Ecological impacts from invasive plants that have been identified include reductions in biodiversity, changes in resource cycling, and disruptions of ecosystem function. To mitigate these negative ecological impacts, managers work to remove invasive plants. However, removal does not necessarily immediately lead to a return to the uninvaded ecological state. Similarly, the accumulation rate of ecological impacts following invader establishment is almost entirely unknown for most species, hindering identification of optimal management times. The accumulation and loss (so-called legacy effects) of impacts following invader establishment and removal represent an “invasion shadow.” To begin to understand invasion shadows, we measured the changes in biotic and abiotic ecological impacts during establishment and following removal of the forest understory invader Japanese stiltgrass. We found that when the abiotic metrics were considered, seeded areas became more functionally similar to the invaded landscape and removed areas became more similar to the uninvaded landscape. However, while the plant community did not change in a 3-yr period during a new invasion, following invader removal, it became less similar to both the invaded and uninvaded landscape altogether, suggesting legacies. Surprisingly, all changes occurred almost immediately and persisted following invader establishment and removal. Our results show, at least in a 3-yr period, that ecosystems can respond to changes in invader abundance, and in some cases simply removing the invader could result in long-term changes to the resident plant community.

Nomenclature: Japanese stiltgrass (Mary's-grass), Microstegium vimineum (Trin.) A. Camus.

Invasive nonnative grasses pose a significant threat to rangelands of the Northern Great Plains. Long-term data from a grazing experiment near Mandan, ND (46°46′11.43″N, 100°54′55.16″W) revealed the invasion of native prairie by Kentucky bluegrass, an exotic grass. We hypothesized that bluegrass invasion altered soil ¹³C and ¹⁵N levels, tracking the increased abundance of invasive cool-season grass aboveground. In 2014, soil samples were collected to depths of 0 to 7.6 cm and 7.6 to 15.2 cm in pastures grazed similarly since 1916. Samples were analyzed for total carbon (C) and nitrogen (N) and ¹³C and ¹⁵N isotopes and compared against archived samples from 1991. Vegetation change from native to exotic grasses changed the isotopic composition of soil C. The soil δ¹³C at the 0- to 7.6-cm depth became more negative between 1991 and 2014. Soil δ¹³C became less negative with increasing stocking rate at both soil depths. Soil δ¹⁵N values at the 0- to 7.6-cm depth decreased between 1991 and 2014. Soil δ¹⁵N increased with increasing stocking rate at the 0- to 7.6-cm depth in 2014. Soil C and N concentrations at 0 to 7.6 cm increased by 35% (12 g C kg^-1) and 27% (0.9 g N kg^-1), respectively, from 1991 to 2014; however, concentrations at the 7.6- to 15.2-cm depth did not change. The shift from native C₄ to invasive C₃ grass did not reduce soil C storage in the long-term prairie pastures. The more deleterious effect of invasion, however, may have been the buildup of dead biomass, which alters vegetation structure and may reduce native species' diversity and abundance.

Nomenclature: Kentucky bluegrass, Poa pratensis L.

Many introduced species are capable of both sexual and vegetative reproduction. Our understanding of the ecology of such species depends on the trade-offs between vegetative and sexual reproduction and the ecological conditions that favor both modes of reproduction and how those factors influence the population ecology of introduced species. Here, we studied the efficacy of propagation via both seeds and rhizomes in Johnsongrass, a widespread invasive grass whose success is due to its prolific production of shattering seeds and rhizomes, the latter of which are readily dispersed by anthropogenic and natural processes. In a common garden in Virginia, we varied the density of seeds and rhizomes and manipulated whether recruits experienced interspecific competition. Johnsongrass recruited from both seeds and rhizomes. We compared the efficacy of seeds and rhizomes on a per propagule basis and by standardizing them according to their total carbon content. Rhizomes were more efficient than seeds on a per propagule basis, but seeds propagated more efficiently than rhizomes on a per unit of carbon basis, establishing in nearly all plots and obtaining much greater biomass than rhizomes. We also found that rhizomes were subject to stronger negative density dependence than seeds and were more sensitive to site variation and competition. Our results suggest that, provided sufficient dispersal, a single Johnsongrass plant produces enough propagules to establish over more than a hectare, even at relatively low propagule densities. Proper understanding of both seed and vegetative propagation is crucial for understanding the ecology of this and other invasive species that utilize multiple reproductive modes.

Nomenclature: Johnsongrass; Sorghum halepense (L.) Pers. SORHA.

Feral cereal rye is an aggressive, persistent winter annual grass. Although feral rye has been documented as a weed in Utah cropland for many years, it has only recently been described as a weed of natural areas in Utah. After feral rye was observed on hillside locations where it had not previously been present, research was conducted to evaluate expansion rates in isolated patches and on a landscape scale. Individual patch measurements indicated expansion rates of 17%, 42%, 44%, and 112% in 2009. The landscape expansion rates were 1%, 4%, 8%, 21%, and 50% in the same year. The spread of feral rye appears to have occurred primarily on south- to west-facing slopes where the density and diversity of native species is limited. The expansion of feral rye into natural, undisturbed areas indicates that this species should be closely monitored. The relatively short seed longevity and current small infestations make it a good candidate for early detection/rapid response efforts.

Nomenclature: Cereal rye, Secale cereale L, SECE.

In this case study, we evaluated a point-mapping method for simultaneously collecting data while controlling three invasive woody plant species: black locust, Chinese privet, and hardy orange. The study in Arkansas Post National Memorial included seven project areas ranging in size from 2.7 to 27.3 ha and spanned six field seasons (2010 to 2015). The control techniques varied depending on plant size and always included the application of herbicide, which also varied over the course of the study to include glyphosate, imazapyr, and triclopyr. Each person responsible for controlling plants simultaneously collected global positioning system point data to estimate the foliar cover of the plants treated. The resulting data demonstrated evidence of decreases in all three plant species in most project areas during the 6-yr period. Initial increases in area treated for some species—area combinations reflected differences in the preliminary efforts required to control invasive plants in entire project areas, but by 2012 six of seven project areas were treated in their entirety. Despite a high level of reduction, in some cases, the plants persisted at low levels even during the sixth year of the project. Our findings support the ability of this method to granularly detect changes in plant abundance while simultaneously controlling invasive plants. With several acknowledged limitations, this streamlined project-based monitoring approach provides data that allow managers to assess the effectiveness of weed control treatments.

Nomenclature: Glyphosate; imazapyr; triclopyr; black locust, Robinia pseudoacacia L.; Chinese privet, Ligustrum sinense Lour.; hardy orange, Poncirus trifoliata (L.) Raf.

Severe leaf blight of Japanese stiltgrass (JSG) from Bipolaris disease, causing significant decline in population density at some locations, has been reported sporadically in the field. Even so, much of the JSG in the mid-Atlantic is not diseased. Six populations of JSG from the field, one that was severely diseased by B. microstegii and the others “healthy,” were tested by artificial inoculation for susceptibility to both B. microstegii (five isolates) and B. drechsleri (three isolates). Populations of JSG in this study differed in their response to the two Bipolaris species, but within species of Bipolaris the plant responses were consistent. Plants from the diseased population of JSG from Frederick, MD, were very susceptible to B. microstegii, and plants from other populations from Maryland (three locations), Delaware, and Indiana were not. In contrast, B. drechsleri caused moderate disease on plants from all accessions but one, and it was significantly less aggressive than was B. microstegii on the susceptible accession of JSG. Results of a limited host range determination only with B. microstegii revealed hypersensitive responses, and therefore high levels of resistance, in corn (four cultivars) and sorghum (three accessions). The native, sympatric grass deertongue was not diseased in these tests. Results reveal a distinct differential response among populations of JSG to disease from B. microstegii, while in contrast, B. drechsleri is capable of causing disease on a broader range of JSG populations.

Nomenclature: Bipolaris microstegii Minnis, Rossman, Kleczewski & S. L. Flory; Bipolaris drechsleri Minnis & Manamgoda; Deertongue, Dichanthelium clandestinum (L.) Gould; Mary's-grass, Japanese stiltgrass, Microstegium vimineum (Trin.) A. Camus var. imberbe (Nees) Honda MCGVM; corn, Zea mays L. ZEAMX; sorghum, Sorghum bicolor (L.) Moench subsp. bicolor SORVU.

The giant reed, Arundo donax is one of the worst invasive alien species globally, including South Africa, where it invades riparian areas across the country. Biological control is being considered to address the invasive potential and negative impacts of the weed. This study investigated the phylogeography of A. donax to guide the biological control program. To determine plant haplotype and genetic diversity, three regions of the chloroplast were sequenced and three microsatellite markers were analyzed in 40 samples from across the plant's distribution in South Africa. It was determined that all populations of A. donax in South Africa were haplotype M1, which is the most widely distributed haplotype worldwide, believed to originate from the Indus Valley, Asia. In addition, no genetic diversity was found, indicating that all the A. donax populations in South Africa are essentially one clone. The results indicate that suitable biological control agents are likely to be found in the ancient native range of haplotype M1. This research has contributed to the global understanding of the phylogeography of A. donax and will guide the biological control program in South Africa.

Nomenclature: Giant reed; Arundo donax L. ABKDO.

Although an increasing number of investigations have been made into the evolution of alien species once introduced, few studies have identified the invasion routes of these introduced species. Because multiple introductions are common in invasive species, failing to take into account the introduced lineages can be misleading when studying evolutionary change in alien species after they begin to extend their ranges. In Japan, diverse lineages of ryegrasses (Lolium spp.) were introduced as forage crops and contaminants in trading grain and have expanded to sandy coasts. We studied the expansion route of populations established along the coasts of three geographic regions within Japan by comparing variations in morphology and nuclear microsatellite and chloroplast DNA in the two habitats where ryegrasses were first introduced: croplands and international seaports. Chloroplast DNA haplotypes did not differ significantly among habitats and regions, but the coastal and seaport populations displayed similar microsatellite genetic compositions and morphological characteristics. Our results revealed that coastal populations originated from seaport populations derived from contaminants. Selective forces from the past, including domestication and naturalization, may have assisted the introduced lineages in colonizing new habitats.

Nomenclature: Italian ryegrass, Lolium multiflorum L.; rigid ryegrass, Lolium rigidum Gaudin.

Woolly distaff thistle is a long-lived winter annual that threatens the ranching and dairy industries within the North Coast counties of California, particularly the organic producers. No peer-reviewed publications have documented effective control options or integrated management approaches for this species. We conducted two experiments, each replicated, in Marin County, California. The first compared several conventional herbicides at two timings and rates, while the second compared a conventional herbicide treatment with organic and integrated organic control methods, including an organic herbicide (mixture of capric and caprylic acids). Results of the conventional herbicide treatments showed most spring applications (March or April) of aminopyralid, aminocyclopyrachlor, clopyralid, and combinations of aminopyralid triclopyr, or aminocyclopyrachlor chlorsulfuron had greater than 99% control of woolly distaff thistle with fewer than 1.5 seedlings per 27-m² plot by the end of the growing season. Higher rates were generally necessary to achieve the same level of control with winter (January) applications. In the organic herbicide treatments, the most consistent treatment was a combination of mowing followed by 9% (v/v) or the organic herbicide. This treatment was slightly less effective compared with aminopyralid but did have better than 95% control of woolly distaff thistle. The results of this study provide control options for both conventional and organic ranching practices where woolly distaff thistle is a problem.

Nomenclature: Aminopyralid; aminocyclopyrachlor; capric acid; caprylic acid; chlorsulfuron; clopyralid; triclopyr; woolly distaff thistle, Carthamus lanatus L.

Kentucky bluegrass (Poa pratensis L.) invades northern Great Plains rangelands. On the Sheyenne National Grassland in southeastern North Dakota, three research sites, each with a different level of Kentucky bluegrass invasion, were chosen to evaluate effectiveness of burning and burning—herbicide combinations to control Kentucky bluegrass. Initial Kentucky bluegrass invasion levels were 37%, 77%, and 91% for LOW, MODERATE, and HIGH invaded sites, respectively. Within each invaded site, four replicated strips (20 by 60 m) were established, with half of each strip burned in late October 2005 and the other half burned in early May 2006. Herbicide treatments of (1) no herbicide, (2) 2.24 kg ha^-1 of glyphosate, and (3) 0.43 kg ha^-1 of imazapic were randomly assigned to 10 by 20 m subplots within each burn. Control plots were established at the same time. Relative basal cover of native grass, native forb, and Kentucky bluegrass was estimated annually using 50 10-point frames within each subplot. On the HIGH site in 2006, fall-burned plots with a spring glyphosate application had three times the native grass cover and only one fourth of the Kentucky bluegrass cover compared with controls. Similar results with the same treatment occurred at the MODERATE site. Native grasses became the most abundant plant community on these plots in the MODERATE and HIGH sites within 1 yr. Treatment differences were transitory, and the LOW site differed from the MODERATE and HIGH sites. In 2007, on fall-burned plots with spring glyphosate application, the amount of Kentucky bluegrass was 14% and 30%, and native grass species were 52% and 42% on the MODERATE and HIGH sites, respectively, which was similar to the initial values on the LOW site. These data emphasize the importance of initial invasion level in developing restoration strategies and provide evidence burning and herbicide combinations can be valuable management tools even on heavily invaded grasslands.

Nomenclature: Glyphosate; imazapic; Kentucky bluegrass, Poa pratensis L. POAPR.

Interest exists in planting mixed forb—grass prairies in the midwestern United States. Aminopyralid or clopyralid can be used to suppress competition from invasive plants prior to seeding prairies. As these active ingredients are known to persist, concern exists that reductions in forb establishment could occur. We tested whether common midwestern forb species could tolerate an application of aminopyralid or clopyralid alone or in combination the summer prior to seeding, and whether fall dormant or spring seeding date influenced establishment. This experiment was performed in Beresford, SD, and Arlington, WI, where aminopyralid (54 or 123 g ae ha^-1), clopyralid (237 and 420 g ae ha^-1), or aminopyralid clopyralid (54 237 g ae ha^-1) were applied to a prepared seedbed in July of 2009. Ten forbs were seeded in November 2009 as a dormant seeding and in April 2010 as a spring seeding at both locations, and establishment was assessed 12 and 24 mo after treatment (MAT). Results were site and species specific. Time of seeding was an important driver of plant counts at both locations 12 and 24 MAT. In Wisconsin at 12 MAT, 60% of species studied exhibited higher counts in the spring seeding. This trend persisted in some, but was not consistent across all 10 species. In South Dakota, 80% of species studied had higher counts at 12 and 24 MAT, but differences were species specific and often differed from those studied in Wisconsin. Those species that had higher counts in spring seeding at 12 MAT, maintained higher counts at 24 MAT. Forbs planted in plots treated with herbicides did not differ from plots left untreated at either location. Results suggest native forbs typically seeded in the upper Midwest can tolerate these herbicides when applied at least 4 mo prior to seeding.

Nomenclature: Aminopyralid; clopyralid.

There are an estimated 400 million hectares of non-cropland in the United States primarily designated as rangeland and pastureland, and there are more than 300 invasive weeds found on these sites, causing an estimated annual loss of $5 billion. Among the most invasive and problematic weeds are Dalmatian toadflax, diffuse knapweed, downy brome, and musk thistle. Currently, herbicides are the most common management strategy for broadleaf weeds and invasive winter annual grasses. Indaziflam, a new herbicide for invasive plant management in non-crop areas, is a cellulose-biosynthesis inhibitor capable of providing residual invasive winter annual grass control up to 3 yr after treatment (YAT). A field experiment was conducted to determine whether residual Dalmatian toadflax and downy brome control by aminocyclopyrachlor, imazapic, and picloram could be extended by tank mixing these herbicides with indaziflam. Indaziflam tank mixed with aminocyclopyrachlor, imazapic, and picloram provided increased Dalmatian toadflax (84% to 91%) and downy brome (89% to 94%) control 4 YAT, compared with treatments excluding indaziflam. Treatments without indaziflam controlled 50% to 68% of Dalmatian toadflax and <25% downy brome 4 YAT. Based on these results, a greenhouse dose-response experiment was conducted with aminocyclopyrachlor, aminopyralid, and indaziflam to compare preemergence control of nine common non-crop weeds. Averaged across species, aminocyclopyrachlor and aminopyralid GR₅₀ values (herbicide concentration resulting in 50% reduction in plant biomass) were 29 and 52 times higher compared with indaziflam, respectively. These data suggest that indaziflam could be used for residual control of non-crop weeds as a tank-mix partner with other foliar-applied broadleaf herbicides.

Nomenclature: Aminocyclopyrachlor; aminopyralid; imazapic; indaziflam; picloram; Dalmatian toadflax, Linaria dalmatica (L.) P. Mill.; diffuse knapweed, Centaurea diffusa Lam.; downy brome, Bromus tectorum L.; musk thistle, Carduus nutans L.

Invasive species management is often more successful if desirable species are seeded after the target weed is controlled. However, control of invasive plants must be maintained following reseeding or the seeded species may fail to establish. A regional study conducted in Minnesota, North Dakota, and South Dakota evaluated the effect of aminopyralid, clopyralid, or picloram applied in the fall prior to fall-dormant seeding or seeding the following spring on cool- and warm-season native grass species establishment. Herbicides were applied at standard rates used to control invasive broadleaf weeds in the upper midwestern tallgrass prairie region of the United States. Cool-season species included Canada wildrye, green needlegrass, and intermediate wheatgrass. Warm-season species included big bluestem, little bluestem, sideoats grama, switchgrass, and Indiangrass. Aminopyralid did not reduce seedling establishment in either fall or spring seeding. Grasses generally were not affected by a pretreatment of the pyridine standards clopyralid or picloram either, with the exception of a slight reduction in fall-seeded establishment of intermediate wheatgrass. Picloram also slightly reduced fall-seeded establishment of Canada wildrye. Application of aminopyralid can safely be used to control susceptible invasive species preceding grass species establishment, with a safety margin similar to or slightly better than that with the pyridine standards clopyralid or picloram.

Nomenclature: Aminopyralid; clopyralid; picloram; big bluestem (Andropogon gerardii Vitman); Canada wildrye (Elymus canadensis L.); green needlegrass [Nassella viridula (Trin.) Barkworth]; indiangrass [Sorghastrum nutans (L.) Nash]; intermediate wheatgrass [Thinopyrum intermedium (Host) Barkworth & D. R. Dewey]; little bluestem [Schizachyrium scoparium (Michx.) Nash]; sideoats grama [Bouteloua curtipendula (Michx.) Torr.]; switchgrass (Panicum virgatum L.).