Cover Of Exotic Species Research Articles

Ecological filtering of exotic plants in an Australian sub‐alpine environment

We investigated some of the factors influencing exotic invasion of native sub‐alpine plant communities at a site in southeast Australia. Structure, floristic composition and invasibility of the plant communities and attributes of the invasive species were studied. To determine the plant characteristics correlated with invasiveness, we distinguished between roadside invaders, native community invaders and non‐invasive exotic species, and compared these groups across a range of traits including functional group, taxonomic affinity, life history, mating system and morphology. Poa grasslands and Eucalyptus‐Poa woodlands contained the largest number of exotic species, although all communities studied appeared resilient to invasion by most species. Most community invaders were broad‐leaved herbs while roadside invaders contained both herbs and a range of grass species. Over the entire study area the richness and cover of native and exotic herbaceous species were positively related, but exotic herbs were more negatively related to cover of specific functional groups (e.g. trees) than native herbs. Compared with the overall pool of exotic species, those capable of invading native plant communities were disproportionately polycarpic, Asteracean and cross‐pollinating. Our data support the hypothesis that strong ecological filtering of exotic species generates an exotic assemblage containing few dominant species and which functionally converges on the native assemblage. These findings contrast with those observed in the majority of invaded natural systems. We conclude that the invasion of closed sub‐alpine communities must be viewed in terms of the unique attributes of the invading species, the structure and composition of the invaded communities and the strong extrinsic physical and climatic factors typical of the sub‐alpine environment.

Ecological filtering of exotic plants in an Australian sub-alpine environment

We investigated some of the factors influencing exotic invasion of native sub-alpine plant communities at a site in southeast Australia. Structure, floristic composition and invasibility of the plant communities and attributes of the invasive species were studied. To determine the plant characteristics correlated with invasiveness, we distinguished between roadside invaders, native community invaders and non-invasive exotic species, and compared these groups across a range of traits including functional group, taxonomic affinity, life history, mating system and morphology. Poa grasslands and Eucalyptus-Poa woodlands contained the largest number of exotic species, although all communities studied appeared resilient to invasion by most species. Most community invaders were broadleaved herbs while roadside invaders contained both herbs and a range of grass species. Over the entire study area the richness and cover of native and exotic herbaceous species were positively related, but exotic herbs were more negatively related to cover of specific functional groups (e.g. trees) than native herbs. Compared with the overall pool of exotic species, those capable of invading native plant communities were disproportionately polycarpic, Asteracean and cross-pollinating. Our data support the hypothesis that strong ecological filtering of exotic species generates an exotic assemblage containing few dominant species and which functionally converges on the native assemblage. These findings contrast with those observed in the majority of invaded natural systems. We conclude that the invasion of closed sub-alpine communities must be viewed in terms of the unique attributes of the invading species, the structure and composition of the invaded communities and the strong extrinsic physical and climatic factors typical of the sub-alpine environment.

Invasion success of exotic plants in natural ecosystems: the role of disturbance, plant attributes and freedom from herbivores

Invasion of natural ecosystems by exotic species is a major threat to biodiversity globally. We assessed two alternative (but not exclusive) hypotheses to explain the success of exotic species in urban bushland on low fertility sandstone-derived soils in Sydney, Australia. These were that success of exotic species is promoted by: (1) plant attributes in particular disturbance types; and (2) freedom from herbivores. We tested these at sites subject to different types of disturbance: nutrient and water enrichment (below stormwater outlets), nutrient enrichment (riparian zones of creeks with an urban catchment) and physical disturbance (tracks), and control sites. At each site we estimated percentage cover of all species and surveyed leaves for damage by herbivores. Species were classified as native, non-invasive exotic or invasive exotic. We found that sites without any disturbance did not support exotic plants. Physically disturbed sites on low fertility soils supported only one exotic species, suggesting that nutrient enrichment is a critical prerequisite for exotic species invasion on low fertility soils. Exotic species cover was highest and native species richness most reduced in areas of highest nutrient enrichment. Both invasive exotic and non-invasive exotic species had significantly lower levels of leaf herbivory than native species, implying that release from pests alone cannot account for the success of invasive species. Specific leaf area of invasive exotic species was consistently higher than specific leaf area of non-invasive exotic and native species, regardless of disturbance type. In physically disturbed sites of higher soil fertility, exotic species were small herbs and grasses of long flowering duration and with small unassisted or wind-dispersed seeds. In sites subject to nutrient-enrichment, exotic species were more likely to be climbers, able to propagate vegetatively, and with seeds dispersed by vertebrates. Thus different plant attributes contribute to exotic species success under different disturbance types. The clearest consistent difference we found between invasive exotic and non-invasive exotic species was in specific leaf area, suggesting that large specific leaf area facilitates invasiveness.

Understory plant species composition 30–50 years after clearcutting in southeastern Wyoming coniferous forests

To evaluate the long-term effects of clearcutting and slash treatment on understory plant species composition in Rocky Mountain coniferous forests, we collected comparable data from 30- to 50-year-old post-harvest stands and adjacent mature (>100-year-old) stands that originated after wildfire. In general, plant species richness was higher in the post-harvest stands than in adjacent mature stands, due in large part to the presence of species that benefit from disturbance. Species composition in post-harvest and mature stands was more similar in montane forests (2710–2805 m elevation) than in subalpine forests (2800–3164 m elevation), suggesting greater resilience of montane understories. Composition in post-harvest and mature stands was least similar when slash was piled and burned, and most similar when slash was lopped and scattered. Eighty-seven percent of the post-harvest stands had at least one exotic species, but total cover of exotic plant species in all post-harvest stands was ≤1%. Exotic species were largely absent from mature forests. Our results suggest that understory species of low-elevation montane forests recover more quickly from disturbance than do those of subalpine forests, with slash treatment and species life history traits strongly influencing this recovery.

A Protocol for Integrated Management, Monitoring, and Enhancement of Degraded Themeda triandra Grasslands based on Plantings of Indicator Species

Abstract Lowland temperate grasslands dominated by Themeda triandra (kangaroo grass) are an endangered ecosystem in southeastern Australia. Grass biomass must be removed frequently to maintain plant diversity, but few studies of the impacts of different biomass removal techniques have been undertaken, and no rapid monitoring schemes have been developed. Low species densities in many reserves (due to past stock grazing) make it difficult to assess the effects of management regimes on plant diversity. Management impacts could be assessed by planting indicator species in replicated enhancement plots and subjecting these plots to adaptive management trials. A protocol for selecting potential indicator species is described, based on a regional quadrat database, using clearly defined criteria. Potential indicator species need to be conspicuous, easy to identify and abundant in high quality diverse grassland remnants, to have relatively broad ecological tolerances, to occur in sites that are relatively species rich and have a comparatively low cover of dominant exotic species, to commonly persist at low densities in long‐grazed reserves, to be responsive to changes in management, and to have been studied ecologically. Only three species from western Victorian grasslands satisfied these criteria: Calocephalus citreus (lemon beauty‐heads), Chrysocephalum apiculatum (common everlasting), and Leptorhynchos squamatus (scaly buttons). All are widespread, herbaceous, hemicryptophytic daisies. Despite a number of caveats, the scheme has the potential to provide a more clearly focused framework for grassland ecosystem management than currently exists.

Exotic plant species of the St Lawrence River wetlands: a spatial and historical analysis

AbstractAim To evaluate the importance (number of species, plant cover) of the exotic flora in seven well‐defined sectors of one of the most important transportation waterways in North America. To determine the impact of exotic species on wetland plant diversity and reconstruct the spread of some invasive species.Location St Lawrence River, southern Québec.Methods The exotic flora (vascular plants) of wetlands bordering the St Lawrence River was studied using 713 sampling stations (25 m2) along a 560‐km long corridor.Results Exotic species represent 13.7% of the vascular flora of the St Lawrence wetlands. The relative plant cover occupied by exotic species is high in some of the fluvial sectors (42–44%), but low (6–10%) in the estuarine sectors. Wetlands (marshes) surrounding islands were particularly susceptible to invasion by exotic plants. Historical, abiotic and landscape factors may explain the differences observed between sites. Purple loosestrife (Lythrum salicaria L.) is the most common exotic species of the St Lawrence wetlands, but other species, namely flowering‐rush (Butomus umbellatus L.) and reed canary grass (Phalaris arundinacea L.) are much more invasive. There is no linear relationship between the exotic species cover and the diversity of wetland plants; low diversity sites can be dominated by either exotic or native plant species. In the other sites, exotic species generally have little impact on plant communities and can contribute to increase diversity. Common reed (Phragmites australis (Cav.) Trin. ex Steudel) and reed canary grass, both considered as exotic species in this study, clearly have a stronger impact on plant diversity than flowering‐rush and purple loosestrife.Main conclusions This study shows that the global impact of an invader cannot be adequately evaluated with only a few highly invaded sites. While nationwide strategies have been developed to control exotic species, large surveys are essential to adapt them to regional particularities.

Roads as Conduits for Exotic Plant Invasions in a Semiarid Landscape

Abstract: Roads are believed to be a major contributing factor to the ongoing spread of exotic plants. We examined the effect of road improvement and environmental variables on exotic and native plant diversity in roadside verges and adjacent semiarid grassland, shrubland, and woodland communities of southern Utah ( U.S.A. ). We measured the cover of exotic and native species in roadside verges and both the richness and cover of exotic and native species in adjacent interior communities ( 50 m beyond the edge of the road cut ) along 42 roads stratified by level of road improvement ( paved, improved surface, graded, and four‐wheel‐drive track ). In roadside verges along paved roads, the cover of Bromus tectorum was three times as great ( 27% ) as in verges along four‐wheel‐drive tracks ( 9% ). The cover of five common exotic forb species tended to be lower in verges along four‐wheel‐drive tracks than in verges along more improved roads. The richness and cover of exotic species were both more than 50% greater, and the richness of native species was 30% lower, at interior sites adjacent to paved roads than at those adjacent to four‐wheel‐drive tracks. In addition, environmental variables relating to dominant vegetation, disturbance, and topography were significantly correlated with exotic and native species richness and cover. Improved roads can act as conduits for the invasion of adjacent ecosystems by converting natural habitats to those highly vulnerable to invasion. However, variation in dominant vegetation, soil moisture, nutrient levels, soil depth, disturbance, and topography may render interior communities differentially susceptible to invasions originating from roadside verges. Plant communities that are both physically invasible ( e.g., characterized by deep or fertile soils ) and disturbed appear most vulnerable. Decision‐makers considering whether to build, improve, and maintain roads should take into account the potential spread of exotic plants.

Is fencing enough? The short‐term effects of stock exclusion in remnant grassy woodlands in southern NSW

Summary Fencing remnant native vegetation has become a widespread activity for arresting declines in biodiversity in agricultural landscapes. However, few data are available on the effectiveness of this approach. The present study investigated the short‐term effects of fencing to exclude livestock on dominant tree and shrub recruitment, plant species cover, litter and soil characteristics in remnant grassy woodlands in southern NSW. Vegetation and soil surveys were undertaken at 47 sites fenced by Greening Australia (NSW) for 2–4 years. Fenced and unfenced areas at each site were compared using split‐plot sampling. Woodlands sampled were dominated by Yellow Box/Blakely’s Red Gum (Eucalyptus melliodora/Eucalyptus blakelyi), Grey Box (Eucalyptus microcarpa) or White Cypress‐pine (Callitris glaucophylla). Significantly higher numbers of tree recruits were found in the fenced sites, with tree recruitment found in 59% of fenced sites compared with 13% of unfenced sites. Fenced sites also had significantly greater cover of native perennial grasses, less cover of exotic annual species and less soil surface compaction. However, outcomes varied among woodland ecosystems and individual sites. Where tree recruitment occurred, there was significantly more tree recruitment where there was greater perennial grass cover and less regeneration where exotic annual grass cover or overstorey crown cover was dense. Few shrubs recruited in fenced or unfenced areas, reflecting the lack of mature shrubs in most sites. Fencing is an important first step for conserving threatened grassy woodlands, but more active management may be needed to enhance woodland recovery, particularly in sites where few or no recruits were found.Key words bush regeneration, fencing, grazing exclusion, rehabilitation, woodland restoration.

Exotic plant invasions over 40 years of old field successions: community patterns and associations

While exotic plant species often come to dominate disturbed communities, long‐term patterns of invasion are poorly known. Here we present data from 40 yr of continuous vegetation sampling, documenting the temporal distribution of exotic plant species in old field succession. The relative cover of exotic species decreased with time since abandonment, with significant declines occurring ≥20 yr post‐abandonment. The number of exotic species per plot also declined with time since abandonment while field‐scale richness of exotics did not change. This suggests displacement occurring at small spatial scales. Life history types changed from short‐lived herbaceous species to long‐lived woody species for both native and exotic plant species. However, shrubs and lianas dominated woody cover of exotic plants while trees dominated native woody cover. The species richness of exotic and native species was positively correlated at most times. In abandoned hay fields, however, the proportion of exotic plant cover per plot was inversely related to total species richness. This relationship suggests that it is not the presence, but the abundance of exotic species that may cause a reduction in community diversity. While the development of closed‐canopy forest appears to limit most introduced plant species, several shade‐adapted exotic species are increasing within the fields. These invasions may cause a reversal of the patterns seen in the first 40 yr of succession and may result in further impacts on community structure.

Exotic and native vegetation establishment following channelization of a western Iberian river

Channelization is often a major cause of human impacts on river systems. It affects both hydrogeomorphic features and habitat characteristics and potentially impacts riverine flora and fauna. Human-disturbed fluvial ecosystems also appear to be particularly vulnerable to exotic plant establishment. Following a 12-year recovery period, the distribution, composition and cover of both exotic and native plant species were studied along a Portuguese lowland river segment, which had been subjected to resectioning, straightening and two-stage bank reinforcement, and were compared with those of a nearby, less impacted segment. The species distribution was also related to environmental data. Species richness and floristic composition in the channelized river segment were found to be similar to those at the more ‘natural’ river sites. Floral differences were primarily consistent with the dominance of cover by certain species. However, there were significant differences in exotic and native species richness and cover between the ‘natural’ corridor and the channelized segment, which was more susceptible to invasion by exotic perennial taxa, such as Eryngium pandanifolium, Paspalum paspalodes, Tradescantia fluminensis and Acacia dealbata. Factorial and canonical correspondence analyses revealed considerable patchiness in the distribution of species assemblages. The latter were associated with small differences in substrate composition and their own relative position across the banks and along the river segments in question. Data was also subjected to an unweighted pair-group arithmetic average clustering, and the Indicator Value methodology was applied to selected cluster noda in order to obtain significant indicator species. Copyright © 2001 John Wiley & Sons, Ltd.

Exotic and native vegetation establishment following channelization of a western Iberian river

AbstractChannelization is often a major cause of human impacts on river systems. It affects both hydrogeomorphic features and habitat characteristics and potentially impacts riverine flora and fauna. Human‐disturbed fluvial ecosystems also appear to be particularly vulnerable to exotic plant establishment. Following a 12‐year recovery period, the distribution, composition and cover of both exotic and native plant species were studied along a Portuguese lowland river segment, which had been subjected to resectioning, straightening and two‐stage bank reinforcement, and were compared with those of a nearby, less impacted segment. The species distribution was also related to environmental data. Species richness and floristic composition in the channelized river segment were found to be similar to those at the more ‘natural’ river sites. Floral differences were primarily consistent with the dominance of cover by certain species. However, there were significant differences in exotic and native species richness and cover between the ‘natural’ corridor and the channelized segment, which was more susceptible to invasion by exotic perennial taxa, such as Eryngium pandanifolium, Paspalum paspalodes, Tradescantia fluminensis and Acacia dealbata. Factorial and canonical correspondence analyses revealed considerable patchiness in the distribution of species assemblages. The latter were associated with small differences in substrate composition and their own relative position across the banks and along the river segments in question. Data was also subjected to an unweighted pair‐group arithmetic average clustering, and the Indicator Value methodology was applied to selected cluster noda in order to obtain significant indicator species. Copyright © 2001 John Wiley & Sons, Ltd.

Identifying the natural flow regime and the relationship with riparian vegetation for two contrasting western Australian rivers

AbstractThe natural flow regime and the relationship between flows and riparian vegetation are described for sites on both the Blackwood River in south‐western Australia and the Ord River in north‐western Australia. Analysis of long‐term flow data showed the historic mean monthly river discharge for the Blackwood River is strongly seasonal and highly predictable with generally low variability each month. The Ord River showed a strong seasonality of flows with about 92% of the (total) yearly flow occurring between December and March. Flow variability was very high (e.g. coefficient of variation >100% for all months) but highly predictable, with this mostly attributed to low but constant dry‐season flows. Water depth, duration of flood events and the number of flood events per year show a significant correlation with aspects of the riparian vegetation within experimental vegetation plots. Results highlight the strong relationship between floristics, life form structure and population dynamics with stream hydrology. On the Blackwood River, species richness and cover of shrubs reduced with increased duration and frequency of flooding, while cover of exotic species and annual herbs increased with increased flooding. Germination of tree seedlings was not influenced by flood regime but size class of tree species increased with flooding frequency. On the Ord River, species richness was not influenced by flooding regime. However, cover of perennial grasses increased with flooding frequency whilst cover of shrubs decreased. There was no relationship between flooding and seedling establishment whilst tree size class decreased with increased flooding. The methods described here can be used to compare the response of different components of the riparian vegetation to different fluvial regimes (e.g. because of impoundment and abstraction). This technique can be expanded for the management of riparian zones and planning rehabilitation programmes. It may also be useful for improving the ecological knowledge base for setting environmental flows in regulated systems. Copyright © 2001 John Wiley & Sons, Ltd.

Patterns of plant invasions: a case example in native species hotspots and rare habitats

Land managers require landscape-scale information on where exotic plant species have successfully established, to better guide research, control, and restoration efforts. We evaluated the vulnerability of various habitats to invasion by exotic plant species in a 100,000 ha area in the southeast corner of Grand Staircase-Escalante National Monument, Utah. For the 97 0.1-ha plots in 11 vegetation types, exotic species richness (log10) was strongly negatively correlated to the cover of cryptobiotic soil crusts (r =− 0.47, P< 0.001), and positively correlated to native species richness (r = 0.22, P< 0.03), native species cover (r = 0.23, P< 0.05), and total nitrogen in the soil (r = 0.40, P< 0.001). Exotic species cover was strongly positively correlated to exotic species richness (r = 0.68, P< 0.001). Only 6 of 97 plots did not contain at least one exotic species. Exotic species richness was particularly high in locally rare, mesic vegetation types and nitrogen rich soils. Dry, upland plots (n = 51) had less than half of the exotic species richness and cover compared to plots (n = 45) in washes and lowland depressions that collect water intermittently. Plots dominated by trees had significantly greater native and exotic species richness compared to plots dominated by shrubs. For the 97 plots combined, 33% of the variance in exotic species richness could be explained by a positive relationship with total plant cover, and negative relationships with the cover of cryptobiotic crusts and bare ground. There are several reasons for concern: (1) Exotic plant species are invading hot spots of native plant diversity and rare/unique habitats. (2) The foliar cover of exotic species was greatest in habitats that had been invaded by several exotic species. (3) Continued disturbance of fragile cryptobiotic crusts by livestock, people, and vehicles may facilitate the further invasion of exotic plant species.

Rapid assessment of butterfly diversity in a montane landscape

We present the results of a rapid assessment of butterfly diversity in the 754 ha Beaver Meadows study area in Rocky Mountain National Park, Larimer County, Colorado. We measured butterfly species richness and relative abundance as part of a landscape-scale investigation of diversity patterns involving several groups of organisms. A stratified random sampling design was used to include replication in both rare and common vegetation types. We recorded 49 butterfly species from the twenty-four 0.1 ha plots that were sampled four times during June, July, and August 1996. Butterfly species richness, diversity, and uniqueness were highest in quaking aspen (Populus tremuloides Michaux) groves and wet meadows, which occupy only a small proportion of the studied landscape. This result supports the suggestion that aspen areas represent ‘hotspots’ of biological diversity in this montane landscape. Patterns of butterfly species richness were positively correlated with total vascular plant species richness (r = 0.69; P < 0.001), and native plant species richness (r = 0.64; P < 0.001). However, exotic plant species richness (r = 0.70; P < 0.001) and the cover of exotic plant species (r = 0.70; P < 0.001) were the best predictors of butterfly species richness.

Size of the local species pool determines invasibility of a C4‐dominated grassland

The size of the local species pool (i.e., species surrounding a community capable of dispersal into that community) and other dispersal limitations strongly influence native plant community composition. However, the role that the local species pool plays in determining the invasibility of communities by exotic plants remains to be evaluated. We hypothesized that the richness and abundance of exotic species would be greater in C4‐dominated grassland communities if the local species pool included a larger proportion of exotic species. We also predicted that an increase in the exotic species pool would increase the invasibility of sites thought to be resistant to invasion (annually burned grassland). To test these hypotheses, study plots were established within two long‐term (&gt;20 yr) fire experiments at a tallgrass prairie preserve in NE Kansas (USA). Study plots were surrounded by either a small pool of exotic species (small species pool (SSP) plots; six species) or a larger exotic species pool (large species pool (LSP) plots; 18 species). We found that richness and absolute cover of exotic species was significantly (P&lt;0.001) lower (∼70 and 90%, respectively) in annually burned compared to unburned plots, regardless of the size of the exotic species pool. As predicted, exotic species richness was higher (P&lt;0.001) for LSP plots (3.9 per 250 m2) than for SSP plots (0.7 per 250 m2); however, absolute cover was unaffected by the size of the exotic species pool. In the absence of fire, plots with a LSP had four times as many exotic species than SSP plots. An increase in the local exotic species pool also increased the invasibility of annually burned grassland. Indeed, richness of exotic plant species in annually burned LSP plots did not differ from unburned plots with a SSP, indicating that a larger pool of exotic species countered the negative effects of fire. These findings have important implications for predicting how the invasion of plant communities may respond to human‐induced global changes, such as habitat fragmentation. Community characteristics or factors such as frequent fires in grasslands may impart resistance to invasions by exotic species in large, intact ecosystems. However, when a large pool of exotic species is present, frequent fire may not be sufficient to limit the invasions of exotic plants in fragmented landscapes.

Seed Banks of Maritime Chaparral and Abandoned Roads: Potential for Vegetation Recovery

germinating were similar in the two habitats. Burning reduced the number of native species and seeds that germinated, and to a lesser extent the number of exotic species and seeds that germinated from the seed bank. In the field survey, intact chaparral areas had greater numbers of native species and greater native cover than roads, whereas road areas had greater numbers and cover of exotic species. Several species were found only in field surveys; in particular, few chaparral shrub seeds germinated from the seed bank. Similarity between the seed bank and vegetation was low in both chaparral and road sites. The results suggest that although many native species of chaparral plants are present in the seed bank, planting of some species may help accelerate recovery in disturbed areas.

The impact of suburbanization on remnant coastal vegetation in Hobart, Tasmania

Abstract. Changes in vascular species composition and abundance were examined in coastal vegetation at Hobart, Tasmania, Australia over a period of 13 yr, during which suburbanization extended to cover a large proportion of its hinterland. There were significant increases in the richness and cover of exotic vascular plant species, most of which derived from the Mediterranean basin or southern Africa. While most of the introduced species were confined to vegetation with a high exotic component, and close to domestic gardens, some, including the South African coastal shrub, Chrysanthemoides monilifera, expanded in otherwise native vegetation relatively remote from gardens. The impact of proximity to gardens on the relative abundance of exotics and natives in coastal vegetation in 1997 best correlated with the pattern of distance from gardens in 1966, suggesting that there is, at least, a 30‐yr lag period in transformation from largely native to substantially exotic vegetation. This lag period provides an opportunity to reduce exotic invasion problems before they become severe.

Using multi-scale sampling and spatial cross-correlation to investigate patterns of plant species richness

Land managers need better techniques to assess exotic plant invasions. We used the cross-correlation statistic, IYZ, to test for the presence of spatial cross-correlation between pair- wise combinations of soil characteristics, topographic variables, plant species richness, and cover of vascular plants in a 754 ha study site in Rocky Mountain National Park, Colorado, U.S.A. Us- ing 25 large plots (1000 m 2 ) in five vegetation types, 8 of 12 variables showed significant spatial cross-correlation with at least one other variable, while 6 of 12 variables showed significant spatial auto-correlation. Elevation and slope showed significant spatial cross-correlation with all variables except percent cover of native and exotic species. Percent cover of native species had significant spatial cross-correlations with soil variables, but not with exotic species. This was probably because of the patchy distributions of vegetation types in the study area. At a finer resolution, using data from ten 1 m 2 subplots within each of the 1000 m 2 plots, all variables showed significant spatial auto- and cross-correlation. Large-plot sampling was more affected by topographic factors than species distribution patterns, while with finer resolution sampling, the opposite was true. However, the statistically and biologically significant spatial correlation of native and exotic species could only be detected with finer resolution sampling. We found exotic plant species invading areas with high native plant richness and cover, and in fertile soils high in nitrogen, silt, and clay. Spatial auto- and cross-correlation statistics, along with the integration of remotely sensed data and geographic information systems, are powerful new tools for evaluating the patterns and distribution of native and exotic plant species in relation to landscape structure.

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.

How Grazing and Soil Quality Affect Native and Exotic Plant Diversity in Rocky Mountain Grasslands

We used multiscale plots to sample vascular plant diversity and soil characteristics in and adjacent to 26 long-term grazing exclosure sites in Colorado, Wyoming, Montana, and South Dakota, USA. The exclosures were 7–60 yr old (31.2 ± 2.5 yr, mean ± 1 se). Plots were also randomly placed in the broader landscape in open rangeland in the same vegetation type at each site to assess spatial variation in grazed landscapes. Consistent sampling in the nine National Parks, Wildlife Refuges, and other management units yielded data from 78 1000-m2 plots and 780 1-m2 subplots. We hypothesized that native species richness would be lower in the exclosures than in grazed sites, due to competitive exclusion in the absence of grazing. We also hypothesized that grazed sites would have higher native and exotic species richness compared to ungrazed areas, due to disturbance (i.e., the intermediate-disturbance hypothesis) and the conventional wisdom that grazing may accelerate weed invasion. Both hypotheses were soundly rejected. Although native species richness in 1-m2 subplots was significantly higher (P < 0.05) in grazed sites, we found nearly identical native or exotic species richness in 1000-m2 plots in exclosures (31.5 ± 2.5 native and 3.1 ± 0.5 exotic species), adjacent grazed plots (32.6 ± 2.8 native and 3.2 ± 0.6 exotic species), and randomly selected grazed plots (31.6 ± 2.9 native and 3.2 ± 0.6 exotic species). We found no significant differences in species diversity (Hill’s diversity indices, N1 and N2), evenness (Hill’s ratio of evenness, E5), cover of various life-forms (grasses, forbs, and shrubs), soil texture, or soil percentage of N and C between grazed and ungrazed sites at the 1000-m2 plot scale. The species lists of the long-ungrazed and adjacent grazed plots overlapped just 57.9 ± 2.8%. This difference in species composition is commonly attributed solely to the difference in grazing regimes. However, the species lists between pairs of grazed plots (adjacent and distant 1000-m2 plots) in the same vegetation type overlapped just 48.6 ± 3.6%, and the ungrazed plots and distant grazed plots overlapped 49.4 ± 3.6%. Differences in vegetation and soils between grazed and ungrazed sites were minimal in most cases, but soil characteristics and elevation were strongly correlated with native and exotic plant diversity in the study region. For the 78 1000-m2 plots, 59.4% of the variance in total species richness was explained by percentage of silt (coefficient = 0.647, t = 5.107, P < 0.001), elevation (coefficient = 0.012, t = 5.084, P < 0.001), and total foliar cover (coefficient = 0.110, t = 2.104, P < 0.039). Only 12.8% of the variance in exotic species cover (log10cover) was explained by percentage of clay (coefficient = −0.011, t = −2.878, P < 0.005), native species richness (coefficient = −0.011, t = −2.156, P < 0.034), and log10N (coefficient = 2.827, t = 1.860, P < 0.067). Native species cover and exotic species richness and frequency were also significantly positively correlated with percentage of soil N at the 1000-m2 plot scale. Our research led to five broad generalizations about current levels of grazing in these Rocky Mountain grasslands: (1) grazing probably has little effect on native species richness at landscape scales; (2) grazing probably has little effect on the accelerated spread of most exotic plant species at landscape scales; (3) grazing affects local plant species and life-form composition and cover, but spatial variation is considerable; (4) soil characteristics, climate, anddisturbances may have a greater effect on plant species diversity than do current levels of grazing; and (5) few plant species show consistent, directional responses to grazing or cessation of grazing.