Mowing Reduces Exotic Annual Grasses but Increases Exotic Forbs in a Semiarid Grassland

Cheatgrass (Bromus tectorum) and other exotic winter‐active plants can be persistent invaders in native grasslands, growing earlier in the spring than native plants and pre‐empting soil resources. Effective management strategies are needed to reduce their abundance while encouraging the reestablishment of desirable native plants. In this 4‐year study, we investigated whether mowing and seeding with native perennial grasses could limit growth of exotic winter‐actives, and benefit growth of native plants in an invaded grassland in Colorado, United States. We established a split‐plot experiment in October 2008 with 3 mowing treatments: control, spring‐mowed, and spring/summer‐mowed (late spring, mid‐summer, and late summer), and 3 within‐plot seeding treatments: control, added B. tectorum seeds, and added native grass seeds. Cover of plant species and aboveground biomass were measured for 3 years. In March and June of 2010, 2011, and March of 2012, B. tectorum and other winter‐annual grasses were half as abundant in both mowing treatments as in control plots; however, cover of non‐native winter‐active forbs increased 2‐fold in spring‐mowed plots and almost 3‐fold in spring/summer‐mowed plots relative to controls. These patterns remained consistent 1 year after termination of treatments. Native cool‐season grasses were most abundant in spring‐mowed plots, and least abundant in control plots. There was higher cover of native warm‐season grasses in spring/summer‐mowed plots than in control plots in July 2011 and 2012. The timing of management can have strong effects on plant community dynamics in grasslands, and this experiment indicates that adaptive management can target the temporal niche of undesirable invasive species.     

Plant demographic responses to mycorrhizal symbiosis in tallgrass prairie

The effects of mycorrhizal symbiosis on seedling emergence, flowering and densities of several grasses and forbs were assessed in native tallgrass prairie and in sown garden populations at the Konza Prairie in northeastern Kansas. Mycorrhizal activity was experimentally suppressed with the fungicide benomyl. Flowering and stem densities of the cool-season grass, Dichanthelium oligosanthes, sedges (Carex spp.), and the forb Aster ericoides were higher in non-mycorrhizal (benomyl-treated) than in mycorrhizal plots and the magnitude of these differences was significantly affected by burning. Mycorrhizae significantly enhanced flowering of the warmseason grasses Andropogon gerardii and Sorghastrum nutans in burned prairie, but not in unburned sites. These patterns suggest that mycorrhizal effects on the dynamics of cool-season graminoid and forb populations are likely to be mediated indirectly through effects of the symbiosis on the competitive dominance of their neighbors. Seedling emergence rates of the cool-season C₃ grasses Elymus canadensis and Koeleria cristata were significantly reduced in the benomyl-treated plots, whereas benomyl treatment had no significant effect on seedling emergence of the warm-season C₄ grasses A. gerardii and Panicum virgatum. The forbs showed variable responses. Seedling emergence of Liatris aspera was greater under mycorrhizal conditions, but that of Dalea purpurea was unaffected by mycorrhizal treatment. These results show that effects of mycorrhizal symbiosis on the population dynamics of co-occurring prairie plants vary significantly both among species and among different life history stages within species. The results also indicate that mycorrhizas and fire interact to influence competitive interactions and demographic patterns of tallgrass prairie plant populations.     

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.     

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.     

Fire Season and Dominance in an Illinois Tallgrass Prairie Restoration

North American prairie remnants and restorations are normally managed with dormant‐season prescribed fires. Growing‐season fire is of interest because it suppresses dominant late‐flowering grasses and forbs, thereby making available light and other resources used by subdominant grasses and forbs that comprise most prairie diversity. Here we report a twofold increase in mean frequency and richness of subdominant species after late‐summer fire. Stimulation of subdominants was indiscriminate; richness of prairie and volunteer species increased in species that flowered in early, mid‐, or late season. Early spring fire, the management tool used on this site until this experiment, had no effect on subdominant richness or frequency. Neither burn treatment affected reproductive tillering of the tallgrasses Sorghastrum nutans or Panicum virgatum. Flowering of Andropogon gerardii increased 4‐fold after early‐spring fires and 11‐fold after late‐summer fires. These preliminary results suggest that frequency and species richness of subdominants can be improved by late growing‐season fire without compromising vigor of warm‐season tallgrasses.     

Using phenological niche separation to improve management in a Northern Glaciated Plains grassland

Many of the remaining patches of untilled (native) prairie in the Northern Glaciated Plains of North America are heavily invaded by the cool‐season grasses, Bromus inermis and Poa pratensis. However, the native vegetation in these patches contains many warm‐season species. This difference in phenology can be used to benefit restoration. We conducted an experiment to examine the efficacy of restoration treatments (mowing and prescribed fire) applied early in the growing season for consecutive years to decrease cool‐season invasive plant biomass without impacting the native warm‐season species. Our treatments were successful at significantly decreasing invasive cool‐season plant biomass and increasing native warm‐season plant biomass. No differences between treatments (mowing and prescribed fire) were found. Results suggest that incorporating differences in phenology between target and nontarget species into management may increase restoration success.     

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.     

Repeated mowing to restore remnant native grasslands invaded by nonnative annual grasses: upsides and downsides above and below ground

California grasslands have been severely impacted by the invasion of nonnative annual grasses, which often limit restoration of this important ecosystem. In this study, we explored the use of mowing as a restoration tool for native perennial grasslands at the Santa Rosa Plateau Ecological Reserve in southern California. We sought to evaluate if, over time, mowing would reduce nonnative annual grass cover and benefit native species, especially the native bunchgrass Stipa pulchra. We hypothesized that repeated mowing, carefully timed to target nonnative annual grasses prior to seed maturation, would reduce nonnative seed inputs into the soil and eventually lead to diminished abundance of these species. We monitored vegetation in mowed and unmowed plots for 4 years, and conducted a seed bank study after 5 years to better understand the cumulative effects of mowing on native and nonnative seed inputs. Consistent with our hypotheses, we found that mowing successfully reduced nonnative annual grass cover and benefitted some native species, including S. pulchra. However, we also found that nonnative forb species showed progressive increases in mowed plots over time. We observed similar patterns of species composition in the soil seed bank. Together, these results suggest that mowing can be used to control nonnative annual grasses and increase the abundance of native bunchgrasses, but that this method may also have the unintended consequence of increasing nonnative forb species.     

Effects of Soil Carbon Amendment on Nitrogen Availability and Plant Growth in an Experimental Tallgrass Prairie Restoration

Restoration of tallgrass prairie on former agricultural land is often impeded by failure to establish a diverse native species assemblage and by difficulties with nonprairie, exotic species. High levels of available soil nitrogen (N) on such sites may favor fast‐growing exotics at the expense of more slowly growing prairie species characteristic of low‐N soils. We tested whether reducing N availability through soil carbon (C) amendments could be a useful tool in facilitating successful tallgrass prairie restoration. We added 6 kg/m² hardwood sawdust to experimental plots on an abandoned agricultural field in the Sandusky Plains of central Ohio, United States, increasing soil C by 67% in the upper 15 cm. This C amendment caused a 94% reduction in annual net N mineralization and a 27% increase in soil moisture but had no effect on total N or pH. Overall, plant mass after one growing season was reduced by 64% on amended compared with unamended soil, but this effect was less for prairie forbs (?34%) than for prairie grasses (?67%) or exotics (?62%). After the second growing season, only exotics responded significantly to the soil C amendment, with a 40% reduction in mass. The N concentration of green‐leaf tissue and of senescent leaf litter was also reduced by the soil C treatment in most cases. We conclude that soil C amendment imparts several immediate benefits for tallgrass prairie restoration––notably reduced N availability, slower plant growth, and lower competition from exotic species.     

CAUSES OF DECREASES IN PRODUCTIVITY IN UNDISTURBED TALL GRASS PRAIRIE

There is abundant evidence that protection of the tall grass prairie from grazing, mowing, or burning for even a few years results in accumulation of much litter and reduced productivity. This reduced productivity has been attributed primarily to either the insulating effect of the litter, a lowered rate of mineral recycling, or a presence of toxins in the litter. Considerable evidence exists in the literature concerning the first suggestion, but no studies have been reported concerning the last two in grasslands. A comprehensive project was conducted during the growing season of 1977 in a tall grass prairie in Oklahoma to investigate the three suggestions. It was clear from our data that increased productivity as a result of mowing and burning was not due to the elimination of toxins nor to an increased rate of turnover of organic matter. Soil moisture stress was greater in the mowed and burned plots than in the controls. Thus, the single factor examined which was consistently different, and likely to explain the increased productivity, was the higher soil temperature in the burned and mowed plots early in the growing season. The four dominant grasses in the study plots are non-Festucoid species which are known to have high optimum temperatures for growth.     

Dionaea muscipula (Venus Flytrap) Establishment, Release, and Response of Associated Species in Mowed Patches on the Rims of Carolina Bays

Carolina bays are depression wetlands of high conservation value that occur across the Southeastern Coastal Plain of the United States. Venus flytrap (Dionaea muscipula) is one rare carnivorous plant that grows in open habitats on the rims of Carolina bays. Without frequent burning, vegetation on bay rims becomes dominated by evergreen shrubs and Venus flytrap populations decline. This project examined the utility of mechanical mowing, soil clearing, transplanting, and seeding as an approach to restoring populations of Venus flytraps when fire is precluded. Mowing of patches on bay rims produced open sites with little ground‐layer vegetation. After two growing seasons, adult Venus flytraps transplanted to mowed patches showed high survivorship and relatively high leaf number/plant. Suppressed Venus flytraps existing on‐site quickly initiated growth in response to mowing. These volunteers and the transplants had higher flowering percentages than plants in reference populations. Seeds of Venus flytraps were scattered in mowed and cleared plots. Seedling establishment was low, but seedlings persisted into the second growing season. Mowing created suitable habitat for growth and flowering of adult Venus flytraps and facilitated establishment of two other carnivorous species, Sundew (Drosera capillaris) and Bladderwort (Utricularia subulata). But, mowing and clearing also facilitated invasion by four species of grasses and rushes; evergreen shrubs resprouted quickly after mowing. Maintaining persistent openings by mowing the rims of Carolina bays will be an ongoing challenge due to availability of potential invaders and rapid regrowth of shrubs.     

Seed mix design and first year management influence multifunctionality and cost‐effectiveness in prairie reconstruction

Agricultural intensification continues to diminish many ecosystem services in the North American Corn Belt. Conservation programs may be able to combat these losses more efficiently by developing initiatives that attempt to balance multiple ecological benefits. In this study, we examine how seed mix design and first year management influence three ecosystem services commonly provided by tallgrass prairie reconstructions (erosion control, weed resistance, and pollinator resources). We established research plots with three seed mixes, with and without first year mowing. The grass‐dominated “Economy” mix had 21 species and a 3:1 grass‐to‐forb seeding ratio. The forb‐dominated “Pollinator” mix had 38 species and a 1:3 grass‐to‐forb seeding ratio. The grass:forb balanced “Diversity” mix, which was designed to resemble regional prairie remnants, had 71 species and a 1:1 grass‐to‐forb ratio. To assess ecosystem services, we measured native stem density, cover, inflorescence production, and floral richness from 2015 to 2018. The Economy mix had high native cover and stem density, but produced few inflorescences and had low floral richness. The Pollinator mix had high inflorescence production and floral richness, but also had high bare ground and weed cover. The Diversity mix had high inflorescence production and floral richness (comparable to the Pollinator mix) and high native cover and stem density (comparable to the Economy mix). First year mowing accelerated native plant establishment and inflorescence production, enhancing the provisioning of ecosystem services during the early stages of a reconstruction. Our results indicate that prairie reconstructions with thoughtfully designed seed mixes can effectively address multiple conservation challenges.     

Field responses to added organic matter, arbuscular mycorrhizal fungi, and fertilizer in reclamation of taconite iron ore tailing

A three season study was conducted to determine the effect of added composted yard waste, arbuscular mycorrhizal (AM) fungi, and fertilizer on plant cover, standing crop biomass, species composition, AM fungal infectivity and spore density in coarse taconite iron ore tailing plots seeded with a mixture of native prairie grasses. Plant cover and biomass, percent seeded species, mycorrhizal infectivity and spore density were greatly increased by additions of composted yard waste. After three seasons, total plant cover was also greater in plots with added fertilizer. Third season plant cover was also greater in plots amended with the higher rate (44.8 Mg ha^{–1) of compost than the moderate rate (22.4 Mg ha-1}). Field inoculation with AM fungi also increased plant cover during the second season and infectivity during the first two seasons. Seeded native species, consisting mostly of the cover species Elymus canadensis, dominated plot vegetation during the second and third seasons. Dispersal of AM fungal propagules into nonmycorrhizal plots occurred rapidly and increased infectivity in compost-amended plots during the third season. In plots with less than 10% plant cover, AM fungal infectivity of inoculated plots was greatly reduced after the second season. The high level of plant cover and the trend of increasing proportion of mycorrhizal-dependent warm-season grasses, along with increases in infectivity, forecast the establishment of a sustainable native grass community that will meet reclamation goals.     

The Role of Seed and Vegetative Reproduction in Plant Recruitment and Demography in Tallgrass Prairie

Recruitment, establishment and survivorship of seed- and vegetatively-derived shoots were quantified biweekly in annually burned and infrequently burned tallgrass prairie to investigate the contributions of seed and vegetative reproduction to the maintenance and dynamics of tallgrass prairie plant populations, the demography of seedlings and ramets, and the influence of fire on the demography of grasses and forbs. Clonally produced grass and forb ramets comprised >99%of all established shoots present at the end of the growing season, whereas established seedlings accounted for <1%,emphasizing the rarity of successful seedling establishment and the importance of vegetative reproduction in driving the annual regeneration and dynamics of aboveground plant populations in tallgrass prairie. Most recruitment from vegetative reproduction occurred early in the growing season and was higher in annually burned than infrequently burned sites, although low levels of new stem recruitment occurred continuously throughout the growing season. Peak recruitment on annually burned prairie coincided with peak recruitment of the dominant C₄ grasses Andropogon gerardii and Sorghastrum nutans prior to prescribed spring fire, with a second peak in recruitment occurring following fire. On infrequently burned prairie, grass and forb recruitment was highest in early April and declined steadily through May. The naturalized C₃ grass, Poa pratensis, was responsible for most of the early recruitment on unburned sites, whereas A. gerardii contributed most to recruitment later in May. Infrequently burned prairie was dominated by these two grasses and contained a larger forb component than annually burned prairie. The principal demographic effect of fire was on ramet natality rather than mortality. Fire regime, plant functional group, or timing of cohort emergence before or after fire did not affect ramet survivorship. C₄ grass shoots that emerged early and were damaged by fire showed similar survivorship patterns to tillers that emerged after fire. Differences in species composition between annually burned and infrequently burned prairie are driven by fire effects on vegetative reproduction and appear to be related principally to the effect of fire and detritus accumulation on the development of belowground vegetative meristems of C₄ grasses and their emergence dynamics.     

Persistence of native and exotic plants 10 years after prairie reconstruction

Prairie reconstructions are a critical component of preservation of the imperiled tallgrass prairie ecosystem in the Midwestern United States. Sustainability of this endeavor depends on establishment of persistent cover of planted native species and resistance to noxious weeds. The goal of this study was to understand the influence of early reconstruction practices on long‐term outcomes. Twelve replicates of three planting methods (dormant‐season broadcast, growing‐season broadcast, and growing‐season drill) and three seed mix richness levels (10, 20, or 34 species), fully crossed in a completely randomized design were planted in 2005 on nine former agricultural fields located in Iowa and Minnesota. Cover by species was estimated in 2005–2007, 2010, and 2015. In 2015, cover of planted species, native nonplanted species, and exotic species were similar to those recorded in 2010. Cover of the noxious weed Cirsium arvense had also declined by an average of 49% without herbicide from a peak in 2007 to low stable levels from 2010 to 2015. Richness of planted forbs, on the other hand, were still increasing in high‐richness broadcast treatments (e.g. 17–59% increase 2010–1015 in Minnesota). Two results in 2015 are reasons for concern: cover of planted species is only slightly over 50% in both Minnesota and Iowa, though with forbs still increasing, this may improve; and the cool‐season exotic grasses Poa pratensis and Bromus inermis are increasing at both Minnesota and Iowa sites. Control of these invasive grasses will be necessary, but care will be needed to avoid negative impacts of control methods on natives.     

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.     

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.     

Control of Exotic Annual Grasses to Restore Native Forbs in Abandoned Agricultural Land

Exotic annual grasses are a major challenge to successful restoration in temperate and Mediterranean climates. Experiments to restore abandoned agricultural fields from exotic grassland to coastal sage scrub habitat were conducted over two years in southern California, U.S.A. Grass control methods were tested in 5 m² plots using soil and vegetation treatments seeded with a mix of natives. The treatments compared grass‐specific herbicide, mowing, and black plastic winter solarization with disking and a control. In year two, herbicide and mowing treatments were repeated on the first‐year plots, plus new control and solarization plots were added. Treatments were evaluated using percent cover, richness and biomass of native and exotic plants. Disking alone reduced exotic grasses, but solarization was the most effective control in both years even without soil sterilization, and produced the highest cover of natives. Native richness was greatest in solarization and herbicide plots. Herbicide application reduced exotics and increased natives more than disking or mowing, but produced higher exotic forb biomass than solarization in the second year. Mowing reduced grass biomass and cover in both years, but did not improve native establishment more than disking. Solarization was the most effective restoration method, but grass‐specific herbicide may be a valuable addition or alternative. Solarization using black plastic could improve restoration in regions with cool, wet summers or winter growing seasons by managing exotic seedbanks prior to seeding. While solarization may be impractical at very large scales, it will be useful for rapid establishment of annual assemblages on small scales.     

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.     

Plant Community Biomass Shifts in Response to Mowing and Fencing in Invaded Oak Meadows with Non‐Native Grasses and Abundant Ungulates

Many herbaceous meadows are dominated by competitive non‐native grasses and subject to ungulate herbivory, ecological processes that shift the proportional biomass of plant groups in the community. Predicting the outcome of restoration is complicated because herbivory and competition can interact. We examined the relationship between herbivory by native black‐tailed deer and domestic sheep and dominance of non‐native grasses in Garry oak meadows, one of North America's most endangered habitat types. A 3‐year factorial experiment tested the effects of mowing and fencing on plant community biomass, categorized into eight groups by geographic origin (native/non‐native), growth form (annual/perennial), and plant type (forb/grass). To test if the rarity of native plant groups was related to herbivory, we estimated ungulate foraging preferences for each plant group. Mowing and fencing treatments interacted for annual and perennial non‐native grasses. Dominance was shifted from non‐native to native grasses only when both mowing and fencing were applied. Fencing increased the total biomass, whereas mowing had no overall effect; however, fencing alone did not affect any individual plant group. Mowing shifted dominance from grasses to forbs, although both native and non‐native forbs benefited from the increased light availability. We also noted that herbivore fecal pellet densities were greatest in the spring, which coincided with the peak season of their preferred plant group, native perennial forbs. Overall, applying both mowing and fencing was the most effective restoration treatment to increase native plant groups and biomass.