Can Carbon and Phosphorous Amendments Increase Native Forbs in a Restoration Process? A Case Study in the Northern Tall‐grass Prairie (U.S.A.)

In the northern Great Plains (United States), sites with less than 20% of native species are difficult to restore. We have experimented with a restoration method that shows some promise. It consists of systematically installing simulated small‐scale patches (8.0 m² in size) over 25% of an old field and then seeding these patches with native species. The working hypothesis is that these patches will generate a constant source of propagules which in time will lead to increases in native species diversity within the surrounding grass matrix. The objective of this paper was to determine whether soil amendments should be used to facilitate the establishment and persistence of native species (primarily forbs) within these patches. We seeded the patches with a mixture of native grass and forb species and applied four soil treatments: P fertilization, C additions, C + P, and a control (no amendments). Results for the first 5 years were as follows: (1) seeded forb richness was mostly unaffected by soil amendments; (2) seeded and nonseeded forb biomass and density were substantially reduced by C additions, whereas they were unaffected or increased under P additions; (3) both seeded and non‐native grass biomass substantially increased with C additions; and (4) there was an inverse relationship between native seeded forbs and non‐native grass biomass. Our conclusions are that: (1) P amendments are a potential tool for enhancing native seeded forb biomass in simulated small‐scale disturbance patches; and (2) C additions, although enhancing seeded grass biomass do not reduce the biomass of non‐native grasses.     

Sowing density effects and patterns of colonization in a prairie restoration

A cost‐effective approach in plant restorations could be to increase sowing density for species known to be challenging to establish, while reducing sowing density for species that easily colonize on their own. Sowing need not occur evenly across the site for rapidly dispersing species. We explored these issues using a prairie restoration experiment on a high‐school campus with three treatments: plots sown only to grasses (G plots), to grasses and forbs (GF1), and to grasses and forbs with forbs sown at twice the density (GF2). In year 2, GF1 and GF2 plots had higher diversity than G plots, as expected, but GF2 treatments did not have twice the sown forb cover. However, high forb sowing density increased forb richness, probably by reducing stochastic factors in establishment. Cover of nonsown species was highest in G plots and lowest in GF2 plots, suggesting suppressive effects of native forbs on weedy species. Colonization of G plots by two sown forbs (Coreopsis tinctoria and Rudbeckia hirta) was apparent after 2.5 years, providing evidence that these species are self‐sustaining. Colonization was greater in edges than in the central areas of G plots. Through construction of establishment kernels, we infer that the mean establishment distance was shorter for R. hirta (6.7 m) compared to C. tinctoria (21.1 m). Our results lead us to advocate for restoration practices that consider not only seed sowing but also subsequent dispersal of sown species. Furthermore, we conclude that restoration research is particularly amenable for outdoor education and university‐high school collaborations.     

Effects of Frequent Mowing on Survival and Persistence of Forbs Seeded into a Species-Poor Grassland

Many early attempts at tallgrass prairie reconstruction failed to achieve the high species diversity of remnant prairies, and instead consist primarily of C₄ grasses. We hypothesized that frequent mowing of established prairie grasses could create sufficient gaps in the aboveground and belowground environment to allow for the establishment of native forbs from seed. We studied forb seedling establishment in a 25‐year‐old prairie planting in northern Iowa that was dominated by native warm‐season grasses. In winter 1999, 23 species of native forbs were broadcast into the recently burned sod at a rate of 350 viable seeds/m². Treatment plots were mowed weekly for either one or two growing seasons, and control plots were unmowed. Mowed plots had greater light availability than controls, especially when warm‐season grasses began to flower. Overwinter seedling mortality was 3% in mowed treatments compared to 29% in the controls. Forbs in mowed plots had significantly greater root and shoot mass than those in control plots in the first and second growing seasons but were not significantly more abundant. By the fourth growing season, however, forbs were twice as abundant in the mowed treatments. No lasting negative impacts of frequent mowing on the grass population were observed. Mowing a second year influenced species composition but did not change total seedling establishment. Experimental evidence is consistent with the idea that mowing reduced competition for light from large established grasses, allowing forb seedlings the opportunity to reach sufficient size to establish, survive, and flower in the second and subsequent years.     

Long‐term experiment manipulating community assembly results in favorable restoration outcomes for invaded prairies

Invasive species are a common problem in restoration projects. Manipulating soil fertility and species arrival order has the potential to lower their abundance and achieve higher abundances of seeded native species. In a 7‐year experiment in Missouri, United States, we tested how nutrient addition and the timing of arrival of the invasive legume Lespedeza cuneata and seeded native prairie grass and forb species influenced overall community composition. Treatments that involved early arrival of seeded forb and grass species and late arrival of L. cuneata were most successful at creating community structure that fulfilled our restoration goals, displaying high abundance of seeded native forb species, low abundances of L. cuneata, and non‐native species. There were few treatment interactions, with the exception that timing seeded native forbs and timing of L. cuneata arrival interactively influenced the abundance of seeded native forbs. This suggests that the individual treatments are supporting the restoration goals, such as creating a community with low abundance of L. cuneate or high abundance of native seeded species, without restricting each other. This study demonstrates the importance of priority effects in disturbed habitats prone to invasion, the lasting effects of initial seeding on long‐term community composition, and the potential for fertilization to positively benefit restoration of degraded grasslands.     

Mycorrhizal suppression alters plant productivity and forb establishment in a grass-dominated prairie restoration

A fundamental goal of restoration is the re-establishment of plant diversity representative of native vegetation. However, many prairie restorations or Conservation Reserve Program sites have been seeded with warm-season grasses, leading to grass-dominated, low-diversity restorations not representative of native grasslands. These dominant grasses are strongly mycotrophic, while many subordinate forb species appear to be less dependent on mycorrhizal symbiosis. Therefore, manipulating arbuscular mycorrhizal fungi (AMF) may be useful in promoting establishment and growth of forb species in grass-dominated prairie restorations. To assess the potential role of mycorrhizae in affecting the productivity and community composition of restored tallgrass prairie, we conducted a 4-year field experiment on an 8-year-old grassland restoration at the Konza Prairie in northeastern Kansas, USA. At the initiation of our study, seeds of 12 forb species varying in degree of mycorrhizal dependence were added to established grass-dominated plots. Replicate plots were treated bi-weekly with a soil drench of fungicide (Topsin-M^®) over four growing seasons and compared to non-treated control plots to assess the role of AMF in affecting plant species composition, productivity, leaf tissue quality, and diversity in restored tallgrass prairie. Topsin applications successfully reduced mycorrhizal colonization of grass roots to approximately 60–80% relative to roots in control plots. Four years of mycorrhizal suppression reduced productivity of the dominant grasses and increased plant species richness and diversity. These results highlight the importance of mycorrhizae as mediators of plant productivity and community dynamics in restored tallgrass prairie and indicate that temporarily suppressing AMF decreases productivity of the dominant C4 grasses and allows for establishment of seeded forb species.     

Are two strategies better than one? Manipulation of seed density and soil community in an experimental prairie restoration

Restoration practitioners have a variety of practices to choose from when designing a restoration, and different strategies may address different goals. Knowledge of how to best use multiple strategies could improve restoration outcomes. Here, we examine two commonly suggested strategies in a single tallgrass prairie restoration experiment: increased forb sowing density and prairie soil inoculation. We designed a study with two different forb seeding densities. Within these densities, we transplanted seedlings into 1‐m² plots that had been grown in either a whole prairie soil inoculum or sterilized prairie soil. After 4 years, we found positive effects of both high forb sowing density and inoculation treatments on the ratio of seeded to nonseeded plant cover in these plots, and negative effects of both treatments on nonseeded plant diversity. No effects of either treatment were seen on seeded plant diversity. Each strategy also affected the plant community in different ways: high forb sowing density increased seeded forb richness and decreased native nonseeded plant cover, while inoculation decreased non‐native cover, and tended to increase average successional stage of the community. These effects on restoration outcome were typically independent of each other, with the result that plots with both manipulations had the most positive effects on restoration outcomes. We thus advocate the combined use of these restoration strategies, and further studies which focus on both seeding and soil community manipulation in restoration.     

Enhancing Native Forb Establishment and Persistence Using a Rich Seed Mixture

Establishing native forbs is crucial for invasive plant management and restoring a desirable plant community. Our objectives were to determine (1) if increasing forb seed density results in increased forb establishment; (2) if a species‐rich mixture of forbs has greater establishment and survivorship than a single species; and (3) if mixtures of forbs are more competitive with Spotted knapweed (Centaurea maculosa) than a forb monoculture. To test our first two objectives, we seeded monocultures of Purple coneflower (Echinacea angustifolia), Arrowleaf balsamroot (Balsamorhiza sagittata), Annual sunflower (Helianthus annuus), Dotted gayfeather (Liatris punctata), Western white yarrow (Achillea millefolium), Sticky geranium (Geranium viscosissimum), as well as a mixture of all forbs. Pots were seeded at 800 or 2,000 seeds/m² and watered twice or thrice weekly. The highest seed density produced the highest plant density, which averaged 4.35 plants/pot. The density of the mixture was similar to the mean density seen for individual species, and it doubled in response to the highest seed density. To test our third objective, Spotted knapweed and Purple coneflower were arranged in an addition series matrix with a maximum total density of 4,000 seeds/pot. We found that the forb mixture was seven times more competitive with Spotted knapweed than Purple coneflower alone. Using a mixture of forbs rather than a single species enhances forb establishment in various and unpredictable environments because the mixture possesses a variety of traits that may match year–year and site–site conditions. Once established, the mixture may have a greater chance of persisting than a monoculture.     

Responses of native and non-native Mojave Desert winter annuals to soil disturbance and water additions

Habitat modification (i.e., disturbance) and resource availability have been identified as possible mechanisms that may influence the invasibility of plant communities. In the Mojave Desert, habitat disturbance has increased dramatically over the last 50 years due to increased human activities. Additionally, water availability is considered to be a main limiting resource for plant production. To elucidate the effects of soil disturbance and water availability on plant invasions, we created experimental patches where we varied the levels of soil disturbance and water availability in a fully crossed factorial experiment at five replicated field sites, and documented responses of native and non-native winter annuals. The treatments did not significantly affect the density (seedlings m⁻²) of the non-native forb, Brassica tournefortii. However, the relationship between silique production and plant height differed among treatments, with greater silique production in disturbed plots. In contrast to Brassica, density of the non-native Schismus spp. increased in soil disturbed and watered plots, and was greatest in disturbed plots during 2009 (the second year of the study). Species composition of the native annual community was not affected by treatments in 2008 but was influenced by treatments in 2009. The native forb Eriophyllum sp. was most dense on water-addition plots, while density of Chaenactis freemontii was highest in disturbed plots. Results illustrate that habitat invasibility in arid systems can be influenced by dynamics in disturbance regimes and water availability, and suggest that invasiveness can differ between non-native annual species and among native annuals in habitats undergoing changing disturbance and precipitation regimes. Understanding the mechanistic relationships between water availability and non-native plant responses will be important for understanding the effects of shifting precipitation and vegetation patterns under predicted climate change in arid ecosystems.     

Establishing Tallgrass Prairie on Grazed Permanent Pasture in the Upper Midwest

The goal of the study was to learn whether native prairie grasses and, eventually, a diverse mixture of native forbs could be incorporated in permanent pastures by means of rotational grazing by cattle. An experiment was established on a farm in northeastern Iowa on a pasture that had never been plowed but had been grazed since the 1880s. One treatment was protected from grazing to test for the presence of remnant vegetation. Andropogon gerardii, Sorghastrum nutans, Panicum virgatum, and Desmanthus illinoensis were introduced in plots first treated with glyphosate; seeds were either drilled (DR) or hand-broadcast and incorporated by controlled cattle trampling (BT). Seedling establishment and aboveground biomass were followed over 3 years. There was no evidence for remnant native plants on uplands, but seven species of native forbs and four native graminoids flowered in exclosures erected within waterways. D. illinoensis initially established up to 12 seedlings/m² but had disappeared from all but one plot by the third year. Variation in native grass establishment among replicate plots within treatments was very high, ranging initially from 0.2 to 9.9 plants/m². In August of the second year, native grasses made up only 8% of the available forage in DR plots and 1% of BT plots. One year later, however, native grasses made up 56% of the available forage in DR plots and 37% of BT plots, and these differences were significant (p = 0.05). A pilot study seeded in late winter (frost seeding) suggested that seeds spread after cattle trampling produced five times more seedlings (2.5/m²) than seeds spread before cattle trampling (0.5/m²). Frost seeding had advantages because it did not require herbicide for sod suppression or tractor access to the site. New plantings could be safely grazed in early spring and late fall, before and after most native grass growth, to offset the negative economic impact of protecting new plantings from burning during the growing season. But this practice precluded subsequent prescribed burning. I propose a strategy for incorporating native wildflowers into the pasture over time with minimum cost.     

Interspecific variation in plant responses to mycorrhizal colonization in tallgrass prairie

Symbiotic associations between plants and arbuscular mycorrhizal fungi are ubiquitous and ecologically important in many grasslands. Differences in species responses to mycorrhizal colonization can have a significant influence on plant community structure. The growth responses of 36 species of warm- and cool-season tallgrass prairie grasses and 59 tallgrass prairie forbs to arbuscular mycorrhizal (AM) fungal colonization were assessed in greenhouse studies to examine the extent of interspecific variation in host-plant benefit from the symbiosis and patterns of mycorrhizal dependence among host plant life history (e.g., annual, perennial) and taxonomic (e.g., grass, forb, legume, nonlegume) groups and phenological guilds. There was a strong and significant relationship between phenology of prairie grasses and mycorrhizal responsiveness, however this relationship was less apparent in forbs. Perennial warm-season C(4) grasses and forbs generally benefited significantly from the mycorrhizal symbiosis, whereas biomass production of the cool-season C(3) grasses was not affected. The root systems of the cool-season grasses were also less highly colonized by the AM fungi, as compared to the warm-season grasses or forbs. Unlike the native perennials, annuals were generally not responsive to mycorrhizal colonization and were lower in percentage root colonization than the perennial species. Plant growth responsiveness and AM root colonization were positively correlated for the nonleguminous species, with this relationship being strongest for the cool-season grasses. In contrast, root colonization of prairie legumes showed a significant, but negative, relationship to mycorrhizal growth responsiveness.     

Effects of Nutrient Manipulations and Grass Removal on Cover,Species Composition,and Invasibility of a Novel Grassland in Colorado

Cover and richness of a 5‐year revegetation effort were studied with ,respect to small‐scale disturbance and nutrient manipulations. The site, originally a relict tallgrass prairie mined for gravel, was replanted to native grasses using a seed mixture of tall‐, mixed‐, and short‐grass species. Following one wet and three relatively dry years, a community emerged, dominated by species common in saline soils not found along the Colorado Front Range. A single species, Alkali sacaton (Sporobolus airoides), composed nearly 50% of relative vegetation cover in control plots exhibiting a negative relationship between cover and richness. Seeded species composed approximately 92% of vegetation cover. The remaining 8% was composed of weeds from nearby areas, seed bank survivors, or mix contaminants. Three years of soil nutrient amendments, which lowered plant‐available nitrogen and phosphorus, significantly increased relative cover of seeded species to 97.5%. Fertilizer additions of phosphate enhanced abundance of introduced annual grasses (Bromus spp.) but did not significantly alter cover in control plots. Unmanipulated 4‐m² plots contained an average of 4.7 planted species and 3.9 nonplanted species during the 5‐year period, whereas plots that received grass herbicide averaged 5.4 nonplanted species. Species richness ranged from an average 6.9 species in low‐nutrient, undisturbed plots to 10.9 species in the relatively high‐nutrient, disturbed plots. The use of stockpiled soils, applied sparingly, in conjunction with a native seed mix containing species uncommon to the preexisting community generated a species‐depauperate, novel plant community that appears resistant to invasion by ruderal species.     

Effects of Established Perennial Grasses on Introduction of Native Forbs in California

Prairie restoration is not complete without the establishment of both grasses and forbs. However, if desirable forbs and grasses are seeded simultaneously, control of broadleaf weeds is problematic. If possible, a two‐step process of introducing forbs after establishing grasses would allow use of broadleaf‐specific herbicides at the critical early stages of grass growth. We conducted experiments to investigate methods for introducing forbs into previously restored native perennial grasslands on rural roadsides in the Sacramento Valley of California. In one experiment, we studied the effects of background vegetation (established perennial grasses or tilled ground) on seven native forb species planted from seed. In a second experiment, we evaluated the effects of background vegetation (existing perennial grasses or tilled ground) and container size (36 ml or 105 ml) with excavation technique (excavation by core removal [core] or by creating an impression [dibble]) on the growth of transplants of the native perennial forbs Asclepias fascicularis and Sisyrinchium bellum. The presence of established perennial grasses reduced the growth of seeded forbs, but did not affect transplants, indicating the vulnerability of seedling forbs to interference. When compared to control plots that had been tilled in the autumn, weed canopy cover was significantly lower in the presence of perennial grasses if seeded with forbs, but not in the presence of perennial grasses alone. Both transplanted species grew better in the large container/core treatment than the small container/dibble treatment; however, existing grasses eliminated these positive effects. Asclepias fascicularis performed better when grown in large containers than in small containers, but its growth was not affected by excavation method; S. bellum performed better when planted with the core method than the dibble method of excavation, but container size made no difference. We attribute differences in the responses of the species to interactions between phenological differences and expansive clay soils that naturally de‐compact upon drying.     

Plant responses to selective grazing by bison: interactions between light,herbivory and water stress

Two abundant tallgrass prairie forb species, Ambrosia psilostachya and Vernonia baldwinii, are commonly found intact in patches where the grasses have been selectively grazed by bison. Microclimatic patterns and physiological responses of these forbs were measured in grazed and ungrazed patches. These experiments demonstrated that bison herbivory indirectly enhanced water availability and productivity of forbs growing in grazed patches. This was due primarily to the reduction in transpiring grass leaf area in grazed patches and an increase in light availability. In grazed patches, incident light at forb mid-canopy height was 53% greater than ungrazed sites at midseason and soil temperatures were always warmer (e.g., 10°C at 5 cm), perhaps enabling forbs to initiate growth earlier in the spring. Enhanced leaf xylem pressure potential and stomatal conductance in plants in grazed areas were most evident when water availability was low (i.e., late in the growing season and over short-term dry periods characteristic of the tallgrass prairie environment). Relative to individuals in ungrazed areas, end-of-season biomass of A. psilostachya was 40% greater and reproductive biomass and head number of V. baldwinii was 45% and 40% greater, respectively, in plants in grazed patches. A favorable growing environment maintained in grazed patches during periods of water limitation enhances carbon gain in forbs leading to increased biomass and potential fitness.     

Forb Species Establishment Increases with Decreased Grass Seeding Density and with Increased Forb Seeding Density in a Northeast Kansas, U.S.A., Experimental Prairie Restoration

Most prairie restorations fail to produce the diversity of species found in unplowed remnants. This lack of restored diversity is hypothesized to be partly due to the inhibition of forb species by high seeding densities of dominant grasses and partly due to the low seeding densities of forbs used in many restorations. We tested this hypothesis by sowing various densities of forb and warm-season grass seeds into a restoration begun on bare soil. This is the first replicated restoration experiment we are aware of that varies grass seeding densities to examine the effects on forbs. Four years after seeding, we found that higher densities of grass seeds decreased forb cover, biomass, and richness, and higher densities of forb seeds increased forb richness. These results suggest that dominant grasses compete strongly with native forb species and that many forb species thrive when they are spatially separated from dominant grasses. The results also suggest that seed availability limits the establishment of some forbs. Forb diversity can therefore be increased by decreasing grass seeding density, by increasing forb seeding density, or both. However, forb seeds are generally expensive, and increasing forb seeding density across the entire area of a restoration may be prohibitively expensive. We therefore recommend a low seeding density of dominant grasses, and we recommend spatially separating forbs from dominant grasses by adding most forb seeds to areas with little to no dominant grasses and by adding the rest of the forb seeds to areas with a low density of dominant grasses.     

Roadside Revegetation in Glacier National Park, U.S.A.: Effects of Herbicide and Seeding Treatments

From 1992 to 1995 we experimentally evaluated the effectiveness of several revegetation treatments along a segment of Going-to-the-Sun Highway in Glacier National Park, U.S.A. This segment, reconstructed during the spring and summer of 1992, is bordered by fescue prairie vegetation and is known to be susceptible to invasion by several alien species, including Centaurea maculosa (spotted knapweed) and Phleum pratense (common timothy). We used a split plot study design to evaluate the effectiveness of herbicide and seeding treatments on assisting recovery of native flora and limiting the establishment of alien species. The herbicide treatment consisted of a yearly herbicide spray application of clopyralid (3,6-dichloropicolinic acid). Five seeding treatments were evaluated, three of which included an indigenous graminoid-forb seed mix. Percent canopy coverages of four species groups—alien graminoids, native graminoids, alien forbs, and native forbs—were determined in July 1995. In addition, community-level patterns in sprayed plots and unsprayed plots were compared with a reference site of native fescue prairie. Herbicide treatments decreased mean canopy coverage of alien forbs (treated = 4.2%, untreated = 23.4%) and increased mean canopy coverage of native graminoids slightly (treated = 6.3%, untreated = 4.0%). But herbicide treatments reduced mean coverage of native forbs (treated = 3.9%, untreated = 8.9%) and likely increased coverage of alien graminoids. Treatments that included a fall 1992 seed mix increased native graminoid coverages 2.8–4.6 times, although coverages were still lower than those attained by alien graminoids. Native and alien forb coverage appeared unaffected by seeding treatments. Species composition was less diverse in sprayed plots and more dominated by alien grasses than in unsprayed plots and the reference site. Areas for additional study are suggested, including seed bank assays to determine treatment effects on recruitment of alien versus native species and the use of native graminoids to create low-diversity communities with high canopy coverages to resist establishment of alien species.     

Disentangling direct and indirect effects of a legume shrub on its understorey community

Direct and indirect interactions among plants contribute to shape community composition through above‐ and belowground processes. However, we have not disentangled yet the direct and indirect soil and canopy effects of dominants on understorey species. We addressed this issue in a semi‐arid system from southeast Spain dominated by the legume shrub Retama sphaerocarpa. During a year with an exceptionally dry spring, we removed the shrub canopy to quantify aboveground effects and compared removed‐canopy plots to open plots between shrubs to quantify soil effects, both with and without watering. We added a grass removal treatment in order to separate direct from indirect shrub effects and quantified biomass, abundance, richness and composition of the forb functional group. With watering, changes in forb biomass were primarily driven by indirect shrub effects, with contrasting negative soil and positive aboveground indirect effects; changes in forb abundance and composition were more influenced by direct shrub soil effects with contrasting species composition between open and Retama patches. As community composition was different between open and Retama patches the indirect effects of Retama on forb species did not concern forbs from the open community but forbs from Retama patches. Indirect effects are, thus, important at the functional group level rather than at the species level. Without watering, there were no significant interactions. Changes in species richness between treatments were weak and seldom significant. We conclude that shrub effects on understorey forbs are primarily due to their influence on soil properties, directly affecting forb species composition but indirectly affecting the biomass of the forbs of the Retama patches, and only with sufficient water.     

Butterfly response to floral resources during early establishment at a heterogeneous prairie biomass production site in Iowa, USA

In the Midwestern USA, current biofuel production systems rely on high input monoculture crops that do little to support native biodiversity. The University of Northern Iowa??s Tallgrass Prairie Center is investigating the feasibility of cultivating and harvesting diverse mixes of native prairie vegetation for use as a sustainable biofuel in a manner that also conserves biodiversity and protects soil and water resources. In 2009, we established 48 research plots on three soil types at an Iowa site with a uniform history of row crop production. We seeded each plot with one of four treatments of native prairie vegetation: (1) switchgrass monoculture, (2) warm-season grass mix (5 grass species), (3) biomass mix (16 species of grasses, legumes, and forbs), or (4) prairie mix (32 species of grasses, legumes, forbs, and sedges). In 2010, we measured vegetation characteristics and studied butterfly use of the plots to investigate the hypothesis that more diverse plant communities would support a greater abundance and diversity of butterflies. Habitat characteristics varied significantly among the plots by treatment and soil type, and butterflies responded rapidly to variation in floral abundance and richness. Averaged over the entire growing season, butterflies were six times more abundant and twice as species rich in the biomass and prairie mix plots compared to the warm-season grass and switchgrass plots. Our results suggest that implementation of biomass production using diverse mixes of native prairie vegetation on marginal lands could have positive effects on the maintenance of butterfly populations in agricultural landscapes.     

Tallgrass prairie restoration: implications of increased atmospheric nitrogen deposition when site preparation minimizes adventive grasses

Site preparation designed to exhaust the soil seedbank of adventive species can improve the success of tallgrass prairie restoration. Despite these efforts, increased rates of atmospheric nitrogen (N) deposition over the next century could potentially promote the growth of nitrophilic, adventive species in tallgrass restoration projects. We used a field experiment to examine how N addition affected species composition and plant productivity over the first 3 years of a tallgrass prairie restoration that was preceded by the planting of glyphosate‐resistant crops and multiple applications of glyphosate to exhaust the pre‐existing seedbank. We predicted that N addition would increase the percent cover of adventive plant species not included in the original seeding. Contrary to our prediction, only the cover of native species increased with N addition; native non‐leguminous forbs increased substantially, with Conyza canadensis (a weedy native species not part of the restoration seed mix) exploiting the combination of high N and bare ground in the first year, and non‐leguminous forbs (in particular Monarda fistulosa) and native C₃ grasses, all of which were seeded, increasing with N addition by the third year. Native legumes was the only functional group that exhibited lower cover in N addition plots than in control plots. There was no significant response by native C₄ grasses to N addition, and adventive grasses remained mostly absent from the plots. Overall, our results suggest that site pre‐treatment with herbicide may continue to be effective in minimizing adventive grasses in restored tallgrass prairie, despite future increases in atmospheric N deposition.     

The roles of exotic grasses and forbs when restoring native species to highly invaded southern California annual grassland

Many semi-arid shrublands in the western US have experienced invasion by a suite of exotic grasses and forbs that have altered community structure and function. The effect of the exotic grasses in this area has been studied, but little is known about how exotic forbs influence the plant community. A 3-year experiment in southern California coastal sage scrub (CSS) now dominated by exotic grasses was done to investigate the influence of both exotic grasses (mainly Bromus spp.) and exotic forbs (mainly Erodium spp.) on a restoration seeding (9 species, including grasses, forbs, and shrubs). Experimental plots were weeded to remove one, both, or neither group of exotic species and seeded at a high rate with a mix of native species. Abundance of all species varied with precipitation levels, but seeded species established best when both groups of exotic species were removed. The removal of exotic grasses resulted in an increase in exotic and native forb cover, while removal of exotic forbs led to an increase in exotic grass cover and, at least in one year, a decrease in native forb cover. In former CSS now converted to exotic annual grassland, a competitive hierarchy between exotic grasses and forbs may prevent native forbs from more fully occupying the habitat when either group of exotics is removed. This apparent competitive hierarchy may interact with yearly variation in precipitation levels to limit restoration seedings of CSS/exotic grassland communities. Therefore, management of CSS and exotic grassland in southern California and similar areas must consider control of both exotic grasses and forbs when restoration is attempted.     

Fire and seedling population dynamics in western Oregon prairies

Abstract. Through an experiment in three prairie vegetation types in western Oregon, USA the effect of prescribed fire on the timing and rates of seedling emergence and mortality was examined. Seeds of common exotic and native prairie species were sown into burned and unburned plots in late September, 1995. Emerged seedlings were censussed the following winter, early spring and late spring. Results indicated that spring population levels could not be forecast by fall seedling flushes, as winter survival was important in seedling establishment. The bulk of emergence for all grass and annual forb species occurred in the fall, followed by low to severe winter mortality. Perennial forbs were more variable in emergence times but, once emerged, perennial forb seedlings were likely to become established. Burning caused a statistically significant increase in seedling accumulation through emergence and survival in 11 of 23 cases. Burning improved seedling winter survival for most grass and short‐lived forb species and increased emergence of perennial forb species. These patterns were most conspicuous on the two sites dominated by exotic species, where burning significantly improved the accumulation of seedlings from most native species tested. Thus, prescribed burning might be a useful restoration tool in these communities. In contrast, two of the three species increased by burning in the native bunchgrass site were exotic pest plants, suggesting that fire should be prescribed with caution.