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1.

Context

Woody plant encroachment—the conversion of grasslands to woodlands—is among the greatest challenges faced by rangelands worldwide. Yet this phenomenon is poorly understood, and complex land use dynamics make interpreting the timing and extent of land cover changes a global challenge.

Objectives

For many regions, the true degree and rate of historical change in woody cover and cropland remain unknown. We quantify these processes and explore the effects of prior cultivation on woody plant distribution.

Methods

In the Lampasas Cut Plain, USA, we measured rangeland transformation using digital classification of aerial imagery 1937–2012. Our study is the first to use data of such high spatial and temporal resolution to address this question at this scale. We also provide some of the first documentation of dramatic regional cropland abandonment.

Results

Although total woody cover remained almost unchanged (1937: 28%, 2012: 27 %), woody cover underwent a major redistribution across the landscape. Formerly open areas attained much greater levels of woody cover, and previously wooded areas lost woody cover. As cropland area declined by 78 %, woody plants invaded former croplands more slowly than the rangeland portions of the area (0.1 % year?1 vs. 0.3 % year?1, respectively).

Conclusions

These findings conflict with widely held assumptions and suggest that woody plant encroachment is more nuanced than often recognized. Multiple dynamics and past conditions interact in complex ways to produce landscape change. Because perceptions of encroachment determine how we respond to this challenge, great care should be taken in interpreting observed woody plant encroachment of the world’s rangelands.
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2.

Context

Woodland and agricultural expansion are major causes of grassland fragmentation. Fire and rainfall play important roles in maintaining grasslands, however, fire activity has been reduced in fragmented landscapes.

Objectives

Quantify the degree to which basic landscape fragmentation metrics could be used as drivers of woody cover potential.

Methods

Woody plant percent cover was calculated between 2004 and 2008 at?>?2000 sites. At each site, we calculated these fragmentation metrics for grassland cover type (classified by the National Land Cover Database); # patches, landscape proportion, edge density, largest patch index, effective mesh size and patch cohesion index within 3 circular areas (10 km2, 360 km2 and 3600 km2) surrounding the sampling site. A quantile regression was performed to identify which metrics were useful at predicting the 25th, 50th, 75th or 95th quantile of woody cover distribution.

Results

Grassland proportion and edge density were significant predictors of the woody plant potential (75th and 95th quantile). Woody cover potential was positively associated with edge density suggesting that fragmented areas (i.e., areas with high number of edges) maintained higher woody cover, while grassland proportion was negatively associated with woody plant potential.

Conclusion

We propose that in addition to a lack of fire, fragmented landscapes may facilitate further woodland expansion by reducing natural land and restricting grasslands to smaller, less connected patches, which can maintain higher woody cover. Given current trends in woodland expansion, special attention should be given to areas that are found within a fragmented landscape and climatically prone to woodland expansion.
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3.

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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4.

Context

Terrestrial ecosystems, including tropical forests, are hypothesized to have tipping points beyond which environmental change triggers rapid and radical shifts to novel alternative states.

Objective

We explored the overarching hypothesis that fire-mediated alternative stable states exist in the semi-deciduous tropical forest zone of Ghana, and that increased fire activity has pushed some forests to a new state in which a novel ecosystem with low tree density is maintained by fire.

Methods

We combined a 30-year time series of remotely-sensed data with field measurements to assess land cover trends, the effects of fire on forest vegetation, and the reciprocal effects of vegetation change on fire regimes, in four forest reserves. We analyzed precipitation trends to determine if shifts in vegetation and fire regime reflected a shift to a drier climate.

Results

Two of the reserves experienced forest loss, were impacted by frequent fires, and transitioned to a vegetation community dominated by shrubs and grasses, which was maintained by fire–vegetation feedbacks. The other two reserves experienced less fire, retained higher levels of forest cover, and resisted fire encroachment from surrounding agricultural areas. Precipitation remained relatively stable, suggesting a hysteresis effect in which different vegetation states and fire regimes coexist within a similar climate.

Conclusion

There is potential for human land use and fire to create novel and persistent non-forest vegetation communities in areas that are climatically suitable for tropical forests. These disturbance-mediated regime shifts should be taken into account when assessing future trajectories of forest landscape change in West Africa.
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5.

Context

We address the issue of adapting landscapes for improved insect biodiversity conservation in a changing climate by assessing the importance of additive (main) and synergistic (interaction) effects of land cover and land use with climate.

Objectives

We test the hypotheses that ant richness (species and genus), abundance and diversity would vary according to land cover and land use intensity but that these effects would vary according to climate.

Methods

We used a 1000 m elevation gradient in eastern Australia (as a proxy for a climate gradient) and sampled ant biodiversity along this gradient from sites with variable land cover and land use.

Results

Main effects revealed: higher ant richness (species and genus) and diversity with greater native woody plant canopy cover; and lower species richness with higher cultivation and grazing intensity, bare ground and exotic plant groundcover. Interaction effects revealed: both the positive effects of native plant canopy cover on ant species richness and abundance, and the negative effects of exotic plant groundcover on species richness were greatest at sites with warmer and drier climates.

Conclusions

Impacts of climate change on insect biodiversity may be mitigated to some degree through landscape adaptation by increasing woody native vegetation cover and by reducing land use intensity, the cover of exotic vegetation and of bare ground. Evidence of synergistic effects suggests that landscape adaptation may be most effective in areas which are currently warmer and drier, or are projected to become so as a result of climate change.
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6.

Context

Quantifying landscape-scale vegetation disturbances by surface coal mining (SCM) is crucial for assessing and mitigating its negative impacts on the environment. Methods for detecting such disturbances in woody ecosystems exist, but these methods do not work well for deserts and grasslands in arid and semiarid regions because of their sensitive responses to precipitation variations.

Objectives

The objective of this study was to develop a new index to reliably detect the locations and spatial extents of SCM-induced vegetation disturbances in dryland regions in the face of fluctuating precipitation.

Methods

We have developed a vegetation disturbance index (VDI) that combines MODIS EVI data with precipitation data to detect vegetation disturbances by SCM on the Mongolian Plateau during 2000–2015. The VDI is computed by comparing vegetation production per unit precipitation for a given year with a multi-year mean, and by considering distances from coal-mining areas.

Results

Our results show that the VDI was able to adequately distinguish vegetation disturbances by SCM from climate-driven vegetation changes in five selected sites across the Mongolian Plateau.

Conclusions

The VDI provides an effective tool for quantifying the locations, spatial extents, and severity of vegetation disturbances by SCM in arid and semiarid regions.
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7.

Context

Traditionally, studies of habitat fragmentation have focused on spatial isolation of habitats. Meanwhile, the role of fragmentation of land ownership and hence of parcelization of habitats remains, particularly in relation to management of semi-natural grasslands, not well understood.

Objective

We propose that, especially in a Danish context, fragmentation of land ownership leads to parcelization of semi-natural grassland habitats. This results in small parcel sizes, obstructing cost effective management in terms of grazing and mowing and consequently leads to encroachment of scrubs, threatening biodiversity.

Methods

We applied national, spatially explicit information about land ownership, management, semi-natural grasslands and vegetation height to examine the relationships between parcel size, management and the proportion of scrubs on semi-natural grasslands.

Results

Results from a regression analysis show that parcel size is significantly negatively related to proportion of scrubs; i.e. small parcels are associated with higher proportions of scrubs compared to large parcels. The results also show that the size of ownership parcels has a stronger explanatory power for the proportion of scrub compared to the size of habitat parcels, where ownership boundaries are not taken into account. Furthermore, parcels, with legal obligations for management, have significantly lower proportion of scrubs compared to parcels without management obligations.

Conclusions

Efforts for conservation of and improvement of biodiversity on semi-natural grassland should pay increasing attention towards the importance of fragmentation of land ownership and parcelization of habitats. Our results point at the need for cross-farm cooperation to secure continuous grassland management to prevent scrub encroachment.
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8.

Context

Plant invasions of native ecosystems are one of the main causes of declines in biodiversity via system-simplification. Restoring native biodiversity can be particularly challenging in landscapes where invasive species have become dominant and where a new set of feedbacks reinforce an invaded state and preclude restoration actions. We lack an understanding of the response of invaded systems to landscape-level manipulations to restore pattern and process relationships and how to identify these relationships when they do not appear at the expected scale.

Objectives

To better understand how fire and grazing influence landscape-level heterogeneity in invaded landscapes, we assess the scale at which grazing pressure and seasonality mediate the success of re-introducing a historical disturbance regime, grazing driven by fire (termed pyric herbivory), to an invasive plant-dominated landscape.

Methods

We manipulated grazing timing and intensity in exotic grass-dominated grasslands managed for landscape heterogeneity with spring fire and grazing. In pastures under patch-burn grazing management, we evaluated the spatial and temporal variability of plant functional groups and vegetation structure among and within patches managed with separate grazing systems: season-long stocking and intensive early stocking.

Results

Warm- and cool-season grasses exhibited greater among-patch variability in invasive-plant dominated grassland under intensive early grazing than traditional season-long grazing, but landscape-level heterogeneity, as measured through vegetation structure was minimal and invariable under both levels of grazing pressure, which contrasts findings in native-dominated systems. Moreover, within-patch heterogeneity for these functional groups was detected; contrasting the prediction that among-patch heterogeneity, in mesic grasslands, manifests from within-patch homogeneity.

Conclusions

In invaded grasslands, manipulation of grazing pressure as a process that drives heterogeneous vegetation patterns influences native and non-native grass heterogeneity, but not heterogeneity of vegetation structure, within and among patches managed with fire. Fire and grazing-moderated heterogeneity patterns observed in native grass-dominated grasslands likely differ from invasive grass-dominated grasslands with implications for using pyric herbivory in invaded systems.
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9.

Context

Natural regenerating forests are rapidly expanding in the tropics. Forest transitions have the potential to restore biodiversity. Spatial targeting of land use policies could improve the biodiversity benefits of reforesting landscapes.

Objective

We explored the relative importance of landscape attributes in influencing the potential of tree cover increase to restore native woody plant biodiversity at the landscape scale.

Methods

We developed land use scenarios that differed in spatial patterns of reforestation, using the Pangor watershed in the Ecuadorian Andes as a case study. We distinguished between reforestation through natural regeneration of woody vegetation in abandoned fallows and planted forests through managed plantations of exotic species on previously cultivated land. We simulated the restoration of woody plant biodiversity for each scenario using LANDIS-II, a process-based model of forest dynamics. A pair-case comparison of simulated woody plant biodiversity for each scenario was conducted against a random scenario.

Results

Species richness in natural regenerating fallows was considerably higher when occurring in: (i) close proximity to remnant forests; (ii) areas with a high percentage of surrounding forest cover; and (iii) compositional heterogeneous landscapes. Reforestation at intermediate altitudes also positively affected restoration of woody plant species. Planted exotic pine forests negatively affected species restoration.

Conclusions

Our research contributes to a better understanding of the recolonization processes of regenerating forests. We provide guidelines for reforestation policies that aim to conserve and restore woody plant biodiversity by accounting for landscape attributes.
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10.

Context

With global change, microclimates become important refuges for temperature-sensitive, range-restricted organisms. In African savannas, woody vegetation on Macrotermes mounds create widely-dispersed microclimates significantly cooler than the surrounding matrix, which buffer against elevated temperatures at the finer scale of mounds, allowing species to persist at the landscape scale. Termite colonies cultivate symbiotic fungi to digest lignin, but the fungi require temperatures between 29 and 32 °C, which termites strive to maintain. Mound-associated vegetation is a hot-spot for elephant herbivory, so removal of woody species cover by elephants could influence mound-associated microclimates, impacting temperature regulation by termites.

Objectives

We explored the interaction between two prominent ecosystem engineers (termites and elephants) to ascertain whether elephant removal of mound woody cover affects (1) external mound-associated microclimate and (2) internal mound temperature.

Methods

We surveyed 44 mounds from three sites in Kruger National Park, South Africa, during an El Niño/Southern Oscillation-induced drought and heatwave, recording whether sub-canopy, external, mound-surface and internal mound temperatures varied with vegetation removal by elephant.

Results

Elephant damage to mound-associated vegetation reduces the fine-scale microclimate effect provided by vegetation on Macrotermes mounds. Despite this, termites were able to regulate internal mound temperatures, whereas internal temperatures of abandoned mounds increased with elevated surface temperatures.

Conclusions

Termites can persist despite loss of mound-associated microclimates, but the loss likely increases energetic costs of mound thermoregulation. Since mound vegetation buffers against drought, loss of widely-dispersed, fine-scale microclimates could increase as megaherbivores remain constrained to protected areas, impacting climate-sensitive organisms and ecosystem function at a range of scales.
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11.

Context

The history of the landscape directly affects biotic assemblages, resulting in time lags in species response to disturbances. In highly fragmented environments, this phenomenon often causes extinction debts. However, few studies have been carried out in urban settings.

Objectives

To determine if there are time lags in the response of temperate natural grasslands to urbanization. Does it differ for indigenous species and for species indicative of disturbance and between woody and open grasslands? Do these time lags change over time? What are the potential landscape factors driving these changes? What are the corresponding vegetation changes?

Methods

In 1995 and 2012 vegetation sampling was carried out in 43 urban grassland sites. We calculated six urbanization and landscape measures in a 500 m buffer area surrounding each site for 1938, 1961, 1970, 1994, 1999, 2006, and 2010. We used generalized linear models and model selection to determine which time period best predicted the contemporary species richness patterns.

Results

Woody grasslands showed time lags of 20–40 years. Contemporary open grassland communities were, generally, associated with more contemporary landscapes. Altitude and road network density of natural areas were the most frequent predictors of species richness. The importance of the predictors changed between the different models. Species richness, specifically, indigenous herbaceous species, declined from 1995 to 2012.

Conclusions

The history of urbanization affects contemporary urban vegetation assemblages. This indicates potential extinction debts, which have important consequences for biodiversity conservation planning and sustainable future scenarios.
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12.

Purpose

Wildlife conservation requires understanding how landscape context influences habitat selection at spatial scales broader than the territory or habitat patch.

Objectives

We assessed how landscape composition, fragmentation, and disturbance affected occurrence and within-season site-fidelity of a declining grassland songbird species (Henslow’s Sparrow, Ammodramus henslowii).

Methods

Our study area encompassed eastern Kansas (USA) and North America’s largest remaining tracts of tallgrass prairie. We conducted 10,292 breeding-season point-count surveys over 2 years, and related occurrence and within-season site-occupancy dynamics of sparrows to landscape attributes within 400-, 800-, and 1600-m radii.

Results

Sparrows inhabited < 1% of sites, appearing and disappearing locally within and between breeding seasons. Early in spring, sparrows responded to landscape attributes most strongly within 400-m radii, settling in areas containing > 50% unburned prairie. Later in summer, sparrows responded to landscape attributes most strongly within 800-m radii, settling in areas containing > 50% unfragmented prairie, including sites burned earlier the same year. Sparrows avoided landscapes containing woody vegetation, disappeared from hayfields after mowing, and were most likely to inhabit landscapes containing Conservation Reserve Program (CRP) fields embedded within rangeland.

Conclusions

Landscape context influenced habitat selection at spatial scales broader than both the territory and habitat patch. Protecting contiguous prairies from agricultural conversion and woody encroachment, promoting CRP enrollment, and maintaining portions of undisturbed prairie in working rangelands each year are critical to reversing the conservation crisis in North America’s remaining grasslands. As landscape change alters natural areas worldwide, effective conservation requires suitable conditions for threatened species at multiple spatial scales.
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13.
14.

Context

Loss and fragmentation of semi-natural grasslands has critically affected many butterfly species in Europe. Habitat area and isolation can have strong effects on the local biodiversity but species may also be strongly affected by the surrounding matrix.

Objectives

We explored how different land cover types in the landscape explained the occurrence of butterfly species in semi-natural grasslands.

Methods

Using data from 476 semi-natural grasslands in Sweden, we analysed the effect of matrix composition on species richness and occurrence. Additionally, we analysed at which spatial scales butterflies responded to matrix types (forests, semi-natural grasslands, arable land and water).

Results

Forest cover showed the strongest positive effect on species richness, followed by semi-natural grasslands. Forest also had a positive effect on red-listed species at local scales. Responses to matrix composition were highly species-specific. The majority of the 30 most common species showed strong positive responses to the amount of forest cover within 200–500 m. There was a smaller group of species showing a positive response to arable land cover within 500–2000 m. Thirteen species showed positive responses to the amount of semi-natural grasslands, generally at larger scales (10–30 km).

Conclusions

Our study showed that surrounding forest is beneficial for many grassland butterfly species and that forests might mitigate the negative effects of habitat loss caused by agricultural intensification. Also, semi-natural grasslands were an important factor for species richness at larger spatial scales, indicating that a landscape consisting mainly of supporting habitats (i.e. forests) are insufficient to sustain a rich butterfly fauna.
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15.
16.

Context

Landscape modification is an important driver of biodiversity declines, yet we lack insight into how ongoing landscape change and legacies of historical land use together shape biodiversity.

Objectives

We examined how a history of agricultural land use and current forest fragmentation influence the abundance of red-backed salamanders (Plethodon cinereus). We hypothesized that historical agriculture and fragmentation cause changes in habitat quality and landscape structure that limit abundance.

Methods

We measured salamander abundance at 95 forested sites in New York, USA, and we determined whether sites were agricultural fields within the last five decades. We used a structural equation model to estimate relationships between historical agriculture and salamander abundance mediated by changes in forest vegetation, microclimate, and landscape structure.

Results

Historical agriculture affected salamander abundance by altering forest vegetation at a local scale and forest cover at a landscape scale. Abundance was lowest at post-agricultural sites with low woody vegetation, leaf litter depth, and canopy cover. Post-agricultural sites had limited forest cover in the surrounding landscape historically, and salamander abundance was positively related to historical forest cover, suggesting that connectivity to source populations affects colonization of regenerating forests. Abundance was also negatively related to current forest fragmentation.

Conclusions

Historical land use can have legacy effects on animal abundance on par with effects of ongoing landscape change. We showed that associations between animal abundance and historical land use can be driven by altered site conditions and surrounding habitat area, indicating that restoration efforts should consider local site conditions and landscape context.
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17.

Context

Wildfire activity in boreal forests is projected to increase dramatically in response to anthropogenic climate change. By altering the spatial arrangement of fuels, land-cover configuration may interact with climate change to influence fire-regime dynamics at landscape and regional scales.

Objectives

We evaluate how land cover interacts with weather conditions to influence boreal-forest burning from 2012 to 2014 in Alaska.

Methods

Using geospatial fire and land-cover data, we quantify relationships between area burned and land cover, and test whether observed patterns of burning differ from random under varying weather conditions and fire sizes.

Results

Mean summer moisture index was correlated with annual area burned (ρ = ?0.78, p < 0.01), the total number of fires (ρ = ?0.68, p = 0.01), and the number of large fires (>500 km2; ρ = ?0.58, p = 0.04). Area burned was related positively to percent cover of coniferous forest and woody wetlands, and negatively to percent cover of shrub scrub, dwarf scrub, and open water and barren areas. Fires preferentially burned coniferous forest, which represented 50.1 % of the area burned in warmer/drier summers and 40.3 % of area burned in cooler/wetter summers, compared to the 34.5 % (±4.2 %) expected by random selection of land-cover classes. Overall vegetation tended to burn more similarly to random in warmer/drier than cooler/wetter years.

Conclusions

Land cover exerted greater influences on boreal fire regimes when weather conditions were less favorable for forest burning. Reliable projections of boreal fire-regime change thus require consideration of the interactions between climate and land cover, as well as feedbacks from land-cover change.
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18.

Context

Resilience in fire-prone forests is strongly affected by landscape burn-severity patterns, in part by governing propagule availability around stand-replacing patches in which all or most vegetation is killed. However, little is known about drivers of landscape patterns of stand-replacing fire, or whether such patterns are changing during an era of increased wildfire activity.

Objectives

(a) Identify key direct/indirect drivers of landscape patterns of stand-replacing fire (e.g., size, shape of patches), (b) test for temporal trends in these patterns, and (c) anticipate thresholds beyond which landscape patterns of burn severity may change fundamentally.

Methods

We applied structural equation modeling to satellite burn-severity maps of fires in the US Northern Rocky Mountains (1984–2010) to test for direct and indirect (via influence on fire size and proportion stand-replacing) effects of climate/weather, vegetation, and topography on landscape patterns of stand-replacing fire. We also tested for temporal trends in landscape patterns.

Results

Landscape patterns of stand-replacing fire were strongly controlled by fire size and proportion stand-replacing, which were, in turn, controlled by climate/weather and vegetation/topography, respectively. From 1984 to 2010, the proportion of stand-replacing fire within burn perimeters increased from 0.22 to 0.27. Trends for other landscape metrics were not significant, but may respond to further increases proportion stand-replacing fire.

Conclusions

Fires from 1984 to 2010 exhibited tremendous heterogeneity in landscape patterns of stand-replacing fire, likely promoting resilience in burned areas. If trends continue on the current trajectory, however, fires may produce larger and simpler shaped patches of stand-replacing fire with more burned area far from seed sources.
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19.

Context

Pasture-woodlands are semi-natural landscapes that result from the combined influences of climate, management, and intrinsic vegetation dynamics. These landscapes are sensitive to future changes in land use and climate, but our ability to predict the impact on ecosystem service provisioning is limited due to the disparate scales in time and space that govern their dynamics.

Objectives

To develop a process-based model to simulate pasture-woodland landscapes and the provisioning of ecosystem services (i.e., livestock forage, woody biomass and landscape heterogeneity).

Methods

We modified a dynamic forest landscape model to simulate pasture-woodland landscapes in Switzerland. This involved including an annual herbaceous layer, selective grazing from cattle, and interactions between grazing and tree recruitment. Results were evaluated within a particular pasture, and then the model was used to simulate regional vegetation patterns and livestock suitability for a ~198,000 ha landscape in the Jura Vaudois region.

Results

The proportion of vegetation cover types at the pasture level (i.e., open, semi-open and closed forests) was well represented, but the spatial distribution of trees was only broadly similar. The entire Jura Vaudois region was simulated to be highly suitable for livestock, with only a small proportion being unsuitable due to steep slopes and high tree cover. High and low elevation pastures were equally suitable for livestock, as lower forage production at higher elevations was compensated by reduced tree cover.

Conclusions

The modified model is valuable for assessing landscape to regional patterns in vegetation and livestock, and offers a platform to evaluate how climate and management impact ecosystem services.
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20.

Context

Landscape and habitat filters are major drivers of biodiversity of small habitat islands by influencing dispersal and extinction events in plant metapopulations.

Objectives

We assessed the effects of landscape and habitat filters on the species richness, abundance and trait composition of grassland specialist and generalist plants in small habitat islands. We studied traits related to functional spatial connectivity (dispersal ability by wind and animals) and temporal connectivity (clonality and seed bank persistence) using model selection.

Methods

We sampled herbaceous plants, landscape (local and regional isolation) and habitat filters (inclination, woody encroachment and disturbance) in 82 grassland islands in Hungary.

Results

Isolation decreased the abundance of good disperser specialist plants due to the lack of directional vectors transferring seeds between suitable habitat patches. Clonality was an effective strategy, but persistent seed bank did not support the survival of specialist plants in isolated habitats. Generalist plants were unaffected by landscape filters due to their wide habitat breadth and high propagule availability. Clonal specialist plants could cope with increasing woody encroachment due to their high resistance against environmental changes; however, they could not cope with intensive disturbance. Steep slopes providing environmental heterogeneity had an overall positive effect on species richness.

Conclusions

Specialist plants were influenced by the interplay of landscape filters influencing their abundance and habitat filters affecting species richness. Landscape filtering by isolation influenced the abundance of specialist plants by regulating seed dispersal. Habitat filters sorted species that could establish and persist at a site by influencing microsite availability and quality.
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