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1.
Wood betony, Orobanchaceae (Pedicularis canadensis) and bastard toadflax, Santalaceae (Comandra umbellata) are two root‐hemiparasitic plant species found in tallgrass prairie communities. Natural resource managers are interested in utilizing these species as “pseudograzers” in grasslands to reduce competitively dominant grasses and thereby increase ecological diversity and quality in prairie restorations and urban plantings. We performed an observational field study at 5 tallgrass prairie sites to investigate the association of hemiparasite abundance with metrics of phylogenetic and ecological diversity, as well as floristic quality. Although no reduction in C4 grasses was detected, there was a significant association between hemiparasite abundance and increased floristic quality at all 5 sites. Hemiparasite abundance and species richness were positively correlated at one restoration site. In a greenhouse mesocosm experiment, we investigated response to parasitism by P. canadensis in 6 species representing different plant functional groups of the tallgrass prairie. The annual legume partridge pea, Fabaceae (Chamaecrista fasciculata) had the greatest significant dry biomass reduction among 6 host species, but the C4 grass big bluestem, Poaceae (Andropogon gerardii) had significantly greater aboveground biomass when grown with the hemiparasite. Overall, host species biomass as a total community was significantly reduced in mesocosms, consistent with other investigations that demonstrate influence on community structure by hemiparasitic plant species. Although hemiparasites were not acting as pseudograzers, they have the potential to influence community structure in grassland restorations and remnants.  相似文献   

2.
Ants are highly influential organisms in terrestrial ecosystems, including the tallgrass prairie, one of the most endangered ecosystems in North America. Through their tunneling, ants affect soil properties and resource availability for animals and plants. Ants also have important ecological roles as consumers of plant tissue and seeds. In the last several decades, various organizations, agencies, and agricultural producers have attempted to create wildlife habitat or reduce soil erosion by seeding thousands of hectares of bare cropland in the central United States with tallgrass prairie seed mixes. Although initially, monitoring of these restorations and of unplowed prairie remnants focused on plants and birds, in recent years the response of invertebrates such as ants has increasingly been the subject of research. An understanding of tallgrass prairie ant communities can help land managers and scientists better monitor the ecological condition of tallgrass prairie and guide management and restoration efforts. Here I review our current knowledge of ant species found within tallgrass prairie, their ecological roles, and their response to management.  相似文献   

3.
Ecological traps occur when habitat selection and habitat suitability (measured in terms of fitness) are decoupled. We developed a graphical model based on isodar theory to distinguish between an ideal distribution and an ecological trap. We tested the model's predictions using data on breeding bird populations in managed tallgrass prairie in Oklahoma. Between 1992 and 1996 we monitored success for 2600 nests of 26 breeding species in undisturbed, grazed, and burned and grazed plots. We also sampled arthropod biomass and nest predator abundance. Using the isodar model we determined that managed plots are ecological traps: compared with success on plots left undisturbed, nest success on plots that were only grazed was lower, and success on plots that were burned and grazed was substantially lower. Yet birds preferred to nest on managed plots, where arthropod abundance was measurably higher. Reptiles were the most abundant taxon of nest predators, and their abundance was highest in managed plots. Consequently, tree-nesting species had higher nest success than shrub- and ground-nesting birds. Nest success also increased with tree height. We concluded that isodar theory is a useful tool for detecting ecological traps if any component of fitness is measured in addition to animal densities. Our study also suggests that (1) human modification of the environment may alter simultaneously food and predator abundance, (2) the former affects nest site selection and the latter nest success, and (3) such ecosystems are likely to become traps for breeding birds.  相似文献   

4.
Theory has predicted, and many experimental studies have confirmed, that resident plant species richness is inversely related to invisibility. Likewise, potential invaders that are functionally similar to resident plant species are less likely to invade than are those from different functional groups. Neither of these ideas has been tested in the context of an operational prairie restoration. Here, we tested the hypotheses that within tallgrass prairie restorations (1) as seed mix species richness increased, cover of the invasive perennial forb, Canada thistle (Cirsium arvense) would decline; and (2) guilds (both planted and arising from the seedbank) most similar to Canada thistle would have a larger negative effect on it than less similar guilds. Each hypothesis was tested on six former agricultural fields restored to tallgrass prairie in 2005; all were within the tallgrass prairie biome in Minnesota, USA. A mixed-model with repeated measures (years) in a randomized block (fields) design indicated that seed mix richness had no effect on cover of Canada thistle. Structural equation models assessing effects of cover of each planted and non-planted guild on cover of Canada thistle in 2006, 2007, and 2010 revealed that planted Asteraceae never had a negative effect on Canada thistle. In contrast, planted cool-season grasses and non-Asteraceae forbs, and many non-planted guilds had negative effects on Canada thistle cover. We conclude that early, robust establishment of native species, regardless of guild, is of greater importance in resistance to Canada thistle than is similarity of guilds in new prairie restorations.  相似文献   

5.
Restored grasslands comprise an ever‐increasing proportion of grasslands in North America and elsewhere. However, floristic studies of restored grasslands indicate that our ability to restore plant communities is limited. Our goal was to assess the effectiveness of restoration seeding for recovery of key plant community components on former exotic, cool‐season pastures using a chronosequence of six restoration sites and three nearby remnant tallgrass prairie sites in West‐Central Iowa. We assessed trends in Simpson's diversity and evenness, richness and abundance of selected native and exotic plant guilds, and mean coefficient of conservatism (mean C). Simpson's diversity and evenness and perennial invasive species abundance all declined with restoration site age. As a group, restoration sites had greater richness of native C3 species with late phenology, but lower richness and abundance of species with early phenology relative to remnant sites. Total native richness, total native abundance (cover), mean C, and abundance of late phenology C3 plants were similar between restoration and remnant sites. Observed declines in diversity and evenness with restoration age reflect increases in C4 grass abundance rather than absolute decreases in the abundance of perennial C3 species. In contrast to other studies, restoration seeding appears to have led to successful establishment of tallgrass prairie species that were likely to be included in seeding mixtures. While several floristic measures indicate convergence of restoration and remnant sites, biodiversity may be further enhanced by including early phenology species in seeding mixes in proportion to their abundance on remnant prairies.  相似文献   

6.
The alteration and fragmentation of native tallgrass prairie in the Midwestern United States has created a need to identify other land types with the ability to support grassland butterfly species. This study examines butterfly usage of marginal grasslands, which consist of semi-natural grasslands existing within in a larger agricultural matrix, compared to grasslands managed for conservation of prairie species. Using generalized linear mixed models we analyzed how land purpose (marginal vs. conservation grasslands) affected butterfly abundance. We found grassland butterfly species to be significantly more common on conservation grasslands, whereas generalist species were significantly more common on marginal grasslands. Results of ordination analyses indicated that while many species used both types of habitats, butterfly species assemblages were distinct between habitat types and that edge to interior ratio and the floristic quality index of sites were important habitat characteristics driving this distinction. Within conservation grasslands we examined the relationship between butterfly abundance and the planting diversity used in restoring each site. We found higher diversity restorations hosted more individuals of butterflies considered habitat generalists, as well as species considered to be of conservation concern.  相似文献   

7.
Understanding the degree to which species assemblages naturally vary over time will be critically important when assessing whether direct management effects or contingency is responsible for species gain or loss. In this study, we tested three predictions related to short‐term variation in prairie moth communities: (1) communities would only exhibit significant temporal variation in newly restored sites (1–3 years old); (2) prairie size and age would positively influence community reassembly, with larger, older restorations sampling a greater proportion of the regional species pool; and (3) older restorations (7–10 years old) would have yet to converge on the community composition of prairie remnants. Moths were sampled from 13 Tallgrass prairie restorations and remnants in central Iowa in 2004–2005. Repeated measures analysis of variance revealed significant effects of sampling year on moth species richness and abundance as well as on the richness of two functional groups, but difference among prairie types was only observed in 2005. Rarefaction analysis revealed that older restorations and prairie remnants supported higher species richness compared to recently planted sites, and nonmetric, multidimensional scaling ordination indicated that restorations older than 7 years were clearly converging on the species composition of remnants. These results suggest that moth communities in restorations and remnants are highly variable in time but that as restorations age, they appear to reaccumulate moth species found in prairie remnants. The long‐term persistence of a particular species assemblage within a given site, however, might be a difficult endpoint to attain in central Iowa prairies because of significant annual variation in species occurrence.  相似文献   

8.
Plant species richness in rural landscapes of northern Europe has been positively influenced by traditional management for millennia. Owing to abandonment of these practices, the number of species‐rich semi‐natural grasslands has decreased, and remaining habitats suffer from deterioration, fragmentation, and plant species decline. To prevent further extinctions, restoration efforts have increased during the last decades, by reintroducing grazing in former semi‐natural grasslands. To assess the ecological factors that might influence the outcome of such restorations, we made a survey of semi‐natural grasslands in Sweden that have been restored during the last decade. We investigated how plant species richness, species density, species composition, and abundance of 10 species that are indicators of grazing are affected by (1) the size of the restored site, (2) the time between abandonment of grazing and restoration, (3) the time elapsed since restoration, and (4) the abundance of trees and shrubs at the restored site. Only two factors, abundance of trees and shrubs and time since restoration, were positively associated with total species richness and species density per meter square at restored sites. Variation in species composition among restored sites was not related to any of the investigated factors. Species composition was relatively similar among sites, except in mesic/wet grasslands. The investigated factors had small effects on the abundance of the grazing‐indicator species. Only Campanula rotundifolia responded to restoration with increasing abundance and may thus be a suitable indicator of improved habitat quality. In conclusion, positive effects on species richness may appear relatively soon after restoration, but rare, short‐lived species are still absent. Therefore, remnant populations in surrounding areas may be important in fully recreating former species richness and composition.  相似文献   

9.
Species abundance data are critical for testing ecological theory, but obtaining accurate empirical estimates for many taxa is challenging. Proxies for species abundance can help researchers circumvent time and cost constraints that are prohibitive for long‐term sampling. Under simple demographic models, genetic diversity is expected to correlate with census size, such that genome‐wide heterozygosity may provide a surrogate measure of species abundance. We tested whether nucleotide diversity is correlated with long‐term estimates of abundance, occupancy and degree of ecological specialization in a diverse lizard community from arid Australia. Using targeted sequence capture, we obtained estimates of genomic diversity from 30 species of lizards, recovering an average of 5,066 loci covering 3.6 Mb of DNA sequence per individual. We compared measures of individual heterozygosity to a metric of habitat specialization to investigate whether ecological preference exerts a measurable effect on genetic diversity. We find that heterozygosity is significantly correlated with species abundance and occupancy, but not habitat specialization. Demonstrating the power of genomic sampling, the correlation between heterozygosity and abundance/occupancy emerged from considering just one or two individuals per species. However, genetic diversity does no better at predicting abundance than a single day of traditional sampling in this community. We conclude that genetic diversity is a useful proxy for regional‐scale species abundance and occupancy, but a large amount of unexplained variation in heterozygosity suggests additional constraints or a failure of ecological sampling to adequately capture variation in true population size.  相似文献   

10.
Belowground bud production mediates many responses of tallgrass prairie grasses to temporal environmental variation, and clonal spread contributes to dominance in successional plant communities. However, we know little about the importance of these regenerative traits for vegetation establishment in grassland restoration and about their regional variability. Further, geographic seed or transplant sources affect plant performance. We investigated the effects of seed source on the number of belowground buds per tiller, clonal spread (total rhizome length adjusted for plant biomass), and population establishment in the native C4 grass Sorghastrum nutans sown into tallgrass prairie restorations using reciprocal common gardens in three sites (in Nebraska, Kansas, and Oklahoma). We characterized buds per tiller and clonal spread at the plant level and related both traits to population establishment at the restoration plot level. Plant-level bud production significantly differed among seed sources and common garden sites, and was positively associated with plot-level establishment (density and canopy cover), but plant-level clonal spread did not differ among seed sources or common garden sites and was not associated with the plot-level establishment. Our results are among the first to link bud production, which varied among source populations, to differences in plant population establishment. These results suggest that we should more often consider differences among sources in traits closely tied to post-seeding population establishment. The use of sources that possess such traits may facilitate successful establishment in restoration settings.  相似文献   

11.
Keystone species restoration, or the restoration of species whose effect on an ecosystem is much greater than their abundance would suggest, is a central justification for many wildlife reintroduction projects globally. Following restoration, plains bison (Bison bison L.) have been identified as a keystone species in the tallgrass prairie ecoregion, but we know of no research to document similar effects in the mixed‐grass prairie where restoration efforts are ongoing. This study addresses whether Northern Great Plains (NGP) mixed‐grass prairie plant communities exhibit traits consistent with four central keystone effects documented for bison in the tallgrass prairie. We collected species composition, diversity, abundance, bare ground cover, and plant height data in three treatments: where livestock (Bos taurus L.) continuously grazed, livestock were removed for 10 years, and bison have been introduced and resident for 10 years. We observed mixed support for bison acting as keystone species in this system. Supporting the keystone role of bison, we observed higher species richness and compositional heterogeneity (β‐diversity) in the bison treatment than either the livestock retention or livestock removal treatments. However, we observed comparable forb, bare ground, and plant height heterogeneity between bison‐restored sites and sites where livestock were retained, contradicting reported keystone effects in other systems. Our results suggest that after 10 years of being restored, bison partially fulfill their role as a keystone species in the mixed‐grass prairie, and we encourage continued long‐term data collection to evaluate their influence in the NGP.  相似文献   

12.
Feather mites are obligatory ectosymbionts of birds that primarily feed on the oily secretions from the uropygial gland. Feather mite abundance varies within and among host species and has various effects on host condition and fitness, but there is little consensus on factors that drive variation of this symbiotic system. We tested hypotheses regarding how within‐species and among‐species traits explain variation in both (1) mite abundance and (2) relationships between mite abundance and host body condition and components of host fitness (reproductive performance and apparent annual survival). We focused on two closely related (Parulidae), but ecologically distinct, species: Setophaga cerulea (Cerulean Warbler), a canopy dwelling open‐cup nester, and Protonotaria citrea (Prothonotary Warbler), an understory dwelling, cavity nester. We predicted that feather mites would be more abundant on and have a more parasitic relationship with P. citrea, and within P. citrea, females and older individuals would harbor greater mite abundances. We captured, took body measurements, quantified feather mite abundance on individuals’ primaries and rectrices, and monitored individuals and their nests to estimate fitness. Feather mite abundance differed by species, but in the opposite direction of our prediction. There was no relationship between mite abundance and any measure of body condition or fitness for either species or sex (also contrary to our predictions). Our results suggest that species biology and ecological context may influence mite abundance on hosts. However, this pattern does not extend to differential effects of mites on measures of host body condition or fitness.  相似文献   

13.
14.
We assessed the determinants and consequences of the expansion of Juniperus virginiana L. (red cedar) populations into central US grasslands using historical aerial photos and field measurements of forest extent, tree growth, fire-induced mortality, and responses in herbaceous species diversity and productivity. Photos from northeast Kansas dating back to 1956 indicate that native tallgrass prairie can be converted to closed-canopy red cedar forest in as little as 40 years (a 2.3% increase in forest cover per year). Mean tree density in 21 forested sites ranged from 130 to 3500 trees/ha, with most sites at more than 800 trees/ha. In younger stands, maximum growth rates of individual red cedar trees exceeded 20 cm/y in height. Land management practices were critical to the establishment and growth of red cedar forest. Grazing reduced the fuel loads by more than 30% in tallgrass prairie. Based on measurements of mortality for more than 1800 red cedar trees, fire-induced mortality in grazed areas averaged 31.6% versus more than 90% at ungrazed sites. When tallgrass prairie was converted to red cedar forest, herbaceous species diversity and productivity were drastically reduced, and most grassland species were virtually eliminated. Consequently, community structure shifted from dominance by herbaceous C4 species to evergreen woody C3 species; this shift is likely to be accompanied by alterations in carbon storage and other ecosystem processes in a relatively short time period. Here we present a conceptual model that integrates the ecological and socioeconomic factors that underlie the conversion of grassland to red cedar forest. Received 29 August 2001; accepted 5 February 2002.  相似文献   

15.
Attempting to control invasive plant species in tallgrass prairie restorations is time-consuming and costly, making improved approaches for predicting and reducing invasion imperative. Both biotic and abiotic factors mediate plant invasions, and can potentially be used by restoration managers to reduce invasion rates. Biotic factors such as plant species richness and phylogenetic diversity of the native community may impact invasion. Relatedness of invading species to those in recipient communities has also been shown to influence invasion success. However, the direction of this influence is variable, reflecting Darwin’s Naturalization Conundrum. Abiotic factors such as fire regime and soil factors may impact invasion by selecting against invasive species or indicating suitable habitats for them. We surveyed 17 tallgrass prairie restorations in Illinois, USA, to investigate the effects of biotic and abiotic factors on invasion by non-native plant species at two different scales. We predicted we would find support for Darwin’s Naturalization Hypothesis at the plot (neighborhood) scale with invasion by distantly related species, and find support for the Pre-adaptation Hypothesis at the site scale. We hypothesized that biotic factors would exert more influence at the neighborhood scale, while abiotic factors would be more influential at a coarser site scale. Contrary to our expectations, at the neighborhood scale we found that closely related invasive species are more likely to invade, supporting the Pre-adaptation Hypothesis. We found that native species richness and age of restoration were negatively correlated with invasion. At the site scale, soil organic matter [SOM] concentrations and heterogeneity in SOM were positively associated with the number of invasive species while pH heterogeneity was negatively associated. Restoration practitioners may be able to reduce plant invasions by increasing native species richness, and non-native species most closely related to the resident community should potentially be prioritized as those most likely to be highly invasive.  相似文献   

16.
Because of the economic and environmental importance of stabilizing fragile sand dune habitats, restoration of dunes has become a common practice. Restoration efforts in the Great Lakes and East Coast regions of North America often consist of planting monocultures of the dominant native grass species, Ammophila breviligulata. We evaluated 18 dune restoration projects in the Great Lakes region conducted over the past 25 years. We characterized attributes of diversity (plants and insects), vegetation structure (plant biomass and cover), and ecological processes (soil nutrients and mycorrhizal fungi abundance) in each restoration, and we compared these measures to geographically paired natural dune communities. Restoration sites were similar to reference sites in most measured variables. Differences between restorations and reference sites were mostly explained by differences in ages, with the younger sites supporting slightly lower plant diversity and mycorrhizal spore abundance than older sites. Plant community composition varied little between restored and reference sites, with only one native forb species, Artemisia campestris, occurring significantly more often in reference sites than restored sites. Although it remains unclear whether more diverse restoration plantings could accelerate convergence on the ecological conditions of reference dunes, in general, traditional restoration efforts involving monoculture plantings of A. breviligulata in Great Lakes sand dunes appear to achieve ecological conditions found in reference dunes.  相似文献   

17.
We set up two alternative hypotheses on how environmental variables could foster nestedness; one of “nested habitats” and another of “nested habitat quality”. The former hypothesis refers to situations where the nestedness of species depends on a nestedness of discrete habitats. The latter considers situations where all species in an assemblage increase in abundance along the same environmental gradient, but differ in specialisation or tolerance. We tested whether litter‐dwelling land snails (terrestrial gastropods) in boreal riparian forest exhibited a nested community structure, whether such a pattern was related to differences in environmental variables among sites, and which of the two hypotheses that best could account for the found pattern. We sampled litter from 100 m2 plots in 29 mature riparian forest sites along small streams in the boreal zone of Sweden. The number of snail species varied between 3 and 14 per site. Ranking the species‐by‐site matrix by PCA scores of the first ordination axis revealed a similarly significant nested pattern as when the matrix was sorted by number of species, showing that the species composition in this meta‐community can be properly described as nested. Several environmental variables, most notably pH index, were correlated with the first PCA axis. All but two species had positive eigenvectors in the PCA ordination and the abundance increased considerably along the gradient for most of the species implying that the hypothesis of “nested habitats” was rejected in favour of the “nested habitat quality” hypothesis. Analyses of nestedness have seldom been performed on equal sized plots, and our study shows the importance of understanding that variation in environmental variables among sites can result in nested communities. The conservation implications are different depending on which of our two hypotheses is supported; a conservation focus on species “hotspots” is more appropriate if the communities are nested because of “nested habitat quality”.  相似文献   

18.
According to the guidelines of the European Water Framework Directive, assessment of the ecological quality of streams and rivers should be based on type-specific reference conditions. Moreover to support biological indicators an hydromorphological analysis is also requested for each river type. The rationale for including an habitat assessment in biomonitoring study is that a biological community can be influenced by habitat quality just as water chemistry.In the present work benthic macroinvertebrates were analysed in a specific river type of Central Italy (small-sized streams, volcanic-siliceous), to identify taxa assemblages at the mesohabitat scale and to test how common measures of benthic community used in biomonitoring differ between riffles and pools in order to evaluate if differences may influence water quality classification.Macroinvertebrates were collected in 10 selected streams, covering the whole quality range present in the geographic area from ‘reference sites’ to human-impacted sites, along a pool–riffle sequence following a multihabitat sampling protocol.We compared assemblage of macroinvertebrates found in different mesohabitats using principal component analysis (PCA). Similar site grouping was obtained in riffle, pool and abiotic analysis.The measures of diversity and abundance were used as replicates in ANOVA analysis to test differences between pools and riffles within the groups of sites. There were no significant differences in terms of taxa richness and total abundance.When we compared the abundance of each taxon we found significant differences only in the group of reference sites with 18 taxa (about 25%) that showed a significant habitat preference.Our findings support that macroinvertebrates assemblages reflected primarily the environmental conditions and differences at mesohabitat scale are strongly correlated to hydromorphological condition and are maximized in reference sites. However such differences do not influence the ecological status assessment in this typology.  相似文献   

19.
A fundamental goal of restoration is the re-establishment of plant diversity representative of native vegetation. However, many prairie restorations or Conservation Reserve Program sites have been seeded with warm-season grasses, leading to grass-dominated, low-diversity restorations not representative of native grasslands. These dominant grasses are strongly mycotrophic, while many subordinate forb species appear to be less dependent on mycorrhizal symbiosis. Therefore, manipulating arbuscular mycorrhizal fungi (AMF) may be useful in promoting establishment and growth of forb species in grass-dominated prairie restorations. To assess the potential role of mycorrhizae in affecting the productivity and community composition of restored tallgrass prairie, we conducted a 4-year field experiment on an 8-year-old grassland restoration at the Konza Prairie in northeastern Kansas, USA. At the initiation of our study, seeds of 12 forb species varying in degree of mycorrhizal dependence were added to established grass-dominated plots. Replicate plots were treated bi-weekly with a soil drench of fungicide (Topsin-M®) over four growing seasons and compared to non-treated control plots to assess the role of AMF in affecting plant species composition, productivity, leaf tissue quality, and diversity in restored tallgrass prairie. Topsin applications successfully reduced mycorrhizal colonization of grass roots to approximately 60–80% relative to roots in control plots. Four years of mycorrhizal suppression reduced productivity of the dominant grasses and increased plant species richness and diversity. These results highlight the importance of mycorrhizae as mediators of plant productivity and community dynamics in restored tallgrass prairie and indicate that temporarily suppressing AMF decreases productivity of the dominant C4 grasses and allows for establishment of seeded forb species.  相似文献   

20.
We propose that patterns of plant functional group occurrences could be a reliable indicator of prairie vegetation quality. A method for assessing tallgrass prairie quality based on density and composition of plant functional groups was developed and tested by comparison with qualitative indices calculated from species data at 17 prairies in Illinois. Species sample data were recorded from quadrats while functional group data were recorded from segments of belt transects overlying the species sample transects. Prairies selected include remnants and restorations and represent a wide range of habitat quality including recognized natural areas, degraded remnants, and prairie plantings of varying age and success. For agglomerative clustering of prairie quality classes, a matrix of habitat indices and metrics was used based on species sample data from all sites. Three groups were identified in cluster analysis that were characteristic of high, medium, and low-quality prairies. Mean Functional Group Density (Mean FGD), an index developed based on the mean products of frequency and density among plant functional groups recorded from belt transects at each site, had the highest correlations to habitat quality indices among two other functional group indices tested. Mean FGD was highly correlated with the mean coefficient of conservatism and floristic quality index, indices calculated from species sample data that have been shown to be reliable indicators of habitat quality. In means comparison tests among prairie quality classes, Mean FGD differentiated high-quality from medium and low-quality prairies, but did not distinguish medium from low-quality sites (only two low-quality sites were identified), although rank order of Mean FGD was as predicted. There is a tradeoff in efficiency and precision between species-level and functional group sample data. Species-level data more precisely discriminate differences in low and medium-quality sites; however, functional group sampling is much more rapid requiring only 20-to-25% of the time required for collecting species-level data. Results from functional group sampling highlight differences in functional group composition among prairie quality classes. High-quality prairies are characterized by greater abundance of sedges and hemi-parasites while lower-quality prairies were affiliated more with non-native perennial forbs and annual/biennial species.  相似文献   

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