Increasing native, but not exotic, biodiversity increases aboveground productivity in ungrazed and intensely grazed grasslands

Species-rich native grasslands are frequently converted to species-poor exotic grasslands or pastures; however, the consequences of these changes for ecosystem functioning remain unclear. Cattle grazing (ungrazed or intensely grazed once), plant species origin (native or exotic), and species richness (4-species mixture or monoculture) treatments were fully crossed and randomly assigned to plots of grassland plants. We tested whether (1) native and exotic plots exhibited different responses to grazing for six ecosystem functions (i.e., aboveground productivity, light interception, fine root biomass, tracer nitrogen uptake, biomass consumption, and aboveground biomass recovery), and (2) biodiversity-ecosystem functioning relationships depended on grazing or species origin. We found that native and exotic species exhibited different responses to grazing for three of the ecosystem functions we considered. Intense grazing decreased fine root biomass by 53% in exotic plots, but had no effect on fine root biomass in native plots. The proportion of standing biomass consumed by cattle was 16% less in exotic than in native grazed plots. Aboveground biomass recovery was 30% less in native than in exotic plots. Intense grazing decreased aboveground productivity by 25%, light interception by 14%, and tracer nitrogen uptake by 54%, and these effects were similar in native and exotic plots. Increasing species richness from one to four species increased aboveground productivity by 42%, and light interception by 44%, in both ungrazed and intensely grazed native plots. In contrast, increasing species richness did not influence biomass production or resource uptake in ungrazed or intensely grazed exotic plots. These results suggest that converting native grasslands to exotic grasslands or pastures changes ecosystem structure and processes, and the relationship between biodiversity and ecosystem functioning.     

Exotic plant species in a C4-dominated grassland: invasibility, disturbance, and community structure

We used data from a 15-year experiment in a C₄-dominated grassland to address the effects of community structure (i.e., plant species richness, dominance) and disturbance on invasibility, as measured by abundance and richness of exotic species. Our specific objectives were to assess the temporal and spatial patterns of exotic plant species in a native grassland in Kansas (USA) and to determine the factors that control exotic species abundance and richness (i.e., invasibility). Exotic species (90% C₃ plants) comprised approximately 10% of the flora, and their turnover was relatively high (30%) over the 15-year period. We found that disturbances significantly affected the abundance and richness of exotic species. In particular, long-term annually burned watersheds had lower cover of exotic species than unburned watersheds, and fire reduced exotic species richness by 80–90%. Exotic and native species richness were positively correlated across sites subjected to different fire (r = 0.72) and grazing (r = 0.67) treatments, and the number of exotic species was lowest on sites with the highest productivity of C₄ grasses (i.e., high dominance). These results provide strong evidence for the role of community structure, as affected by disturbance, in determining invasibility of this grassland. Moreover, a significant positive relationship between exotic and native species richness was observed within a disturbance regime (annually burned sites, r = 0.51; unburned sites, r = 0.59). Thus, invasibility of this C₄-dominated grassland can also be directly related to community structure independent of disturbance. Received: 9 February 1999 / Accepted: 12 May 1999     

Competing drivers lead to non-linear native–exotic relationships in endangered temperate grassy woodlands

Relationships between the diversity and abundance of native versus exotic species underpin management of disturbance regimes for conservation. Theory predicts negative, positive or neutral relationships depending on respective drivers, with greatest potential benefit when natives and exotics show opposing responses to management. We examined drivers of exotic plant cover and relationships with native plant richness using 12-year burning, mowing and grazing experiments in two representative temperate grassy eucalypt woodlands with contrasting histories of frequent versus infrequent disturbance. We hypothesized that disturbance and high resources favour exotics, and assessed whether natives and exotics covary positively due to common external drivers or negatively due to contrasting external drivers and/or competition. Positive relationships with rainfall and disturbance explained >80 % of the variation in exotic cover at both sites, supporting our first hypothesis. Native–exotic relationships were non-linear, with native richness first increasing rapidly with increasing exotic cover, then levelling and beginning to decrease. Common external drivers, particularly inter-annual rainfall, explained initial positive relationships, highlighting a prevalence of positive relationships at long temporal (as well as large spatial) scales. At the historically frequently-burnt site, a concomitant increase in native richness and exotic cover after fire contributed to the positive relationship, indicating a management trade-off. At the long-unburnt site, exotics increased but natives decreased with fire, suggesting dual benefits of low fire frequency. We conclude that relationships between exotic cover and native richness emerge from interactions among external drivers and competitive responses, with responses to external drivers dominating at low resources and negative interactions gaining importance as resources increase.     

What determines disturbance‐productivity‐diversity relationships? The effect of scale,species and environment on richness patterns in an Australian woodland

Much of the observed variation in relationships between diversity and disturbance or productivity may be attributed to scale, species characteristics, or environment. We used exclusion fences to create gradients of grazing (by native and introduced herbivores), cover, and standing crop in temperate Eucalypt woodlands. We investigated patterns of native, exotic, and total plant species richness at two scales (1 m² and 625 m²). Richness patterns were similar at both scales, though species richness at 1m² was more strongly affected by our grazing treatments. Season and rainfall explained more variation in richness than did surrogate measures of productivity or disturbance by herbivores. The richness-herbivory relationship depended strongly on rainfall, season, and species origin, and altering these factors produced the entire range of observed diversity-disturbance relationships. Richness-biomass and richness-cover relationships were consistently hump-shaped, and related to species origin with native richness negatively related and exotic richness positively related. The ability of weedy annuals to pre-empt space after death may have contributed to the observed unimodal responses.     

Can root trait diversity explain complementarity effects in a grassland biodiversity experiment?

Aims The positive relationship between plant biodiversity and community productivity is well established. However, our knowledge about the mechanisms underlying these positive biodiversity effects is still limited. One of the main hypotheses is that complementarity in resource uptake is responsible for the positive biodiversity effects: plant species differ in resource uptake strategy, which results in a more complete exploitation of the available resources in space and time when plant species are growing together. Recent studies suggest that functional diversity of the community, i.e. the diversity in functional characteristics ('traits') among species, rather than species richness per se, is important for positive biodiversity effects. However, experimental evidence for specific trait combinations underlying resource complementarity is scarce. As the root system is responsible for the uptake of nutrients and water, we hypothesize that diversity in root traits may underlie complementary resource use and contribute to the biodiversity effects.Methods In a common garden experiment, 16 grassland species were grown in monoculture, 4-species mixtures differing in root trait diversity and 16-species mixtures. The 4-species mixtures were designed to cover a gradient in average rooting depth. Above-ground biomass was cut after one growing season and used as a proxy for plant productivity to calculate biodiversity effects.Important findings Overall, plant mixtures showed a significant increase in biomass and complementarity effects, but this varied greatly between communities. However, diversity in root traits (measured in a separate greenhouse experiment and based on literature) could not explain this variation in complementarity effects. Instead, complementarity effects were strongly affected by the presence and competitive interactions of two particular species. The large variation in complementarity effects and significant effect of two species emphasizes the importance of community composition for positive biodiversity effects. Future research should focus on identifying the traits associated with the key role of particular species for complementarity effects. This may increase our understanding of the links between functional trait composition and biodiversity effects as well as the relative importance of resource complementarity and other underlying mechanisms for the positive biodiversity effects.     

Long-term grazing impacts on vegetation diversity,composition, and exotic species presence across an aridity gradient in northern temperate grasslands

Little is known about the specific role of exotic species on measures of grassland plant diversity, including how this may vary with climatic conditions or large mammal herbivory. This study examined vegetation responses to long-term livestock grazing, including plant richness and diversity, as well as the contribution of exotic species to these metrics, across a network of 107 northern temperate grasslands in Alberta, Canada, spanning a broad aridity gradient. Exposure to grazing modestly increased plant richness, but did not alter Shannon’s diversity, Simpson’s diversity, or evenness, suggesting stability in floral diversity relative to grazing. However, grazing did increase grass cover while reducing shrub cover, the latter of which was only apparent in mesic grasslands. Unlike total plant diversity, exotic species richness and cover, together with exotic plant contributions to diversity, varied jointly with grazing and aridity. While long-term grazing increased exotic species, this response was most apparent in wetter areas, and non-grazed grasslands remained more resistant to the presence of exotics. Several exotic species were positive indicators of grazing in wetter grasslands, and coincided with lower native species cover, indicating grazing may be facilitating a shift from native to exotic vegetation under these conditions. Overall, our results indicate that while long-term grazing has altered the composition and cover of certain functional groups, including favoring exotics and minimizing woody vegetation in mesic areas, overall changes to plant diversity were limited. Additionally, these findings suggest that semi-arid northern temperate grasslands remain relatively resistant to grazing effects, including their susceptibility to exotic plant encroachment. These results improve our understanding of how ongoing grazing exposure may impact grassland diversity, including efforts to conserve native vegetation, as well as the important role of climate in altering fundamental grassland responses to grazing.     

Relationships between arthropod richness,evenness, and diversity are altered by complementarity among plant genotypes

Biodiversity is quantified via richness (e.g., the number of species), evenness (the relative abundance distribution of those species), or proportional diversity (a combination of richness and evenness, such as the Shannon index, H′). While empirical studies show no consistent relationship between these aspects of biodiversity within communities, the mechanisms leading to inconsistent relationships have received little attention. Here, using common evening primrose (Oenothera biennis) and its associated arthropod community, we show that relationships between arthropod richness, evenness, and proportional diversity are altered by plant genotypic richness. Arthropod richness increased with O. biennis genotypic richness due to an abundance-driven accumulation of species in response to greater plant biomass. Arthropod evenness and proportional diversity decreased with plant genotypic richness due to a nonadditive increase in abundance of a dominant arthropod, the generalist florivore/omnivore Plagiognathas politus (Miridae). The greater quantity of flowers and buds produced in polycultures—which resulted from positive complementarity among O. biennis genotypes—increased the abundance of this dominant insect. Using choice bioassays, we show that floral quality did not change in plant genotypic mixtures. These results elucidate mechanisms for how plant genotypic richness can modify relationships between arthropod richness, evenness, and proportional diversity. More broadly, our results suggest that trophic interactions may be a previously underappreciated factor controlling relationships between these different aspects of biodiversity.     

Recovery of Native Plant Community Characteristics on a Chronosequence of Restored Prairies Seeded into Pastures in West‐Central Iowa

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.     

Biodiversity, phenology and temporal niche differences between native- and novel exotic-dominated grasslands

Many exotic species have been introduced or have escaped into grasslands where they form ‘novel ecosystems’ of species with no evolutionary history of interaction. Novel ecosystems are good model systems for understanding how diversity maintenance mechanisms might differ between species with a history of interaction (natives) and species without a history (exotics) in cases where exotics originated from several continents. We tested for lower species diversity and richness in exotic grasslands and found a negative correlation between species diversity measures and proportion of exotic species across 15 grasslands in an observational study in Texas. We then planted 9-species mixtures of all native or all exotics under ambient or elevated summer precipitation to compare dynamics of diversity and to test if exotic species respond more strongly to altered resource availability. Species diversity was lower in communities of exotic than native species by the second year. Reduced diversity in exotic communities resulted from lower complementarity and higher temporal niche overlap among species and occurred in both ambient and irrigated plots. In general, summer irrigation had additive positive effects and did not interact with native–exotic status. Exotic species and communities had much earlier green-up during spring than natives, and altered inter-correlations among phenology variables. There were no differences in flowering dates. Taken together, our results suggest that rapid and synchronous growth may increase niche overlap among exotic species and reduce local diversity in exotic-dominated grassland communities. Earlier green-up by exotics may complicate attempts to ascertain relationships between phenology and climate. An increase in exotic species may cause earlier green-up regardless of any climate change effects and our results suggest that phenology networks should take a species-based rather than an ecosystem approach to evaluate green-up if the abundance of exotics increases within the time-frame in question. These differences between native and exotic species and communities should be considered in future management and restoration projects.     

Relationship between land uses and diversity of dung beetles (Coleoptera: Scarabaeinae) in the southern Atlantic forest of Argentina: which are the key factors?

The loss of natural habitats is one of the main drivers of biodiversity decline. Anthropogenic land uses preserving biotic and abiotic conditions of the native ecosystem are more suitable to preserve the native biodiversity. In this study, we explored changes in species richness and composition in different land uses of the southern Atlantic forest, considering three independent factors: (1) canopy (presence–absence), (2) type of vegetation (native–exotic) and (3) livestock (presence–absence). We expected a gradient of response in the richness and composition of the native forest dung beetle community, from land uses preserving canopy and native vegetation to open land uses with exotic vegetation. Dung beetles were sampled in protected native forests and four land uses, using two potential food resources: human dung and carrion. The species richness and composition of each habitat, as well as differences in composition and the influence of factors over diversity, were then analyzed. As expected, our results showed that land uses preserving canopy and native vegetation maintain the dung beetle diversity of the native forest. Moreover, while the three factors analyzed influenced dung beetle diversity, canopy cover was the main driver of dung beetle diversity loss. The main conclusion of this study is that the conservation of canopy (either native or exotic) is determinant to preserve highly diverse dung beetle communities and subsequently, the ecological functions performed by this taxon. However, the ecophysiological mechanism behind the response of dung beetles to habitat disturbance is poorly understood.     

Biophysical and human influences on plant species richness in grasslands: Comparing variegated landscapes in subtropical and temperate regions

A survey of grassy woodlands in the Queensland subtropics was conducted, recording herbaceous species richness at 212 sites on three properties (2756 ha). A range of habitats typical of cattle grazing enterprises was sampled and site variables included lithology, slope position, tree density, soil disturbance, soil enrichment and grazing. Results were compared with a previously published survey of temperate grasslands. Lithology, slope position and tree density had relatively minor effects on plant species richness, although in both surveys there was some evidence of lower species richness on the more fertile substrates. Soil disturbance and soil enrichment significantly reduced the richness of native species in both surveys, while exotic species were insensitive (subtropics) or increased (temperate) with disturbance. Rare native species were highly sensitive to disturbances, including grazing, in the temperate study. Although some trends were similar for rare species in the subtropics, the results were not significant and there were complex interactions between grazing, lithology and slope position. Grazing did not have a negative effect on native species richness, except in the closely grazed patches within pastures, and then only on the most intensively developed property. At the scale recorded (30 m²), the native pastures, roadsides and stock routes sampled in the subtropics appear to be among the most species‐rich grasslands ever reported, both nationally and globally. Native species richness was approximately 50% higher than the temperate survey figures across all the comparable habitats. While there are no clear reasons for this result, potential explanations are proposed.     

Rapid short-tussock grassland decline with and without grazing, Marlborough, New Zealand

Species abundance, species richness, and ground cover were measured over 10 years on nine paired grazed and exclosure plots in short-tussock grassland in the early stages of invasion by Hieracium species. With and without grazing, H. pilosella and H. caespitosum increased markedly and H. lepidulum increased locally. In contrast, 50% of all other common species and species groups, and total, native, and exotic species richness declined significantly. Exclusion increased or had no effect on rates of increase in Hieracium species and rates of decline in short tussocks, and did not reduce rates of decline in other species. Exclusion had no effect on decline in native species richness, but mainly accelerated declines in total and exotic richness. Declines in 13 key vegetation variables were significantly predicted by increase in Hieracium abundance, suggesting competitive exclusion. With or without grazing, Hieracium species will become more dominant and other species will continue to decline. The effects of large herbivores on plant species diversity can often be predicted from site productivity. Our results indicate the need also to account for species origin, spatial scale, time, and exotic invasion.     

Temporal changes in native-exotic richness correlations during early post-fire succession

The relationship between native and exotic richness has mostly been studied with respect to space (i.e., positive at larger scales, but negative or more variable at smaller scales) and its temporal patterns have rarely been investigated. Although some studies have monitored the temporal trends of both native and exotic richness, how these two groups of species might be related to each other and how their relative proportions vary through time in a local community remains unclear. Re-analysis of early post-fire successional data for a California chaparral community shows that, in the same communities and at small spatial scales, the native-exotic correlations varied through time. Both exotic richness and exotic fraction (i.e., the proportion of exotic species in the flora) quickly increased and then gradually declined, during the initial stages of succession following fire disturbance. This result sheds new light on habitat invasibility and has implications for timing the implementation of effective management actions to prevent and/or mitigate species invasions.     

Tropical paradox: a multi-scale analysis of the invasion paradox within Miami Rock Ridge tropical hardwood hammocks

The invasion paradox describes the scale dependence of native-exotic richness relationships (NERRs), where NERRs are negative at neighborhood scales and positive at landscape scales. However, a lack of tropical surveys and past failures to isolate potential confounding variables contribute to significant gaps in our understanding of the processes producing these patterns. We surveyed the vascular flora of 13 tropical hardwood hammocks for community characteristics (e.g., native and exotic species richness, vegetative cover) with a hierarchical sampling design. Using model selection, we determined which variables best predicted patterns of exotic species richness at each spatial scale of consideration. We found that native and exotic species richness were positively correlated at neighborhood scales, but negatively correlated at landscape scales. The latter result stands in stark opposition to the patterns published in the literature thus far. We found that natural disturbance history (as approximated by vegetative cover) was positively correlated with exotic species richness at intermediate and landscape scales only. Overall, hammock identity was the most important factor driving exotic species richness patterns at all spatial scales. Hammocks with highly-disturbed hydrologies, brought about by water management, had fewer native species and more exotic species than hammocks with more natural hydrological conditions. Our results are among the first from examination of subtropical communities, and may support the hypothesis that tropical and subtropical communities are subject to more intense biotic interactions. However, given our unique sampling design, our results do not reject the hypothesis that environmental heterogeneity drives the relationship between native and exotic species richness patterns.     

Landscape-scale patterns of biological invasions in shoreline plant communities

Little is known about the patterns and dynamics of exotic species invasions at landscape to regional spatial scales. We quantified the presence (identity, abundance, and richness) and characteristics of native and exotic species in estuarine strandline plant communities at 24 sites in Narragansett Bay, Rhode Island, USA. Our results do not support several fundamental predictions of invasion biology. Established exotics (79 of 147 recorded plant species) were nearly indistinguishable from the native plant species (i.e. in terms of growth form, taxonomic grouping, and patterns of spatial distribution and abundance) and essentially represent a random sub-set of the current regional species pool. The cover and richness of exotic species varied substantially among quadrats and sites but were not strongly related to any site-level physical characteristics thought to affect invasibility (i.e. the physical disturbance regime, legal status, neighboring habitat type, and substrate characteristics). Native and exotic cover or richness were not negatively related within most sites. Across sites, native and exotic richness were positively correlated and exotic cover was unrelated to native richness. The colonization and spread of exotics does not appear to have been substantially reduced at sites with high native diversity. Furthermore, despite the fact that the Rhode Island strandline system is one of the most highly-invaded natural plant communities described to date, exotic species, both individually and as a group, currently appear to pose little threat to native plant diversity. Our findings are concordant with most recent, large-scale investigations that do not support the theoretical foundation of invasion biology and generally contradict small-scale experimental work.     

Native weeds and exotic plants: relationships to disturbance in mixed-grass prairie

Disturbance frequently is implicated in the spread of invasive exotic plants. Disturbances may be broadly categorized as endogenous (e.g., digging by fossorial animals) or exogenous (e.g., construction and maintenance of roads and trails), just as weedy species may be native or exotic in origin. The objective of this study was to characterize and compare exotic and native weedy plant occurrence in and near three classes of disturbance – digging by prairie dogs (an endogenous disturbance to which native plants have had the opportunity to adapt), paved or gravel roads (an exogenous disturbance without natural precedent), and constructed trails (an exogenous disturbance with a natural precedent in trails created by movement of large mammals) – in three geographically separate national park units. I used plant survey data from the North and South Units of Theodore Roosevelt National Park and Wind Cave National Park in the northern mixed-grass prairie of western North and South Dakota, USA, to characterize the distribution of weedy native and exotic plants with respect to the three disturbance classes as well as areas adjacent to them. There were differences both in the susceptibility of the disturbance classes to invasion and in the distributions of native weeds and exotic species among the disturbance classes. Both exotic and native weedy species richness were greatest in prairie dog towns and community composition there differed most from undisturbed areas. Exotic species were more likely to thrive near roadways, where native weedy species were infrequently encountered. Exotic species were more likely to have spread beyond the disturbed areas into native prairie than were weedy native species. The response of individual exotic plant species to the three types of disturbance was less consistent than that of native weedy species across the three park units. This revised version was published online in June 2006 with corrections to the Cover Date.     

城市化对植物多样性影响的研究进展

本文综述了城市化对植物多样性影响的研究进展。随着全球特别是发展中国家城市化水平的提高, 城市化对生物多样性的影响逐渐引起了人们的重视。城市化造成了本土植物物种的丢失和外来物种的增加。在空间分布上, 城市化还常使城区本土植物多样性沿着远郊农区－城郊－城区梯度性下降; 但是由于引进大量外来物种, 总体植物多样性反而升高, 沿着远郊农区－城郊－城区梯度性升高。城市化对植物种类组成也有很大影响,优势种在远郊农区、城郊和城区呈现不同。城市化对植物多样性影响的机制主要有人为引入外来物种、小生境改变以及景观格局的变化三个方面。以下四个研究方向将越来越重要: (1) 不同区域、不同方法、不同学科的系统整合研究;(2) 城市化扩张与植物多样性变化过程的定位监测研究; (3) 本土植物多样性丢失以及性状改变的内在机制研究, 特别是外来物种与本地物种的相互作用过程和机制的研究; (4) 城市植物多样性保护研究。     

Climate warming moderates the impacts of introduced sportfish on multiple dimensions of prey biodiversity

Human‐assisted introductions of exotic species are a leading cause of anthropogenic change in biodiversity; however, context dependencies and interactions with co‐occurring stressors impede our ability to predict their ecological impacts. The legacy of historical sportfish stocking in mountainous regions of western North America creates a unique, natural quasiexperiment to investigate factors moderating invasion impacts on native communities across broad geographic and environmental gradients. Here we synthesize fish stocking records and zooplankton relative abundance for 685 mountain lakes and ponds in the Cascade and Canadian Rocky Mountain Ranges, to reveal the effects of predatory sportfish introduction on multiple taxonomic, functional and phylogenetic dimensions of prey biodiversity. We demonstrate an innovative analytical approach, combining exploratory random forest machine learning with confirmatory multigroup analysis using multivariate partial least‐squares structural equation models, to generate and test hypotheses concerning environmental moderation of stocking impacts. We discovered distinct effects of stocking across different dimensions of diversity, including negligible (nonsignificant) impacts on local taxonomic richness (i.e. alpha diversity) and trophic structure, in contrast to significant declines in compositional uniqueness (i.e. beta diversity) and body size. Furthermore, we found that stocking impacts were moderated by cross‐scale interactions with climate and climate‐related land‐cover variables (e.g. factors linked to treeline position and glaciers). Interactions with physical morphometric and lithological factors were generally of lesser importance, though catchment slope and habitat size constraints were relevant in certain dimensions. Finally, applying space‐for‐time substitution, a strong antagonistic (i.e. dampening) interaction between sportfish predation and warmer temperatures suggests redundancy of their size‐selective effects, meaning that warming will lessen the consequences of introductions in the future and stocked lakes may be less impacted by subsequent warming. While both stressors drive biotic homogenization, our results have important implications for fisheries managers weighing the costs/benefits of stocking—or removing established non‐native populations—under a rapidly changing climate.     

Factors Controlling Compositional Changes in a Northern Andean Páramo (La Rusia,Colombia)

In this study, we aimed to assess the processes controlling compositional change in a Northern Andean páramo highly affected by human‐induced disturbances over the last few decades (La Rusia, Colombia). Along the 3000–3800 m asl altitudinal range, we randomly sampled fifty 10 × 10 m plots. Therein, we measured altitude and variables related to soil conditions (i.e., moisture, nutrient contents, bulk density, and texture), occurrence of human‐induced disturbances (i.e., fire, vegetation clearing, potato cultivation, and cattle grazing), and land‐use history. We also recorded richness and abundance of plant species, identifying them as exotic or native. We differentiated four groups of plots according to their species composition. The groups had significant differences in altitude, soil conditions, land‐use history, and particularly, in richness of exotic species and exotic/native cover ratio. They could be ascribed to shrub‐ and grass‐páramo vegetation types based on their relative dominance of woody and herbaceous species; however, these groups were not arranged according to the hypothetical composition of altitudinal belts, but rather formed a mosaic of patches. This mosaic was determined not only by altitude but also by soil conditions and disturbance history of sites. Our results corroborate recent findings which highlight shrub‐ and grass‐páramo vegetation types as patches of contrasting species composition and structure that depend on local environmental variables, as well as human‐induced disturbances as a major determinant of compositional discontinuities in these ‘high mountain’ tropical ecosystems.     

Native species richness buffers invader impact in undisturbed but not disturbed grassland assemblages

Many systems are prone to both exotic plant invasion and frequent natural disturbances. Native species richness can buffer the effects of invasion or disturbance when imposed in isolation, but it is largely unknown whether richness provides substantial resistance against invader impact in the face of disturbance. We experimentally examined how disturbance (drought/burning) influenced the impact of three exotic invaders (Centaurea stoebe, Linaria dalmatica, or Potentilla recta) on native abundance across a gradient of species richness, using previously constructed grassland assemblages. We found that invaders had higher cover in experimentally disturbed plots than in undisturbed plots across all levels of native species richness. Although exotic species varied in cover, all three invaders had significant impacts on native cover in disturbed plots. Regardless of disturbance, however, invader cover diminished with increasing richness. Invader impacts on native cover also diminished at higher richness levels, but only in undisturbed plots. In disturbed plots, invaders strongly impacted native cover across all richness levels, as disturbance favoured invaders over native species. By examining these ecological processes concurrently, we found that disturbance exacerbated invader impacts on native abundance. Although diversity provided a buffering effect against invader impact without disturbance, the combination of invasion and disturbance markedly depressed native abundance, even in high richness assemblages.