The widespread recognition that nonnative plants can have significant biological and economic effects on the habitats they invade has led to a variety of strategies to remove them. Removal alone, however, is often not sufficient to allow the restoration of altered communities or ecosystems. The invasive plant's effects may persist after its removal thus exerting a “legacy” that influences community composition or the ecosystem properties or both over some ensuing period. Here, we review evidence of such legacy effects on plant and soil communities, soil chemistry, and soil physical structure. We discuss this evidence in the context of efforts to restore community composition and ecosystem function in invaded habitats. Legacies are especially likely to develop in cases where invasive species cause local extirpations of resident species, alter resource pools, and interact with other aspects of global change including land-use changes, atmospheric N deposition, acid rain, and climate change. In cases where legacies of invasive plants develop, the removal of the nonnative species must also be accompanied by strategies to overcome the legacies if restoration goals are to be achieved.

The distribution and abundance of invasive species can be strongly influenced by habitat suitability and by corridors that facilitate dispersal. We synthesize results from a large-scale invasive plant survey with a patch-scale expansion experiment. The large-scale survey involved transects up to 250 m away from of all roads in a 32,000 ha forest. The patch experiment involved initiating invasions in different habitat types (roadside, wetland, disturbed, and intact forests), and then fitting statistical models to patch spread rates. The large-scale survey highlighted the importance of roads in predicting the presence of invasive plants, also revealing that one invasive plant, Microstegium vimineum, has spread rapidly since its purported introduction in 1994. The patch-scale experiments focused on Microstegium and demonstrated that spread rates are higher in roadsides than in forested and wetland patches, even in the absence of major disturbances. These results highlight the importance of landscape features when designing prevention and management practices aimed at limiting invasive plant abundance and spread.

The rate at which plant invasions occur is accelerating globally, and a growing amount of recent research uses genetic analysis of invasive plant populations to better understand the histories, processes, and effects of plant invasions. The goal of this review is to provide natural resource managers with an introduction to this research. We discuss examples selected from published studies that examine intraspecific genetic diversity and the role of hybridization in plant invasion. We also consider the conflicting evidence that has emerged from recent research for the evolution of increased competitiveness as an explanation for invasion, and the significance of multiple genetic characteristics and patterns of genetic diversity reported in the literature across different species invasions. High and low levels of genetic diversity have been found in different invading plant populations, suggesting that either selection leading to local adaptation, or pre-adapted characteristics such as phenotypic plasticity, can lead to aggressive range expansion by colonizing nonnative species. As molecular techniques for detecting hybrids advance, it is also becoming clear that hybridization is a significant component of some plant invasions, with consequences that include increased genetic diversity within an invasive species, generation of successful novel genotypes, and genetic swamping of native plant gene pools. Genetic analysis of invasive plant populations has many applications, including predicting population response to biological or chemical control measures based on diversity levels, identifying source populations, tracking introduction routes, and elucidating mechanisms of local spread and adaptation. This information can be invaluable in developing more effectively targeted strategies for managing existing plant invasions and preventing new ones. Future genetic research, including the use of high throughput molecular marker systems and genomic approaches such as microarray analysis, has the potential to contribute to better understanding and more effective management of plant invasions.

Invasion by cheatgrass and the associated high fire frequency can displace native plant communities from a perennial to an annual grass driven system. Our overall objective of this study was to determine the potential to favor desired native perennial bunchgrasses over annual grasses by altering plant available mineral nitrogen (N). In the first study, we grew cheatgrass and three native bunch grasses (native grasses were combined in equal proportions) in an addition series experimental design and applied one of three N treatments (0, 137, and 280 mg N/kg soil). Regression models were used to derive the effects of intra- and interspecific competition on individual plant yield of cheatgrass and the native bunch grasses (combined). In our second study, we compared the absolute growth rate of the four plant species grown in isolation in a randomized complete block design for 109 days under the same soil N treatments as the competition study. Predicted mean average weight of isolated individuals increased with increasing soil N concentrations for both cheatgrass and the three native perennials (P < 0.05). Biomass of cheatgrass and its competitive ability increased with increasing soil N concentrations (P < 0.0001) compared to the combined native bunchgrasses. However, the greatest resource partitioning occurred at the 137 mg N/kg soil N treatment compared to the 0 (control) and 280 mg N/kg soil treatments, suggesting there may be a level of N that minimizes competition. In the second study, the absolute growth of cheatgrass grown in isolation also increased with increasing N levels (P = 0.0297). Results and ecological implications of this study suggest that increasing soil N leads to greater competitive ability of cheatgrass, and that it may be possible to favor desired plant communities by modifying soil nutrient levels.

Preamble: Executive Order (EO) 13112—defines an invasive species as “an alien species whose introduction does or is likely to cause economic or environmental harm or harm to human health.” In the Executive Summary of the National Invasive Species Management Plan (NISMP) the term invasive species is further clarified and defined as “a species that is non-native to the ecosystem under consideration and whose introduction causes or is likely to cause economic or environmental harm or harm to human health.” To provide guidance for the development and implementation of the NISMP, the National Invasive Species Council (NISC) and the Invasive Species Advisory Committee (ISAC) adopted a set of principles outlined in Appendix 6 of the NISMP. Guiding Principle #1 provides additional context for defining the term invasive species and states “many alien species are non-invasive and support human livelihoods or a preferred quality of life.”

Large-statured invasive grasses (LSIGs) constitute a distinct functional group with characteristic life history traits that facilitate colonization and aggressive growth in aquatic ecosystems, particularly those modified by human activities. These species typically form monocultures in the systems they invade and have wide-ranging and negative impacts on biodiversity and ecosystem processes. In March 2008, a special symposium was held as part of the Western Society of Weed Scientists annual meeting to synthesize our current knowledge of the ecological impacts and management of two notorious LSIGs: Arundo donax and Phragmites australis. In this volume of Invasive Plant Science and Management, symposium participants provide articles summarizing existing knowledge, recent research progress, and research needs for these two taxa. Here, we summarize the basic biology of these species and suggest the use of a more holistic approach to deal with the effects and management of LSIG invasions.

Kentucky bluegrass, a nonnative species, has invaded rangelands in the United States and is currently present in most rangelands across the Northern Great Plains. Despite its accelerated expansion, the consequences of Kentucky bluegrass on the diversity of native plant species and on ecosystem services remain largely unknown. We synthesized the available data related to Kentucky bluegrass and how it affects native plant diversity and ecosystem services. We found that invasion may bring negative consequences to ecosystem services, such as pollination, habitat for wildlife species, and alteration of nutrient and hydrologic cycles, among others. To maintain the flow of ecosystem goods and services from these rangeland ecosystems, range science must adapt to the challenge of introduced, cool-season grass dominance in mixed-grass prairie. Based on our findings, we identify research needs that address ecosystem changes brought on by Kentucky bluegrass invasion and the corresponding effects these changes have on ecosystem services. We are dealing with novel ecosystems, and until we have better answers, adaptive management strategies that use the best available information need to be developed to adapt to the invasion of this pervasive invasive species.

The rapid progression of an invasive genotype of common reed along roads and other linear infrastructures in North America provides one of the most spectacular examples of the role of transportation corridors as invasion pathways. In this paper, we discuss ecological patterns and processes in roadside habitats important for understanding the invasion dynamics of common reed from coastal areas inland. Frequent disturbances in roadsides combined with potentially high levels of nutrients from adjacent land and stress conditions (from deicing salt and other pollutants) mimic the conditions unfortunately found more and more in natural wetlands. The novel contribution of roads is the creation of linear wetlands with an unprecedented level of connectivity. Genetic evidence shows that invasion inland coincides with the intensification of the road network. Time series analysis of remote sensing data reveals impressive rates of invasion of roadsides and other linear infrastructures, suggesting prime conditions for common reed in these novel habitats. Whereas reed dispersal along roads was thought to be largely due to rhizome transport, new evidence suggests a significant contribution of sexual reproduction and seedling establishment, likely enhanced by climate warming at northern latitudes. There is little evidence that other wetland plants can slow down vegetative expansion of common reed in roadside habitats, but plant cover could prevent seedling establishment and shading by shrubs and trees limit lateral clonal expansion. The fact that common reed possibly provides water treatment and other ecosystem services in roadsides must be carefully weighed against the threat to biodiversity in natural systems. All this begs for investigating urgently if, where, and how we should intervene without compromising the great value of wetlands of conservation interest intersected by roads.

In this paper, we describe five successful classical biological weed control agents released in the United States. For each of the five arthropod species, we compared data from prerelease studies that experimentally predicted the agent's host range with data collected postrelease. In general, experimental host range data accurately predicted or overestimated risks to nontarget plants. We compare the five cases with insects recently denied for introduction in the United States and conclude that none of the discussed agents would likely be approved if they were petitioned today. Three agents would be rejected because they potentially could attack economic plants, and two because of potential attack on threatened or endangered plants. All five biocontrol agents have contributed significantly to the successful management of major weeds with no or minimal environmental risk. We believe that the United States may miss opportunities for sustainable and environmentally benign management of weeds using biological control if the regulatory framework only considers the risks of agents as potential plant pests and treats any host-range data regarding economic or threatened and endangered species as a binary decision (i.e., mandates rejection if there is any chance of feeding or development). As a way forward we propose the following: (1) the addition of risk and benefit analyses at the habitat level with a clear ranking of decision-making criteria as part of the U.S. Department of Agriculture Animal and Plant Health Inspection Service Technical Advisory Group's evaluation process of biocontrol agents; (2) recognition of the primacy of realized host range data for potential agents that considers the insect's host selection behavior instead of emphasizing fundamental host range data during release evaluations, and (3) development of formalized postrelease monitoring of target and nontarget species as part of the release permit. These recommendations may initially be advanced through reassessment of current policies but may in the longer term require the implementation of dedicated biocontrol legislation.

Plant invasions can have large effects on ecosystem services. Some plant invaders were introduced specifically to restore key services to ecosystems, and other invaders are having unintended, detrimental effects on services, such as the quantity and quality of water delivered, flood control, erosion control, and food production. Many ecosystem services are difficult to measure directly, and although there are extensive studies on plant invaders and ecosystem processes, a number of challenges prevent us from confidently extrapolating those processes as proxies for services. To extrapolate local, short-term measures of processes to ecosystem services, we must: (1) determine which processes are the key contributors to a service, (2) assess how multiple processes interact to provide a given service, (3) determine how vegetation types and species affect those processes, and (4) explicitly assess how ecosystem services and their controls vary over space and time, including reliance of ecosystem services on “hot spots” and “hot moments” and a minimum size of a vegetation type in the landscape. A given invader can have positive effects on some services and negative effects on others. It is important to consider that, in some systems, shifting environmental conditions may no longer support native species and that invasive species may be critical contributors to the resilience of ecosystem services.

We developed a protocol for categorizing nonnative plants according to their negative impacts on biodiversity in a large area such as a state, nation, or ecological region. Our objective was to provide a tool that makes the process of identifying, categorizing, and listing nonnative plants that cause negative impacts to biodiversity analytic, transparent, and equitable and that yields lists that are useful to researchers, land managers, regulators, consumers, and commercial interests such as the nursery industry. The protocol was designed to distinguish between species that cause high, medium, low, or insignificant negative impacts to native biodiversity within the state, region, or nation of interest. It consists of 20 multiple-choice questions grouped into four sections, which each address a major aspect of a species' total impact and when combined yield an overall “Invasive Species Impact Rank” or “I-Rank” (high, medium, low, or insignificant). The nonprofit organization NatureServe is now using this protocol to assess the estimated 3,500 nonnative vascular plant species that are established in the United States to create a national list prioritized by negative impact on biodiversity. The protocol and additional information are available on the Internet at http://www.natureserve.org/getData/plantData.jsp, and over 500 completed species assessments are available through NatureServe Explorer (http://www.natureserve.org/explorer/).

Screening tools that effectively predict which nonnative species are likely to become invasive are necessary because of the disproportionate ecological and economic costs associated with invaders. We tested the effectiveness of the Australian Weed Risk Assessment system (WRA) in distinguishing plant species that are major invaders, minor invaders, and noninvaders in Florida. The test included 158 annuals and perennials in six growth forms from 52 families in 27 orders. The WRA with a secondary screen met all hypothesized accuracy levels: it correctly rejected 92% of test species that have been documented to be invasive in Florida and correctly accepted 73% of the noninvaders. The incorrect rejection of noninvaders was 8% with the remaining 19% of noninvaders falling into the “evaluate further” outcome. Only 10% of the 158 species required further evaluation. Invaders of natural areas and agricultural systems were identified with equal accuracy. Receiver operating characteristic analysis demonstrated high separation of invaders from noninvaders. The degree to which the WRA is precautionary may be adjusted by altering the cutoff scores that define the “accept, evaluate further,” and “reject” outcomes. This approach could be adopted in Florida as a screening mechanism to reduce importation of new invaders.

Genetic diversity and reproductive characteristics may play an important role in the invasion process. Here, we review the genetic structure and reproductive characteristics of common reed and giant reed, two of the most aggressive, large-statured invasive grasses in North America. Common reed reproduces both sexually and asexually and has a complex population structure, characterized by three subspecies with overlapping distributions; of which, one is introduced, one native, and the third is of unknown origins. These three subspecies show varying levels of genetic diversity, with introduced common reed having high levels of nuclear diversity, indicating that multiple introductions have likely occurred. In contrast, giant reed has low genetic diversity and appears to reproduce solely via asexual fragments yet is highly aggressive in parts of its introduced range. Both species are well-adapted for growth in human-dominated landscapes, which is presumably facilitated by their rhizomatous growth habit.

The common reed invasion in North America has spanned two centuries and is still ongoing. This expansion comprises two main forms: an introduced Eurasian lineage (identified here as “Introduced Phragmites”) and a Gulf Coast lineage of unknown origin (identified here as “Gulf Coast Phragmites”). Both lineages are spreading beyond their current ranges and are colonizing Southwestern and Gulf Coast ecosystems where they have not previously existed. As a result, the native North American lineage of common reed (hereafter “native Phragmites”) has declined in many places. The recent invasion of the U.S. Southwest by Introduced and Gulf Coast Phragmites lineages has made this the only region in the world colonized by all three lineages. Along the central Gulf Coast where Gulf Coast Phragmites remains the dominant form, Introduced Phragmites has also recently invaded the Mississippi River delta. The consequences of these new invasions are uncertain, but a rapid response is needed to protect native species and ecosystems and reduce future control costs.

Kentucky bluegrass was introduced into the present-day United States in the 1600s. Since that time, Kentucky bluegrass has spread throughout the United States and Canada becoming prolific in some areas. In the past century, Kentucky bluegrass has been a presence and often a dominant species in some prairies in the Northern Great Plains. Sometime within the past few decades, Kentucky bluegrass has become the most-common species on the untilled, native prairie sites of much of North and South Dakota. In this article, we hypothesize how Kentucky bluegrass has come to dominate one of the most endangered ecosystems in North America—the prairie—through a historical, ecological, and climatological lens. We urge others to start addressing the invasion of Kentucky bluegrass with both new research and management strategies.

Invasion by annual grasses, such as cheatgrass, into the western U.S. sagebrush-steppe is a major concern of ecologists and resource managers. Maintaining or improving ecosystem health depends on our ability to protect or re-establish functioning, desired plant communities. In frequently disturbed ecosystems, nutrient status and the relative ability of species to acquire nutrients are important drivers of invasion, retrogression, and succession. Thus, these processes can potentially be modified to direct plant community dynamics toward a desired plant community. The overall objective of this review paper is to provide the ecological background of invasion by exotic plants and propose a concept to facilitate the use of soil nitrogen (N) management to achieve desired plant communities that resist invasion. Based on the literature, we propose a model that predicts the outcome of community dynamics based on N availability. The model predicts that at low N levels, native mid- and late-seral species are able to successfully out-compete early-seral and invasive annual species up to some optimal level. However, at some increased level of N, early-seral species and invasive annual grasses are able to grow and reproduce more successfully than native mid- and late-seral species. At the high end of N availability to plants, the community is most susceptible to invasion and ultimately, increased fire frequency. Soil N level can be managed by altering microbial communities, grazing, mowing, and using cover crops and bridge species during restoration. In these cases, management may be more sustainable since the underlying cause of invasion and succession is modified in the management process.

The herbicide imazapic is registered for use on rangelands and provides effective short-term control of certain invasive annual grasses. However, details about optimal application rates for downy brome and susceptibility of simultaneously seeded species are lacking. Thus, we investigated downy brome and seeded species responses to variable rates of imazapic (0, 35, 70, 105, and 140 g ai/ha) in two plant communities (salt desert shrub and Wyoming big sagebrush). In autumn 2003, plots were treated with imazapic and seeded with one of five perennial plant materials (Siberian wheatgrass [‘Vavilov’ and the experimental source Kazak]; prostrate kochia [‘Immigrant’ and the experimental source 6X], and Russian wildrye [‘Bozoisky II’]). Downy brome cover and seeded species establishment were evaluated in spring 2004 and 2006. Downy brome cover in 2004 decreased with increasing imazapic rate at both sites, although more so at the Wyoming big sagebrush site. In 2006, no difference in downy brome cover existed among herbicide rates at the Wyoming big sagebrush site. At the salt desert shrub site, the high rate of imazapic reduced downy brome cover by about 25% compared to untreated plots. ‘Vavilov’ Siberian wheatgrass was the only seeded species with lower downy brome cover in 2006 than 2004. Seeded species establishment increased with imazapic rate in the salt desert shrub community, but in the Wyoming big sagebrush community it peaked at intermediate rates and declined at higher rates. Variation in downy brome control and seeded species establishment might have been associated with differences in precipitation, soil organic matter, and disturbance history between sites. Overall, imazapic was useful for helping establish desirable perennial species, but unless downy brome is reduced below a critical threshold, favorable precipitation can return sites to pretreatment levels within two years.

There has been recent interest in determining the upper limits to the feasibility of weed eradication. Although a number of disparate factors determine the success of an eradication program, ultimately eradication feasibility must be viewed in the context of the amount of investment that can be made. The latter should reflect the hazard posed by an invasion, with greater investment justified by greater threats. In simplest terms, the effort (and hence investment) to achieve weed eradication comprises the detection effort required to delimit an invasion plus the search and control effort required to prevent reproduction until extirpation occurs over the entire infested area. The difficulty of estimating the required investment at the commencement of a weed eradication program (as well as during periodic reviews) is a serious problem. Bioeconomics show promise in determining the optimal approach to managing weed invasions, notwithstanding ongoing difficulties in estimating the costs and benefits of eradication and alternative invasion management strategies. A flexible approach to the management of weed invasions is needed, allowing for the adoption of another strategy when it becomes clear that the probability of eradication is low, owing to resourcing or intractable technical issues. Whether the considerable progress that has been achieved towards eradication of the once massive witchweed invasion can be duplicated for other weeds of agricultural systems will depend to a large extent upon investment (> $250 million over 50 yr in this instance). Weeds of natural ecosystems seem destined to remain more difficult eradication targets for a variety of reasons, including higher impedance to eradication, more difficulty in valuing the benefits arising from eradication, and possibly less willingness to pay from society at large.

Chinese privet is a major invasive shrub within riparian zones throughout the southeastern United States. We removed privet shrubs from four riparian forests in October 2005 with a Gyrotrac® mulching machine or by hand-felling with chainsaws and machetes to determine how well these treatments controlled privet and how they affected plant community recovery. One year after shrub removal a foliar application of 2% glyphosate was applied to privet remaining in the herbaceous layer. Three “desired-future-condition” plots were also measured to assess how well treatments shifted plant communities toward a desirable outcome. Both methods completely removed privet from the shrub layer without reducing nonprivet shrub cover and diversity below levels on the untreated control plots. Nonprivet plant cover on the mulched plots was > 60% by 2007, similar to the desired-future-condition plots and higher than the hand-felling plots. Both treatments resulted in higher nonprivet plant cover than the untreated controls. Ordination showed that after 2 yr privet removal plots were tightly grouped, suggesting that the two removal techniques resulted in the same plant communities, which were distinctly different from both the untreated controls and the desired-future-condition. Both treatments created open streamside forests usable for recreation and other human activities. However, much longer periods of time or active management of the understory plant communities, or both, will be required to change the forests to typical mature forest plant communities.

Downy brome (Bromus tectorum L.)—dominated communities can remain as stable states for long periods, even with frequent disturbance by grazing and fire. The objective of this study was to determine the effectiveness of using targeted cattle grazing and late-season prescribed burning, alone and in combination, to reduce B. tectorum seed bank input and seed bank density and thus alter aboveground community dynamics (species composition) on a B. tectorum–dominated landscape in northern Nevada. Cattle removed 80 to 90% of standing biomass in grazed plots in May of 2005 and 2006 when B. tectorum was in the boot (phenological) stage. Grazed and ungrazed plots were burned in October 2005 and 2006. The combined grazing–burning treatment was more effective than either treatment alone in reducing B. tectorum seed input and seed bank density, and in shifting species composition from a community dominated by B. tectorum to one composed of a suite of species, with B. tectorum as a component rather than a dominant. This study provides a meso-scale precursor for landscape-scale adaptive management using grazing and burning methodologies.