species at a faster rate than landscapes (z = 0.37 vs. 0.26, respectively), producing domains of incongruity in the SAR. Landscape richness (S L) was greater than patch richness (S P) below 30 % habitat, but S P > S L above 60 % habitat. Landscape configuration contributed to this incongruity below 30 % habitat (fragmented S L > clumped S L), but landscape context (whether the largest patch was embedded in a fragmented or clumped landscape) was important above 60 % habitat for understanding the SAR in this domain.

Conclusions

Landscape configuration exerts both direct (<30 % habitat) and indirect (>60 % habitat) effects on the SAR. Because patch-based and landscape-based SARs may not be congruent, we should exercise care when extrapolating from patches to landscapes to make inferences about the effects of habitat loss and fragmentation on species richness.
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8.

Context

Abundance and diversity of bumblebees have been declining over the past decades. To successfully conserve bumblebee populations, we need to understand how landscape characteristics affect the quantity and quality of floral resources collected by colonies and subsequently colony performance.

Objectives

We therefore investigated how amount and composition of pollen collected by buff-tailed bumblebee Bombus terrestris colonies was affected by the surrounding landscape (i.e. the proportion of forest, urban, semi-natural habitats) and how they were related to colony growth.

Methods

Thirty B. terrestris colonies were placed at grassland sites differing in surrounding landscape. Colonies were established in spring when availability of flowering plants was highest, and their weight gain was monitored for 1 month. We additionally recorded the quantity and compared plant taxonomic composition and nutritional quality (i.e. amino acid composition) of pollen stored.

Results

Bumblebee colonies varied little in the pollen spectra collected despite differences in surrounding landscape composition. They collected on average 80 % of pollen from woody plants, with 34 % belonging to the genus Acer. Early colony growth positively correlated with total amount of woody pollen and protein collected and decreased with increasing proportions of semi-natural habitats and total amino acid concentrations.

Conclusions

Our results suggest that woody plant species represent highly important pollen sources for the generalist forager B. terrestris early in the season. We further show that colony growth of B. terrestris is predominantly affected by the quantity, not quality, of forage, indicating that several abundant plant species flowering throughout the bumblebees’ foraging season may cover the colonies’ nutritional needs.
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9.

Context

Interactions between landscape-scale processes and fine-grained habitat heterogeneity are usually invoked to explain species occupancy in fragmented landscapes. In variegated landscapes, however, organisms face continuous variation in micro-habitat features, which makes necessary to consider ecologically meaningful estimates of habitat quality at different spatial scales.

Objectives

We evaluated the spatial scales at which forest cover and tree quality make the greatest contribution to the occupancy of the long-horned beetle Microplophorus magellanicus (Coleoptera: Cerambycidae) in a variegated forest landscape.

Methods

We used averaged data of tree quality (as derived from remote sensing estimates of the decay stage of single trees) and spatially independent pheromone-baited traps to model the occurrence probability as a function of multiple cross-scale combinations between forest cover and tree quality (with scales ranging between 50 and 400 m).

Results

Model support and performance increased monotonically with the increasing scale at which tree quality was measured. Forest cover was not significant, and did not exhibit scale-specific effects on the occurrence probability of M. magellanicus. The interactive effect between tree quality and forest cover was stronger than the independent (additive) effects of tree quality and particularly forest cover. Significant interactions included tree quality measured at spatial scales ≥200 m, but cross-scale interactions occurred only in four of the seven best-supported models.

Conclusions

M. magellanicus respond to the high-quality trees available in the landscape rather than to the amount of forest per se. Conservation of viable metapopulations of M. magellanicus should consider the quality of trees at spatial scales >200 m.
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10.

Context

Habitat fragmentation generates a loss of functional connectivity detrimental to the persistence of biodiversity. The French agricultural intensification initiated in the 1950s has caused a decline in field margins.

Objectives

As field margins may facilitate species dispersal while providing socio-economic benefits, it is of interest to assess their contribution to the functional connectivity of insect-pollinated plants in agro-ecosystems. This will help develop appropriate management strategies mitigating fragmentation.

Methods

We addressed this issue by studying the links between landscape structure and the patterns of abundance and pollen dispersal (using fluorescent dye particles) for two contrasted insect-pollinated plants occurring in field margins (Crepis sancta and Euphorbia serrata). We investigated the influence of field margins quality and of the surrounding matrix on pollen dispersal and compared the relevance of the least-cost algorithm with a straight-line approach to depict pollinators’ movements.

Results

The influence of landscape structure on plant abundance is species and scale-specific. Pollen dispersal decreases with distance from the source. For E. serrata, it was preferentially dispersed via field margins, confirming the relevance of the least-cost algorithm, while C. sancta dispersal followed a straight-line.

Conclusions

Euphorbia serrata, which grows strictly on field margins with a greater dispersal ability and a more diversified pollinator guild than C. sancta, is less affected by land-use changes. Our study demonstrates the contrasting contributions of field margins to pollen dispersal as they may act as functional corridors favouring pollinators’ movement depending on the species of interest.
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11.

Context

In deserts, many plant species exhibit a patchy spatial distribution within a harsh habitat matrix, where the likelihood of propagule dispersal among patches is uncertain, but may be promoted by landscape corridors or dispersal vectors.

Objectives

We examine the connectivity of a representative desert plant species (Acacia (Senegalia) greggii), and the ability of three major factors (animal dispersal agents, water flow along dry-washes, and climate) to facilitate dispersal within four watersheds in the Mojave National Preserve.

Methods

We genotyped 323 individuals sampled across 22 one-hectare sites using ten nuclear microsatellite markers.

Results

A hierarchical AMOVA revealed no significant differentiation among watersheds (F RT = 0.00, P > 0.10), and very little genetic structure among all sites (F ST = 0.03, P < 0.001), indicating regional connectivity. Mantel tests indicated distance along dry-washes best explained genetic distance between sites (r = 0.47, P < 0.05) when compared to Euclidean distance (P > 0.05), a distance measure based on rodent dispersal (P > 0.05), and a distance measure avoiding inhospitable climate (P > 0.05). An AIC comparison of generalized linear models found that within site genetic diversity (H E and allelic richness) and average relatedness were best explained by slope (which increases seed dispersal potential via water flow) and area of the upstream watershed (which determines the number of potential seed donors), rather than plant density or habitat suitability.

Conclusions

Together, these findings indicate that dry-washes are key landscape features that enhance dispersal and regional connectivity in this patchy desert plant.
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12.

Context

Beyond the recognized importance of protecting large areas of contiguous habitat, conservation efforts for many species are complicated by the fact that patch suitability may also be affected by characteristics of the landscape within which the patch is located. Currently, little is known about the spatial scales at which species respond to different aspects of the landscape surrounding an occupied patch.

Objectives

Using grassland bird point count data, we describe an approach to evaluating scale-specific effects of landscape composition on patch occupancy.

Methods

We used data from 793 point count surveys conducted in idle and grazed grasslands across Wisconsin, USA from 2012 to 2014 to evaluate scale-dependencies in the response of grassland birds to landscape composition. Patch occupancy models were used to evaluate the relationship between occupancy and landscape composition at scales from 100 to 3000 m.

Results

Bobolink (Dolichonyx oryzivorus) exhibited a pattern indicating selection for grassland habitats in the surrounding landscape at all spatial scales while selecting against other habitats. Eastern Meadowlark (Sturnella magna) displayed evidence of scale sensitivity for all habitat types. Grasshopper Sparrow (Ammodramus savannarum) showed a strong positive response to pasture and idle grass at all scales and negatively to cropland at large scales. Unlike other species, patch occupancy by Henslow’s Sparrow (A. henslowii) was primarily influenced by patch area.

Conclusions

Our results suggest that both working grasslands (pasture) and idle conservation grasslands can play an important role in grassland bird conservation but also highlight the importance of considering species-specific patch and landscape characteristics for effective conservation.
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13.
14.

Context

Agroecosystems are dynamic, with yearly changing proportions of crops. Explicit consideration of this temporal heterogeneity is required to decipher population and community patterns but remains poorly studied.

Objectives

We evaluated the impact on the activity-density of two dominant carabid species (Poecilus cupreus and Anchomenus dorsalis) of (1) local crop, current year landscape composition, and their interaction, and (2) inter-annual changes in landscape composition due to crop rotations.

Methods

Carabids were sampled using pitfall-traps in 188 fields of winter cereals and oilseed rape in three agricultural areas of western France contrasting in their spatial heterogeneity. We summarized landscape composition in the current and previous years in a multi-scale perspective, using buffers of increasing size around sampling locations.

Results

Both species were more abundant in oilseed rape, and in landscapes with a higher proportion of oilseed rape in the previous year. P. cupreus abundance was negatively influenced by oilseed rape proportion in the current year landscape in winter cereals and positively by winter cereal proportion in oilseed rape. A. dorsalis was globally impacted at finer scales than P. cupreus.

Conclusions

Resource concentration and dilution-concentration processes jointly appear to cause transient dynamics of population abundance and distribution among habitat patches. Inter-patch movements across years appear to be key drivers of carabids’ survival and distribution, in response to crop rotation. Therefore, the explicit consideration of the spatiotemporal dynamics of landscape composition can allow future studies to better evidence ecological processes behind observed species patterns and help developing new management strategies.
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15.

Context

Emissions of greenhouse gases in urban areas play an important role in climate change. Increasing attention has been given to urban landscape structure–emission relationships (SERs). However, it remains unknown if and to what extent SERs are dependent on observational scale.

Objective

To assess how changing observational scales (in terms of spatial and thematic resolutions) of urban landscape structure affect SERs.

Methods

We examined correlations between 16 landscape metrics and greenhouse gas emissions across 52 European cities, through (1) systematic manipulation of spatial and thematic resolutions of the urban land use/cover (ULUC) dataset, and (2) comparison between available standard ULUC datasets with different spatial resolutions.

Results

Our analyses showed that the observed SERs significantly depend on both thematic and spatial resolutions of the ULUC data. For the 16 landscape metrics, we found diverse spatial/thematic scaling relations exhibiting monotonic, hump-shaped or scale-invariant trends. For different landscape metrics, the SERs were strongest at different spatial scales, suggesting that there is no consistent scaling relation over those observational scales.

Conclusions

SERs are highly sensitive to spatial and thematic resolutions of landscape data. To avoid the problem of ‘ecological fallacy,’ important caveats should be taken for interpretations based on single landscape metrics. Particular consideration should be paid to metrics that are easily interpretable, predictable in scaling behaviors, and important for shaping SERs, such as PLAND, ED, and LPI. Systematic investigations on scaling behaviors of SERs over well-defined scale domains are encouraged in future studies linking greenhouse gas emissions and urban landscape structure.
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16.

Context

Conservation planning is increasingly using “coarse filters” based on the idea of conserving “nature’s stage”. One such approach is based on ecosystems and the concept of ecological integrity, although myriad ways exist to measure ecological integrity.

Objectives

To describe our ecosystem-based index of ecological integrity (IEI) and its derivative index of ecological impact (ecoImpact), and illustrate their applications for conservation assessment and planning in the northeastern United States.

Methods

We characterized the biophysical setting of the landscape at the 30 m cell resolution using a parsimonious suite of settings variables. Based on these settings variables and mapped ecosystems, we computed a suite of anthropogenic stressor metrics reflecting intactness (i.e., freedom from anthropogenic stressors) and resiliency metrics (i.e., connectivity to similar neighboring ecological settings), quantile-rescaled them by ecosystem and geographic extent, and combined them in a weighted linear model to create IEI. We used the change in IEI over time under a land use scenario to compute ecoImpact.

Results

We illustrated the calculation of IEI and ecoImpact to compare the ecological integrity consequences of a 70-year projection of urban growth to an alternative scenario involving securing a network of conservation core areas (reserves) from future development.

Conclusions

IEI and ecoImpact offer an effective way to assess ecological integrity across the landscape and examine the potential ecological consequences of alternative land use and land cover scenarios to inform conservation decision making.
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17.

Context

Humans have altered grasslands in recent decades through crop conversion, woody encroachment, and plant invasions. Concurrently, grassland birds have experienced range-wide declines. Studies have reported effects of plant invasions and land conversion on nest ecology, but few have assessed relative impacts of these changes.

Objectives

We compared impacts of invasive plants and landscape context on nest survival of a grassland songbird, the dickcissel (Spiza americana). We also compared effects on parasitism by brown-headed cowbirds (Molothrus ater) and tested whether parasitism affects survival.

Methods

From 2013–2016, we monitored 477 dickcissel nests. We measured nest-site vegetation (including woody plants, tall fescue Schedonorus arundinaceous, and other invasive grasses) and measured landscape context at broad scales.

Results

Nest survival declined with increasing tall fescue cover at nest sites, and parasitism was more common at nests with greater fescue and woody cover. Some evidence suggested a negative effect of row-crop cover within 1000 m on nest survival, but no landscape patterns unambiguously affected survival. Woodland cover and wooded-edge prevalence were associated with reduced parasitism risk. Parasitized nests had smaller clutches, failed more frequently, and produced fewer fledglings than non-parasitized nests.

Conclusions

Determining the impacts of invasive plants and other anthropogenic changes on grassland birds will aid in prioritizing management to improve habitat quality. Our results indicate that optimizing landscape context around habitats may not affect dickcissel nest survival strongly, except perhaps through effects on parasitism. In contrast, controlling tall fescue and shrubs within grasslands could benefit birds by increasing nest success and reducing parasitism.
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18.

Context

Interactions among disturbances, climate, and vegetation influence landscape patterns and ecosystem processes. Climate changes, exotic invasions, beetle outbreaks, altered fire regimes, and human activities may interact to produce landscapes that appear and function beyond historical analogs.

Objectives

We used the mechanistic ecosystem-fire process model FireBGCv2 to model interactions of wildland fire, mountain pine beetle (Dendroctonus ponderosae), and white pine blister rust (Cronartium ribicola) under current and future climates, across three diverse study areas.

Methods

We assessed changes in tree basal area as a measure of landscape response over a 300-year simulation period for the Crown of the Continent in north-central Montana, East Fork of the Bitterroot River in western Montana, and Yellowstone Central Plateau in western Wyoming, USA.

Results

Interacting disturbances reduced overall basal area via increased tree mortality of host species. Wildfire decreased basal area more than beetles or rust, and disturbance interactions modeled under future climate significantly altered landscape basal area as compared with no-disturbance and current climate scenarios. Responses varied among landscapes depending on species composition, sensitivity to fire, and pathogen and beetle suitability and susceptibility.

Conclusions

Understanding disturbance interactions is critical for managing landscapes because forest responses to wildfires, pathogens, and beetle attacks may offset or exacerbate climate influences, with consequences for wildlife, carbon, and biodiversity.
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19.

Context

Projected increases in human population size are expected to increase forest loss and fragmentation in the next century at the expense of forest-dwelling species.

Objectives

We estimated landscape carrying capacity (N k) for Ovenbirds in urban, suburban, exurban, and rural areas for the years 2000 and 2050, and compared changes in N k with changes in occupancy probability.

Methods

Maximum clique analysis, a branch of mathematical graph theory, was used to estimate landscape carrying capacity, the maximum potential number of territories a given landscape is capable of supporting (N k). We used occupancy probability maps as inputs for calculating Ovenbird N k in the northeastern USA and a spatially explicit growth model to forecast future development patterns in 2050. We compared occupancy probability with estimates of N k for urban, suburban, exurban, and rural areas for the years 2000 and 2050.

Results

In response to human population growth and development, Ovenbird N k was predicted to decrease 23% in urban landscapes, 28% in suburban landscapes, 43% in exurban landscapes, and 20% in rural landscapes. These decreases far exceeded decreases in mean occupancy probabilities that ranged between 2 and 5% across the same development categories. Thus, small decreases in occupancy probability between 2000 and 2050 translated to much larger decreases in N k.

Conclusions

For the first time, our study compares occupancy probability with a species population metric, N k, to assess the impact of future development. Maximum clique analysis is a tool that can be used to estimate N k and inform landscape management and communication with stakeholders.
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20.

Context

The pasture-woodlands of Central Europe are low-intensity grazing systems in which the structural richness of dynamic forest-grassland mosaics is causal for their high biodiversity. Distinct mosaic patterns in Picea abies- and Fagus sylvatica-dominated pasture-woodlands in the Swiss Jura Mountains suggest a strong influence of tree species regeneration ecology on landscape structural properties. At the landscape scale, however, cause-effect relationships are complicated by habitat selectivity of livestock.

Objectives

We asked which tree species regeneration traits and what kind of feedbacks among local-scale vegetation dynamics and landscape-scale herbivore behavior are causal for the contrasted landscape structural characteristics of Picea- and Fagus-dominated pasture-woodlands.

Methods

We performed simulation experiments of mosaic pattern formation in both pasture-woodland types. The regeneration traits, namely dispersal distance, resistance to browsing and tolerance to shade, and the rules for habitat selection of cattle were modified and the corresponding shifts in landscape structure were analyzed.

Results

Dispersal distance showed a significant, but only local, effect promoting forest fringe formation. Saplings’ resistance to browsing mainly determined overall tree cover, but did not influence landscape structure. At the landscape scale, both shade tolerance of saplings and selective habitat use by cattle were responsible for forest-grassland segregation: high shade tolerance triggered segregation, whereas non-selective habitat use hindered it.

Conclusions

Existing local-scale theory on pasture-woodland dynamics is complemented by an herbivore-vegetation feedback among spatial scales. In low-intensity pastures, where large herbivores are preferentially “grazers” and trees form dense canopies, an intrinsic trend towards forest-grassland segregation at the expense of forest-grassland ecotones is predicted.
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