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1.
Pamela Graff  Martín R. Aguiar 《Oikos》2011,120(7):1023-1030
Since many arid ecosystems are overstocked with domestic herbivores, biotic stress could have a stronger influence in modulating the balance of species interactions than expected from the stress gradient hypothesis (SGH). Here we tested a priori predictions about the effect of grazing on species interactions and fine scale spatial structure of grasses in water‐limited ecosystems. We used detailed vegetation mapping and spatial analysis, and performed a field experiment where the direct and indirect components of positive interactions were disentangled to provide evidence of links between process and pattern. We found associational resistance (biotic refuge) to be the dominant process in grazing situations, while competition, instead of direct facilitation, seemed to govern grass spatial patterns when herbivore pressure was relaxed. These results suggest that facilitation between grasses in arid communities may be related to herbivory rather than nurse plant effects. Associational resistance tends to have the strongest effect on spatial aggregation of species at intermediate grazing pressure. Results suggest that contrary to SGH, this physical clustering of species decreased when grazing pressure reached their maximum levels. Positive associations remained significant only when palatability differences between neighbours is large, suggesting that managing stocking rate is a key factor determining the persistence of herbivory refuges. These refuges are potential foci to initiate population recovery of high quality forage species in arid degraded areas.  相似文献   

2.
Heterogeneous disturbance patterns are fundamental to rangeland conservation and management because heterogeneity creates patchy vegetation, broadens niche availability, increases compositional dissimilarity, and enhances temporal stability of aboveground biomass production. Pyrodiversity is a popular concept for how variability in fire as an ecological disturbance can enhance heterogeneity, but mechanistic understanding of factors that drive heterogeneity is lacking. Mesic grasslands are examples of ecosystems in which pyrodiversity is linked strongly to broad ecological processes such as trophic interactions because grazers are attracted to recently burned areas, creating a unique ecological disturbance referred to as the fire–grazing interaction, or pyric herbivory. But several questions about the application of pyric herbivory remain: What proportion of a grazed landscape must burn, or how many patches are required, to create sufficient spatial heterogeneity and reduce temporal variability? How frequently should patches burn? Does season of fire matter? To bring theory into applied practice, we studied a gradient of grazed tallgrass prairie landscapes created by different sizes, seasons, and frequencies of fire, and used analyses sensitive to nonlinear trends. The greatest spatial heterogeneity and lowest temporal variability in aboveground plant biomass, and greatest plant functional group beta diversity, occurred in landscapes with three to four patches (25%–33% of area burned) and three‐ to four‐year fire return intervals. Beta diversity had a positive association with spatial heterogeneity and negative relationship with temporal variability. Rather than prescribing that these results constitute best management practices, we emphasize the flexibility offered by interactions between patch number and fire frequency for matching rangeland productivity and offtake to specific management goals. As we observed no differences across season of fire, we recommend future research focus on fire frequency within a moderate proportion of the landscape burned, and consider a wider seasonal burn window.  相似文献   

3.
In this paper, we address the question whether and through which mechanisms herbivores can induce spatial patterning in savanna vegetation, and how the role of herbivory as a determinant of vegetation patterning changes with herbivore density and the pre-existing pattern of vegetation. We thereto developed a spatially explicit simulation model, including growth of grasses and trees, vertical zonation of browseable biomass, and spatially explicit foraging by grazers and browsers. We show that herbivores can induce vegetation patterning when two key assumptions are fulfilled. First, herbivores have to increase the attractiveness of a site while foraging so that they will revisit this site, e.g. through an increased availability or quality of forage. Second, foraging should be spatially explicit, e.g. when foraging at a site influences vegetation at larger spatial scales or when vegetation at larger spatial scales influences the selection and utilisation of a site. The interaction between these two assumptions proved to be crucial for herbivores to produce spatial vegetation patterns, but then only at low to intermediate herbivore densities. High herbivore densities result in homogenisation of vegetation. Furthermore, our model shows that the pre-existing spatial pattern in vegetation influences the process of vegetation patterning through herbivory. However, this influence decreases when the heterogeneity and dominant scale of the initial vegetation decreases. Hence, the level of adherence of the herbivores to forage in pre-existing patches increases when these pre-existing patches increase in size and when the level of vegetation heterogeneity increases. The findings presented in this paper, and critical experimentation of their ecological validity, will increase our understanding of vegetation patterning in savanna ecosystems, and the role of plant–herbivore interactions therein.  相似文献   

4.
There is a long tradition of grazing by semi‐domestic reindeer and sheep in alpine and sub‐alpine Scandinavian habitats, but present management regimes are questioned from a conservation point of view. In this review we discuss plant diversity patterns in the Scandinavian mountains in a global, regional and local perspective. The main objective was to identify processes that influence diversity at different spatial scales with a particular focus on grazing. In a global perspective the species pool of the Scandinavian mountains is limited. partly reflecting the general latitudinal decline of species but also historical and ecological factors operating after the latest glaciation. At the local scale, both productivity and disturbance are primary factors structuring diversity, but abiotic factors such as soil pH, snow distribution and temperature are also important. Although evidence is scarce, grazing favours local species richness in productive habitats, whereas species richness decreases with grazing when productivity is low. Regional patterns of plant diversity is set by, 1) the species pool. 2) the heterogeneity and fragmentation of communities, and 3) local diversity of each plant community. We suggest that local shifts in community composition depend both on the local grazing frequency and the return‐time of the plant community after a grazing session. In addition, an increasing number of grazing‐modified local patches homogenises the vegetation and is likely to reduce the regional plant diversity. The time scale of local shifts in community composition depends on plant colonisation and persistence, From a mechanistic point of view, diversity patterns at a regional scale also depend on the regional dynamics of single species. Colonisation is usually a slow and irregular process in alpine environments, whereas the capacity for extended local persistence is generally high. Although the poor knowledge of plant regional dynamics restricts our understanding of how grazing influences plant diversity, we conclude that grazing is a key process for maintaining biodiversity in the Scandinavian mountains.  相似文献   

5.
Soil properties (i.e. soil organic carbon, SOC; soil organic nitrogen, SON; and soil C/N ratio) and vegetation in a semiarid grassland of Inner Mongolia, northern China, were studied with the method of geostatistical analysis. We examined the spatial heterogeneity of soil and plants, and possible impacts of land use on their heterogeneity and on the relationship between soil resources and plant richness. Land use affected small scale spatial heterogeneity in plants and soil. SOC, SON and C/N ratio displayed autocorrelation over a range of ~2 m under most circumstances on sites where livestock grazing had been excluded. The uncontrolled grazing site (UG, i.e. unregulated grazing by excessive livestock) displayed an increased range of spatial autocorrelation and the total amount of variability in soil nitrogen over the other land use types. Plant life forms and plant species exhibited spatial autocorrelation over a range of about 2 m on the grazing exclusion (GE) and mowed (MW) sites, while pattern of spatial autocorrelation for several less common species on the UG site were difficult to predict. Plant species richness was positively related with spatial heterogeneity of SOC, SON and C/N on both GE and MW sites, and with only SOC heterogeneity on the UG site. These suggest that spatial soil heterogeneity plays an active role in maintaining plant species richness. However, we call for caution in generalization of the control of spatial soil heterogeneity over plant richness when multiple modes of disturbances are present, as we found in this study that higher total amount of variation in soil nitrogen and C/N ratio on the over-grazed UG site did not lead to increased plant species richness, and that land use had apparent effects on the patterns of spatial heterogeneity in both vegetation and soil.  相似文献   

6.
Herbivores influence spatial heterogeneity in soil resources and vegetation in ecosystems. Despite increasing recognition that spatial heterogeneity can drive species richness at different spatial scales, few studies have quantified the effect of grazing on spatial heterogeneity and species richness simultaneously. Here we document both these variables in a rabbit-grazed grassland. We measured mean values and spatial patterns of grazing intensity, rabbit droppings, plant height, plant biomass, soil water content, ammonia and nitrate in sites grazed by rabbits and in matched, ungrazed exclosures in a grassland in southern England. Plant species richness was recorded at spatial scales ranging between 0.0001 and 150 m(2). Grazing reduced plant height and plant biomass but increased levels of ammonia and nitrate in the soil. Spatial statistics revealed that rabbit-grazed sites consisted of a mixture of heavily grazed patches with low vegetation and nutrient-rich soils (lawns) surrounded by patches of high vegetation with nutrient-poor soils (tussocks). The mean patch size (range) in the grazed controls was 2.1 +/- 0.3 m for vegetation height, 3.8 +/- 1.8 m for soil water content and 2.8 +/- 0.9 m for ammonia. This is in line with the patch sizes of grazing (2.4 +/- 0.5 m) and dropping deposition (3.7 +/- 0.6 m) by rabbits. In contrast, patchiness in the ungrazed exclosures had a larger patch size and was not present for all variables. Rabbit grazing increased plant species richness at all spatial scales. Species richness was negatively correlated with plant height, but positively correlated to the coefficient of variation of plant height at all plot sizes. Species richness in large plots (<25 m(2)) was also correlated to patch size. This study indicates that the abundance of strong competitors and the nutrient availability in the soil, as well as the heterogeneity and spatial pattern of these factors may influence species richness, but the importance of these factors can differ across spatial scales.  相似文献   

7.
Both arthropods and large grazing herbivores are important components and drivers of biodiversity in grassland ecosystems, but a synthesis of how arthropod diversity is affected by large herbivores has been largely missing. To fill this gap, we conducted a literature search, which yielded 141 studies on this topic of which 24 simultaneously investigated plant and arthropod diversity. Using the data from these 24 studies, we compared the responses of plant and arthropod diversity to an increase in grazing intensity. This quantitative assessment showed no overall significant effect of increasing grazing intensity on plant diversity, while arthropod diversity was generally negatively affected. To understand these negative effects, we explored the mechanisms by which large herbivores affect arthropod communities: direct effects, changes in vegetation structure, changes in plant community composition, changes in soil conditions, and cascading effects within the arthropod interaction web. We identify three main factors determining the effects of large herbivores on arthropod diversity: (i) unintentional predation and increased disturbance, (ii) decreases in total resource abundance for arthropods (biomass) and (iii) changes in plant diversity, vegetation structure and abiotic conditions. In general, heterogeneity in vegetation structure and abiotic conditions increases at intermediate grazing intensity, but declines at both low and high grazing intensity. We conclude that large herbivores can only increase arthropod diversity if they cause an increase in (a)biotic heterogeneity, and then only if this increase is large enough to compensate for the loss of total resource abundance and the increased mortality rate. This is expected to occur only at low herbivore densities or with spatio‐temporal variation in herbivore densities. As we demonstrate that arthropod diversity is often more negatively affected by grazing than plant diversity, we strongly recommend considering the specific requirements of arthropods when applying grazing management and to include arthropods in monitoring schemes. Conservation strategies aiming at maximizing heterogeneity, including regulation of herbivore densities (through human interventions or top‐down control), maintenance of different types of management in close proximity and rotational grazing regimes, are the most promising options to conserve arthropod diversity.  相似文献   

8.
放牧和刈割条件下草山草坡群落空间异质性分析   总被引:13,自引:1,他引:12  
采用变异矩分析和分形方法,研究了草山草坡群落在放牧和刈割条件下的空间异质性及空间自相关性,结果表明,群落空间格局有尺度依赖性,刈割条件下空物异质性及空间相关性弱,多样性梯度即β多样性小,放牧消除地形引起的样地差异,因而使空间异质性简单化。  相似文献   

9.
微斑块变化是草原退化过程中的活跃成分。分析了呼伦贝尔克氏针茅草原逆行演替过程中微斑块土壤全碳、全氮和碱解氮含量的空间异质性,提出了"养分聚集效应"的概念。研究结果表明:随着群落退化演替的加剧,土壤全碳、全氮和碱解氮的含量均表现为演替前期演替后期演替中期(P0.05)。从土壤全碳、全氮和碱解氮的变异系数和变异函数综合分析来看,10 cm×10 cm微尺度上,草原退化演替过程中土壤全碳、全氮和碱解氮的空间异质性具有明显的不一致性;全碳的空间异质性表现为演替中期演替前期演替后期,全氮表现为演替后期演替前期演替中期,碱解氮表现为演替中期演替后期演替前期。草原退化过程中土壤养分在微斑块上的富积和迁移表现出尺度依赖性和变异性。  相似文献   

10.
Spatial patterns of species diversity have important influences on the functioning of ecosystems, and the effect of livestock grazing on spatial heterogeneity can differ depending on the scale of the analysis. This study examined the effects of grazing on the spatial patterns of species distributions and whether the effects of grazing on the spatial distributions of a species vary with the scale of the analysis. Data were collected at three locations in the subalpine grasslands of Ordesa-Monte Perdido National Park and Aísa Valley, Central Pyrenees, Spain, which differed in mean stocking rates. Aspect explained about one-third of the environmental variation in species distributions. In flat areas, spatial variation in species composition varied with grazing intensity at two scales. At a coarse scale (among vegetation patches), grazing promoted patchiness, and among-transect variation in species diversity and grazing intensity were positively correlated. At a fine scale (within vegetation patches), the disruption of the self-organizing processes of the species spatial distributions resulted in a reduction in the long-range spatial autocorrelations of some of the characteristic species and in the homogenization of species spatial distributions. The presence of encroaching Echinospartum horridum had a significant influence on the effect of grazing on south-facing grassland slopes.  相似文献   

11.
The relative importance of extrinsic and intrinsic causes of variability is among the oldest unresolved problems in ecology. However, the interaction between large-scale intrinsic variability in species abundance and environmental heterogeneity is still unknown. We use a metacommunity model with disturbance-recovery dynamics to resolve the interaction between scales of environmental heterogeneity, biotic processes and of intrinsic variability. We explain how population density increases with environmental variability only when its scale matches that of intrinsic patterns of abundance, through their ability to develop in heterogeneous environments. Succession dynamics reveals how the strength of local species interactions, through its control of intrinsic variability, can in turn control the scale of metapopulation response to environmental scales. Our results show that the environment and species density might fail to show any correlation despite their strong causal association. They more generally suggest that the spatial scale of ecological processes might not be sufficient to build a predictive framework for spatially heterogeneous habitats, including marine reserve networks.  相似文献   

12.
The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.  相似文献   

13.
Questions: Does grazing have the same effect on plant species richness at different spatial scales? Does the effect of spatial scale vary under different climatic conditions and vegetation types? Does the slope of the species‐area curve change with grazing intensity similarly under different climatic conditions and vegetation types? Location: Pastures along a climatic gradient in northeastern Spain. Methods: In zones under different regimes of sheep grazing (high‐, low‐pressure, abandonment), plant species richness was measured in different plot sizes (from 0.01 to 100 m2) and the slope of the species‐area curves was calculated. The study was replicated in five different locations along a climatic gradient from lowland semi‐arid rangelands to upland moist grasslands. Results: Species richness tended to increase with grazing intensity at all spatial scales in the moist upland locations. On the contrary, in the most arid locations, richness tended to decrease, or remain unchanged, with grazing due to increased bare soil. Grazing differentially affected the slope (z) of the species‐area curve (power function S=c Az) in different climatic conditions: z tended to increase with grazing in arid areas and decrease in moist‐upland ones. ß‐diversity followed similar pattern as z. Conclusions: Results confirm that the impact of grazing on plant species richness are spatial‐scale dependent. However, the effects on the species‐area relationship vary under different climatic conditions. This offers a novel insight on the patterns behind the different effects of grazing on diversity in moist vs. arid conditions reported in the literature. It is argued that the effect of spatial scale varies because of the different interaction between grazing and the intrinsic spatial structure of the vegetation. Variations in species‐area curves with grazing along moisture gradients suggest also a different balance of spatial components of diversity (i.e. a‐ and ß‐diversity).  相似文献   

14.
Veen GF  Geuverink E  Olff H 《Oecologia》2012,168(2):511-518
Aboveground and belowground organisms influence plant community composition by local interactions, and their scale of impact may vary from millimeters belowground to kilometers aboveground. However, it still poorly understood how large grazers that select their forage on large spatial scales interact with small-scale aboveground–belowground interactions on plant community heterogeneity. Here, we investigate how cattle (Bos taurus) modify the effects of interactions between yellow meadow ants (Lasius flavus) and European brown hares (Lepus europaeus) on the formation of small-scale heterogeneity in vegetation composition. In the absence of cattle, hares selectively foraged on ant mounds, while under combined grazing by hares and cattle, vertebrate grazing pressure was similar on and off mounds. Ant mounds that were grazed by only hares had a different plant community composition compared to their surroundings: the cover of the grazing-intolerant grass Elytrigia atherica was reduced on ant mounds, whereas the relative cover of the more grazing-tolerant and palatable grass Festuca rubra was enhanced. Combined grazing by hares and cattle, resulted in homogenization of plant community composition on and off ant mounds, with high overall cover of F. rubra. We conclude that hares can respond to local ant–soil–vegetation interactions, because they are small, selective herbivores that make their foraging decisions on a local scale. This results in small-scale plant patches on mounds of yellow meadow ants. In the presence of cattle, which are less selective aboveground herbivores, local plant community patterns triggered by small-scale aboveground–belowground interactions can disappear. Therefore, cattle modify the consequences of aboveground–belowground interactions for small-scale plant community composition.  相似文献   

15.
Recent theoretical studies have shown that spatial redistribution of surface water may explain the occurrence of patterns of alternating vegetated and degraded patches in semiarid grasslands. These results implied, however, that spatial redistribution processes cannot explain the collapse of production on coarser scales observed in these systems. We present a spatially explicit vegetation model to investigate possible mechanisms explaining irreversible vegetation collapse on coarse spatial scales. The model results indicate that the dynamics of vegetation on coarse scales are determined by the interaction of two spatial feedback processes. Loss of plant cover in a certain area results in increased availability of water in remaining vegetated patches through run-on of surface water, promoting within-patch plant production. Hence, spatial redistribution of surface water creates negative feedback between reduced plant cover and increased plant growth in remaining vegetation. Reduced plant cover, however, results in focusing of herbivore grazing in the remaining vegetation. Hence, redistribution of herbivores creates positive feedback between reduced plant cover and increased losses due to grazing in remaining vegetated patches, leading to collapse of the entire vegetation. This may explain irreversible vegetation shifts in semiarid grasslands on coarse spatial scales.  相似文献   

16.
This research investigates the formation of banded vegetation patterns on hillslopes affected by interactions between sediment deposition and vegetation growth. The following two perspectives in the formation of these patterns are taken into consideration: (a) increased sediment deposition from plant interception, and (b) reduced plant biomass caused by sediment accumulation. A spatial model is proposed to describe how the interactions between sediment deposition and vegetation growth promote self-organization of banded vegetation patterns. Based on theoretical and numerical analyses of the proposed spatial model, vegetation bands can result from a Turing instability mechanism. The banded vegetation patterns obtained in this research resemble patterns reported in the literature. Moreover, measured by sediment dynamics, the variation of hillslope landform can be described. The model predicts how treads on hillslopes evolve with the banded patterns. Thus, we provide a quantitative interpretation for coevolution of vegetation patterns and landforms under effects of sediment redistribution.  相似文献   

17.
植物群落中物种小尺度空间结构研究   总被引:27,自引:0,他引:27       下载免费PDF全文
植物群落中, 物种小尺度空间结构影响着种群或群落的动态及有关的生态学过程。植物主要是和它同种或异种的邻近个体相互作用,植物个体周围的局部环境和大空间尺度下群落的平均水平是完全不同的。群落中的许多过程都影响小尺度空间结构的形成和动态,同样,局部空间结构反过来影响着植物的生长、更新和凋亡等重要过程。鉴于目前对小尺度空间结构进行的大量研究以及其重要性,有必要对其研究进展进行适当的总结,以期明确将来进一步的研究方向。该文以此为出发点,首先介绍了植物群落中物种小尺度空间结构产生的6个原因:1)生境的空间异质性;2)植物繁殖体的传播;3)植物之间的相互作用;4)生物环境(动物和微生物)的作用;5)外界干扰的作用;6)多因子综合作用。然后阐述了小尺度空间结构意义及对生物多样性、植物种群遗传学和恢复生态学研究的影响。最后对目前物种小尺度空间结构研究存在的几点问题及将来的研究方向作以下归纳:1)大尺度植被动态的研究应该整合小尺度空间结构的信息;2)不论从生物学还是生态学上来讲, 植被小尺度空间结构的研究应该把植物作为中心,确定适当的尺度和采取合理的空间统计方法;3)充分重视小尺度空间结构在退化生态系统恢复中的应用意义;4)注重从小尺度的局部格局研究入手对群落总体特征进行整合;5)植物群落动态研究中,物种小尺度空间结构与平均场假说相结合的必要性。  相似文献   

18.
Non‐random patterns of species segregation and aggregation within ecological communities are often interpreted as evidence for interspecific interactions. However, it is unclear whether theoretical models can predict such patterns and how environmental factors may modify the effects of species interactions on species co‐occurrence. Here we extend a spatially explicit neutral model by including competitive effects on birth and death probabilities to assess whether competition alone is able to produce non‐random patterns of species co‐occurrence. We show that transitive and intransitive competitive hierarchies alone (in the absence of environmental heterogeneity) are indeed able to generate non‐random patterns with commonly used metrics and null models. Moreover, even weak levels of intransitive competition can increase local species richness. However, there is no simple rule or consistent directional change towards aggregation or segregation caused by competitive interactions. Instead, the spatial pattern depends on both the type of species interaction and the strength of dispersal. We conclude that co‐occurrence analysis alone may not able to identify the underlying processes that generate the patterns.  相似文献   

19.
分析不同放牧强度下植物群落中物种的空间分布特征, 有助于阐明群落在放牧胁迫下的演替规律。该研究基于幂函数法则, 探讨了不同放牧强度下短花针茅(Stipa breviflora)荒漠草原群落植物的频率和空间异质性。结果表明: 不同放牧强度下物种空间分布与幂函数法则能很好地吻合; 不同物种空间异质性具有特异性, 随着放牧强度的增加, 提高群落空间异质性的物种分别由无芒隐子草(Cleistogenes songorica)、冷蒿(Artemisia frigida)、短花针茅、银灰旋花(Convolvulus ammannii)等多个物种逐渐转变为以无芒隐子草、短花针茅为主的少数物种, 同时, 物种空间异质性大于群落空间异质性的物种数逐渐减少。  相似文献   

20.
Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing‐induced legacy effects in the soil and we explore whether these plant responses can help us to understand long‐term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field‐conditioned soils from grazed and ungrazed grassland. We relate these responses to long‐term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing‐induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback.  相似文献   

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