We investigated the relative importance of stand and landscape characteristics in the invasion of a nonnative shrub, Amur honeysuckle, in 40 woodlots in an agricultural matrix in southwest Ohio. We quantified stand characteristics that could influence invasibility, the intrinsic susceptibility of an area to invasion, including woodlot size, perimeter-to-area ratio, tree basal area, and stand age. At the landscape scale we included factors that potentially influence propagule rain (the contribution of seeds from individuals established outside the focal area), including the land cover and road density in a 1,500-m buffer around each woodlot, as well as the extent to which the perimeter was forested at two points in the past, and latitude (based on an apparent south-to-north invasion in this region). Based on stepwise regression, we determined that honeysuckle cover was determined primarily by landscape parameters, particularly the percent of the buffer comprised of cropland. Woodlots surrounded by more cropland had less honeysuckle cover, which we attribute to paucity of nearby seed sources and/or minimal movement of seed-dispersing animals. From these findings, we argue that impediments to propagule rain are more important in shaping the invasion of this exotic shrub than are characteristics of the woodlots themselves, i.e., community invasibility.

One of Australia's most serious weeds, Chrysanthemoides monilifera subsp. rotundata (bitou bush) was recently found for the first time in Western Australia as a well established population in Kwinana, a major port and industrial area south of Perth, the State's capital. This population is remote from other bitou bush infestations in Australia and had escaped detection despite extensive surveys in the same State for the other subspecies that is present in Australia, Chrysanthemoides monilifera subsp. monilifera (boneseed). The main reasons it went undetected are thought to be the tightly controlled access to this area because of mineral processing and port activities, the unusual invasion route via a heavy industrial area and the morphological similarity to a native species when it is not flowering. Two surveys defined the core population of 1038 plants that are spread along the coast over a 25-ha semi-circle with about a 500-m (1640 ft) diameter. Subsequent surveys of first a 500 m buffer zone and later a 1-km (0.621 mi) buffer found four additional plants, indicating that there is considerable potential for dispersal. We concluded that the survey has not delimited the distribution because of the potential and evidence for long distance dispersal. Cooperation by the various land managers has led to all plants being killed, as an initial step to management of this species. Other steps to be undertaken include an awareness campaign in the area that would need to be surveyed for delimitation of the spatial distribution and seed bank assessment to measure potential dispersal both in space and through time. It remains to be determined what is the best strategic response: eradication or containment.

Prioritizing management of invasive plants is important for large land management entities, such as federal and state public land stewards, because management resources are limited and multiple land uses and management objectives are differentially impacted. Management decisions also have important consequences for the likelihood of success and ultimate cost of control efforts. We applied multi-criteria decision analysis methods in a geographic information system using natural resource and land use data from Fort Bragg, North Carolina. Landscape-scale prioritization was based on a hierarchical model designed to increase invasive plant management efficiencies and reduce the risk of impacts to key installation management goals, such as training-land management and protected species conservation. We also applied spatial sensitivity analyses to evaluate the robustness of the prioritization to perturbations of the model weights, which were used to describe the relative importance of different elements of the hierarchical model. Based on stakeholders' need for confidence in making management investments, we incorporated the results of the sensitivity analysis into the decision-making process. We identified high-priority sites that were minimally affected by the weight perturbations as being suitable for up-front management and evaluated how adopting this strategy affected management area, locations, and costs. We found that incorporating the results of the sensitivity analysis led to a reduced management area, different target locations, and lower costs for an equal area managed. Finally, we confirmed the distinctiveness of the approach by comparing this same subset of prioritized sites with locations representing species-centric strategies for three invasive plants and their aggregate distribution. By supplying pragmatic information about the localized effects of weighting uncertainty, spatial sensitivity analyses enhanced the invasive plant management decision-making process and increased stakeholder confidence.

Many restoration projects rely on invasive plant removal to restore ecosystems. However, success of restoration efforts relying on invasive removal can be jeopardized, because in addition to displacing native plants, invasives can also dramatically impact soils. Many studies have documented invasives’ effects on soil chemistry and microbiota. While European beachgrass [Ammophila arenaria (L.) Link] is a worldwide invasive problem in coastal dunes outside northern Europe, little attention has been paid to effects of this species on soil chemistry following invasion, even though it establishes persistent, dense monocultures. In our study, we evaluated effects of A. arenaria invasion on soil chemistry of coastal dunes at Point Reyes National Seashore (PRNS); persistence of effects following removal by mechanical or herbicide treatment (legacy effects); and effects of treatment independent of invasion. Dune restoration efforts at PRNS have met with mixed success, especially in herbicide-treated backdunes, where decomposition of dead A. arenaria has been greatly delayed. Based on results, invasion impacted 74% of 19 variables assessed, although there was a significant interaction in many cases with successional status (earlier vs. later). Almost 60% of invasion effects persisted after restoration, with legacy effects prevalent in herbicide-treated backdunes where sand deposition from adjacent beaches could not mitigate effects as it could in herbicide-treated foredunes. Mechanical removal—or inversion of invaded surface soils with less-contaminated subsoils—resulted in fewer legacy effects, but more treatment effects, primarily in backdunes. Soil chemistry may decelerate decomposition of A. arenaria due to the limited nitrogen (N) available to enable microbial breakdown of the high carbon(C):N (70.8:1) material, but microbial factors probably play a more important role. Success of restoration at PRNS may not be fully realized until legacy effects are resolved through additional actions such as inoculation with healthy microbiomes or necromass reduction through controlled burning.

Field and garden experiments were performed to investigate if pathogens, generalist herbivores, or a specialist herbivore (the tortricid moth Pristerognatha fuligana) have the potential to control the invasive Himalayan balsam. Impacts of generalist herbivores and pathogenic fungi were excluded by using thiacloprid and trifloxystrobin. Results show no effect of any antagonist in the field experiment, though we found a modest impact of the combined influence of generalist herbivores and pathogenic fungi in the garden experiment, i.e., under suboptimal growing conditions. Limited information from the native area (the Himalayas) suggests a strong impact of specialist herbivores, thus confirming the enemy release hypothesis. We predict that impact studies of native specialized herbivores will clearly indicate antagonists from the native area with a strong impact on Himalayan balsam in its invaded area.

Chemical, cultural, and biological methods have been developed to control leafy spurge in a variety of environments. Aphthona spp. biological control agents have established throughout the northern Great Plains and Rocky Mountain region and successfully controlled leafy spurge in many areas, but notable exceptions include areas with sandy soils. Leafy spurge control can be improved by combining methods such as chemical, biological, or cultural treatments, compared to a single method used alone. The effects of Aphthona spp., imazapic herbicide, and interseeded native grass species alone or in combination for leafy spurge control were evaluated at two locations in southeastern North Dakota for 5 yr. Both the Sheyenne National Grassland (SNG) and Walcott, ND, study locations had greater than 80% sand soil. Leafy spurge stem density, canopy cover, and yield were reduced for 1 to 2 yr in all treatments that included imazapic, with no difference in control between single and combination treatments. Aphthona spp. and interseeded native grasses alone or combined did not reduce leafy spurge density or cover. Aphthona spp. population remained low throughout the study at both locations. Forb yield increased during the study at the SNG but not the Walcott location. Conversely, warm-season grass yield increased at Walcott but not at the SNG. Leafy spurge stem density declined from 92 to 50 stems/m2 in 5 yr at the SNG site. The decline could not be attributed to specific treatments applied in this study and may be due to self-limitation or soil pathogens.

Yellowflag iris, native to Europe, is a rhizomatous, emergent, invasive plant found in pond margins, ditches, and other wetland sites in much of the United States. In water depths up to approximately 50 cm, it forms dense stands, which displace native sedges and rushes, reducing waterfowl habitat and water flow. The rhizomes can reach 6 m in lateral spread, making it very difficult to control by mechanical methods. In addition, conventional boom-sprayer applications are often impractical in most aquatic systems. Drizzle application is a technique for directed treatment of hard-to-reach invasive plants. It uses low volumes (26 to 104 L ha—1) of concentrated herbicide solution, applied using a spray gun emitting a thin stream of solution with an effective range of 6 m. In this study, conducted along the margins of two ponds at the University of California, Davis, we compared drizzle applications of glyphosate, imazapyr, and triclopyr to applications using a conventional boom sprayer. Although both glyphosate and imazapyr provided excellent control (> 96 %) of yellowflag iris with either treatment technique, only the drizzle treatments of imazapyr at 2.26 and 4.52% ae (10 and 20% product) at spray volumes of 52 and 26 L ha—1, respectively, were below the maximum labeled rate and still gave > 98 % control. Furthermore, a cost analysis indicated that the most economical application for effective control of yellowflag iris was a drizzle application of imazapyr at 4.52 % ae (20 % product) at 26 L ha—1. This study demonstrates that drizzle application with imazapyr can be a practical application method for yellowflag iris control in aquatic systems in which broadcast treatments with conventional boom sprayers may be difficult.

The Detroit Lakes chain of lakes consists of five basins in northwest Minnesota adjacent to the town of Detroit Lakes. Flowering rush has been established in these basins since the 1960s. We evaluated the distribution of flowering rush in the five basins using a point intercept method, with 830 points distributed in a grid with points 150 m apart. These data were analyzed to determine whether invasive and native species frequencies were different between 2010 and 2011. We also assessed co-occurrence of flowering rush with native hardstem bulrush. The distribution of both flowering rush and hardstem bulrush was unchanged from 2010 to 2011. Flowering rush is invading areas with native plants and not establishing in unvegetated areas. Although flowering rush is found as deep as 4.5 m, it is most frequent at a depth of 1.3 m. We also examined the distribution of biomass and growth across a depth gradient from 0.3 to 3.0 m in 0.3-m intervals. At each 0.3-m interval, three biomass samples were collected at each of 10 transects for a total of 30 samples per depth interval or 300 biomass samples. At each point, leaf height, emergent leaf height, water depth, number of ramets, and number of rhizome buds were counted. Biomass samples were collected in a 0.018-m2 core sampler, sorted to shoots and belowground biomass. We found that flowering rush height and biomass peaked at 1.3 m and declined with greater depth. Bud density was negatively related to water depth. Bud density averaged 300 buds m–2, which was three times the average ramet density (100 ramets m–2).

The genus Asparagus includes at least six invasive species in Australia. Asparagus aethiopicus and A. africanus are invasive in subtropical Australia, and a third species, A. virgatus is naturalized and demonstrates localized spread in south east Queensland. To better understand how the attributes of these species contribute to their invasiveness, we compared fruit and seed traits, germination, seedling emergence, seed survival, and time-to-maturity. We further investigated dispersal ecology of A. africanus, examining the diet of a local frugivore, the figbird (Sphecotheres viridis) and the effect of gut passage on seedling emergence. Overall, A. aethiopicus was superior in germination and emergence, with the highest mean germination (98.8%) and emergence (94.5%) under optimal conditions and higher emergence (mean of 73.3%) across all treatments. In contrast, A. africanus had the lowest germination under optimal conditions (71.7%) and low mean seedling emergence (49.5%), but had fruits with the highest relative yield (ratio of dry pulp to fruit fresh weight) that were favored by a local frugivore. Figbirds consumed large numbers of A. africanus fruits (∼30% of all non-Ficus fruits), and seedling germination was not significantly affected by gut passage compared to unprocessed fruits. Asparagus virgatus germinated poorly under cool, light conditions (1.4%) despite a high optimum mean (95.0%) and had low mean performance across emergence treatments (36.3%). The species also had fruits with a low pulp return for frugivores. For all species, seed survival declined rapidly in the first 12 mo and fell to < 3.2% viability at 36 mo. On the basis of the traits considered, A. virgatus is unlikely to have the invasive potential of its congeners. Uniformly short seed survival times suggest that weed managers do not have to contend with a substantial persistent soil-stored seed bank, but frugivore-mediated dispersal beyond existing infestations will present a considerable management challenge.

Callery pear (Pyrus calleryana Decne.) was introduced to North America as an ornamental tree in the early 1900s. Due to widespread planting, P. calleryana has become common throughout the eastern United States and has invaded natural areas, especially disturbed areas. Prescribed fire is a common management technique in prairie ecosystems to mimic natural disturbances. We tested the effectiveness of prescribed fire as a control technique for P. calleryana in a managed prairie system. Fire top-killed all established P. calleryana individuals. However, these individuals responded to fire with 3 to 4 epicormic sprouts each. Similar sprouting behavior occurred in 2-yr-old seedlings. Exposed seeds, fruits, and 1-yr-old seedlings were killed by fire. Established P. calleryana were single-stemmed individuals before exposure to fire. After the prescribed fire, they all were multistemmed, which increased the potential flower-bearing stems within the prairie. We conclude that fire alone is not a suitable technique for managing P. calleryana invasion. Cut and herbicide application methods are labor-intensive. However, combining cut and spray methods with prescribed fire may be effective. Fire removes standing grass and forb biomass, leaving exposed P. calleryana stems, which would make locating individuals and directly applying herbicides easier.

Several studies have been carried out to evaluate the main drivers behind biological invasions and their ecological consequences. Nevertheless, it is still extremely difficult to acquire a full understanding of the invasion process due to its high level of complexity. The problems that complicate invasion studies are low detection during the early stages of invasion, and the high survey cost of working over large, inaccessible, and rugged areas. The studies that develop efficient tools to reduce costs and time will help to control or mitigate the invaders' damaging effects. Hieracium pilosella is an aggressive invader of grasslands worldwide. The aim of this work was to conduct a regional assessment for the suitability of H. pilosella invasion in the Fuegian Steppe by combining field surveys, spatial modeling, and geographic information system tools. We recorded the invader cover in 167 sample sites and we determined the relationship between environmental variables and the probability of invasion. This was carried out by the selection of alternative generalized linear models. The best model indicates that disturbance and plant community were the main drivers to predict invasion suitability on a regional scale. Therefore, these two variables were used as main inputs to construct a regional invasion suitability map, which identifies the most critical areas for prevention, control, and further monitoring. This approach corresponds to a tool that permits us to evaluate the suitability of invasion even over large and inaccessible areas. The results especially warn about soil disturbance associated with usual management practices in Fuegian rangelands.

The common edible fig is a subcanopy tree that has invaded many of the remnant riparian forests of California's Central Valley. Fig is unusual in its ability to invade low-light, low-disturbance, native-plant–dominated environments. Dendrochronology combined with regression and spatial analyses allowed us to empirically quantify the expansion rate and spatial pattern of the fig invasion into the native plant community at Caswell Memorial State Park (Ripon, CA) over a 70-year invasion period. Fig uses a combination of short-distance dispersal, which results in constant, linear expansion at source population sites and long-distance dispersal, which eventually leads to high recruitment of satellite populations in ideal environments. Although fig initially experienced a long lag in its invasion rate, at the time of this study, it was expanding at an exponential rate at the landscape scale in Caswell. We identified a number of characteristics intrinsic to the fig population (shade suppression, pollinator presence, highly specialized reproduction, and propagule pressure) as well as extrinsic characteristics of the receiving environment (hydrologic alteration from the construction of a dam, safe sites for juvenile recruitment, and target effects from environmental heterogeneity) that may have influenced the rate and pattern of fig invasion. The Central Valley riparian forests have been reduced to less than 6% of their original area, and invasive fig is a significant threat to the remaining fragments of this important vegetation community. We include suggestions for fig eradication based on knowledge gained in this study.

The importance of dispersal in the establishment and proliferation of exotic populations make this life history stage critical in the prediction and management of biological invasions. We observed the dispersal of seeds by patches of smooth brome invading northern fescue prairies and applied an inverse power function model to explore its potential invasion patterns. Based on our observations of two northern fescue prairies in Riding Mountain National Park, Manitoba, Canada, patterns of potential invasion were contingent upon the dispersal of seeds as individual florets or aggregated within spikelets and panicles. For example, although the majority of dispersed seeds were intercepted within one meter, inside and outside the margins of invading patches, slopes of the log–log plots of seed number against their dispersal distance were steeper for seeds dispersed as spikelets than individual florets. Despite the observed aggregation of seeds along the margins of invading patches, the number of dispersed seeds was poorly correlated with that germinated from the seed bank. The shallow dispersal gradient of individual florets and spikelets, combined with the steeper gradient of panicles suggest that smooth brome is capable of simultaneously invading along dense fronts and by establishing isolated foci. Although low correlations between the number of dispersed seeds and their recruitment from the seed bank might suggest postdispersal transport of seeds, other mechanisms, including seed predation and pathogens, remain unexplored. Conservation and restoration of northern fescue prairies must include efforts to control the dispersal of smooth brome seeds and reduce opportunities for their establishment.

Hybridization between species has the potential to change invasion dynamics. Field observations suggest that spotted knapweed and diffuse knapweed, two ecologically and economically destructive invasive plants, hybridize in their introduced range. As a first step towards understanding whether hybridization has affected the dynamics of the invasion of these species, we conducted field surveys in the introduced (North American) and native (European) ranges to discern patterns of hybridization and measured fitness-related traits among field hybrids and parental species. In North America we detected plants with hybrid morphology in 97% of the diffuse knapweed sites (n = 40); such hybrid plants were taller and more often exhibited polycarpy than plants with typical diffuse knapweed morphology. Hybrids were not detected in North American spotted knapweed sites (n = 22). In most regions surveyed in Europe, diffuse knapweed and spotted knapweed were isolated from each other and existed as distinct, nonhybridizing species. However, in Ukraine, the two species frequently coexisted within a site, resulting in hybrid swarms. On average, the plants from the North American diffuse knapweed sites (including plants with both diffuse and hybrid morphology), were larger than the apparently pure diffuse knapweed in the native range. The cross-continental patterns of hybridization likely are explained by differences in cytology. It recently has been confirmed that the spotted knapweed in North America is tetraploid whereas the diffuse knapweed is diploid. Genetic incompatibilities associated with these two cytotypes likely prevent ongoing hybridization. We hypothesize that hybrid individuals were introduced to North America along with diffuse knapweed. Because plants with hybrid morphology are found in nearly all North American diffuse knapweed sites, the introduction of hybrids likely occurred early in the invasion of diffuse knapweed. Thus, although the presence of hybrids might facilitate the ongoing invasion of diffuse knapweed into North America, elevated concern regarding their presence might not be warranted. Because such individuals are not likely to represent a new hybridization event, currently effective management strategies used in diffuse knapweed sites should not need alteration.

Parrotfeather is an invasive, aquatic plant in the United States that is native to South America. It has impaired the use of water bodies throughout the United States and is difficult to control, despite using a variety of management techniques. Our objectives were to examine the efficacy of subsurface applications of seven herbicides labeled for aquatic use and to compare those applications to herbicides that can also be applied to emergent foliage. A replicated mesocosm study was conducted in 378-L (100-gal) tanks beginning in August 2007 and repeated during the same period in 2008. The maximum and half-maximum labeled rates of copper chelate, diquat, endothall, fluridone, triclopyr, and carfentrazone-ethyl were applied to the water column in designated mesocosms. The maximum labeled rate for foliar applications of diquat, triclopyr, and 2,4-D were used to compare treatment methods. Six weeks after treatment (WAT), copper, endothall, fluridone, and carfentrazone-ethyl did not achieve 90% control; in fact, control was less than 50% for each herbicide, and therefore, the herbicides were not considered efficacious for controlling parrotfeather. Diquat at all rates and application methods resulted in 70 to 90% biomass reduction. Triclopyr, with both the highest aqueous concentration and foliar application, resulted in an 84 and 86%, respectively, reduction in biomass at 6 WAT. The foliar application of 2,4-D was the only herbicide and application method that resulted in ≥ 90% biomass reduction of parrotfeather. In these studies, regrowth occurred in all tanks regardless of herbicide or treatment method, indicating multiple applications would be necessary to provide longer-term plant control. Future research should identify possible herbicide combinations or timing of applications to maximize treatment efficacy.

Strong competitive ability of introduced plant species has been frequently stated as a key factor promoting successful invasion. The dynamics of invasive species may depend on their abilities to compete for resources and exploit disturbances relative to native species. This study compares the growth performance of the invasive blackberry (Rubus niveus Thunb.) with four of the most common woody native species of the Scalesia forest in the Galapagos Islands. Using a series of greenhouse and field studies, the growth rate and biomass production of native species alone and in competition with R. niveus was compared under different water and light stress conditions. Rubus niveus showed a faster growth rate and biomass production than the native species as well as a broad tolerance to light and water stress conditions. Competitive ability was also assessed by looking at the seedbank and regeneration processes after herbicide control in the field. Although the number of R. niveus seedlings that germinated from the soil samples was considerably larger than that of native species, recruitment of the invasive on the experimental plots was limited. Overall, R. niveus showed superior competitive ability over native species with comparable growth forms, suggesting a possible mechanism that enables R. niveus to successfully invade a wide range of habitats in the Galapagos Islands. Implementing an integrated management strategy including biological control, seedbank reduction, and active revegetation, should improve the efficiency of R. niveus management, enabling the restoration of degraded vegetation in the Galapagos.

Wild chervil is an invasive biennial or short-lived perennial weed introduced into North America that negatively impacts forage production and degrades habitat for native plant species. A 2-yr study using prebloom mowing followed by combinations of herbicide, tillage, and grass seeding was conducted in the Pacific Northwest to identify an effective integrated weed management strategy for this species. By 2 mo after herbicide treatment (MAHT), wild chervil control with glyphosate + ammonium sulfate (AMS) and clopyralid was 83 and 73%, respectively. Tillage with or without herbicide pretreatment resulted in 92 to 98% wild chervil control at 2 MAHT, whereas herbicide without tillage gave only 45% control across all treatments. Tillage with or without subsequent grass seeding reduced wild chervil density four-fold compared to herbicide alone at 9 MAHT. Herbicide + tillage + grass seeding resulted in similar wild chervil cover (1 to 5% cover) as herbicide + tillage (1 to 6% cover) without subsequent grass seeding. Wild chervil biomass at 1 yr after herbicide treatment (YAHT) was reduced to 487 kg ha−1 (439 lb ac−1) with herbicide + tillage compared to 4,256 kg ha−1 for herbicide treatment alone. Herbicide + tillage + grass seeding increased grass dry weight at 1 YAHT from 201 kg ha−1 for herbicide + tillage to 1,575 kg ha−1, compared to 351 kg ha−1 in herbicide-only plots.

Cutleaf teasel (Dipsacus laciniatus) is an exotic, invasive plant that infests roadsides and other minimally disturbed areas. Plants in established stands appear to be a mixture of rapidly growing rosettes and rosettes with developing reproductive structures. Research that is focused on seed characteristics and their contribution to the spread of plants may be a key to precluding spread of cutleaf teasel in the field. Field studies were conducted to determine the viability and germinability of seeds after flowering, seedling emergence patterns, and seed persistence. Flowering (60% of anthesis) was observed under natural conditions on July 24, 2004, and July 16, 2005. Seeds harvested 12 d after flowering exhibited 43% viability and 2.5% germination. Seed weight and viability were greatest 30 d after flowering, but germination was < 32%. Seedling emergence was monitored over a 12-mo period with the greatest emergence in April and October with 33% of seeds germinating. Seed persistence was evaluated over a 3-yr period under field conditions. Up to 84% of the germinated seeds had germinated during the first year, with 6% of seeds remaining viable after 3 yr. Although seed persistence was relatively short, the rapid development of seeds following flowering as well as seedling emergence in both fall and spring suggests management practices are needed throughout the year to restrict reestablishment spread of cutleaf teasel.

The interactive effects of herbivory, exotic species, and other human-mediated changes to the biosphere are reducing species diversity and altering ecosystem services globally. In this study, we tested whether facilitation could be used as a management technique to restore a degraded northeast Wisconsin forest understory facing high white-tailed deer (Odocoileus virginianus) browse pressure and high abundance of the exotic herb garlic mustard [Alliaria petiolata (Bieb.) Cavara & Grande]. Specifically, we attempted to facilitate native understory restoration by planting or seeding native herbs into three different matrix densities of the native, browse-tolerant grass Virginia wildrye (Elymus virginicus L.), which were either protected from (fenced), or accessible to, deer browsing. Deer had minimal impacts on E. virginicus but significantly reduced the cover, survival, and flower production of white snakeroot [Ageratina altissima (L.) King & H.E. Robins.] transplants, largely independent of the density of E. virginicus plantings in open-access plots. In contrast, the richness and abundance of native-seeded species were not affected by deer access but were reduced by increasing E. virginicus densities. Alliaria petiolata cover was significantly higher in plots accessible to deer but declined significantly with increasing E. virginicus planting density in both open-access and fenced plots. These results were largely corroborated by results from an associated observational study, with the exception that natural E. virginicus stands supported slightly higher native-species richness than did adjacent areas lacking E. virginicus. Thus, although we found little support that establishing E. virginicus facilitated browse-susceptible native understory herbs during our short-term experimental study, restored E. virginicus successfully established, thereby increasing native species cover and significantly reducing the cover of the exotic A. petiolata. We suggest the planting of browse-tolerant native species, such as E. virginicus, as a viable restoration technique in heavily browsed, A. petiolata–invaded woodlands.

Hardwood forests in eastern North America are being colonized by multiple nonnative plant and animal species. Colonization rates can be affected by stand structure and distance from edge. We sampled earthworm densities and understory plant species cover in transects located in paired old- and second-growth forests in Indiana. Two 100-m transects were established within each forest stand during late April to early May in each year. One transect was placed parallel to and within 5 m of a south- or west-facing edge. The second transect was placed parallel to the first. but at no less than 100 m from any edge. Nonnative earthworms and plants were found in forest edge and interior regardless of structural stage (second-growth vs. old-growth). The number of native plant species decreased linearly as the densities of adult Lumbricus and Aporrectodea earthworms and the percent cover of multiflora rose (an invasive plant species) increased. Densities of L. terrestris and Aporrectodea earthworms and percent cover of multiflora rose cumulatively explained 39% of the variation in the number of native plant species found in transects across the state. However, multivariate analyses suggested that the species composition of Indiana understory plant communities was affected more by geography than by earthworm densities. Our results suggest that nonnative earthworms and plants are ubiquitous in Indiana hardwood forests and that they may reduce the number of native plant species.