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.     

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.     

Interspecific nutrient transfer in a tallgrass prairie plant community

Interplant nutrient transfer may be an important ecological process in grasslands, and may significantly influence plant neighborhood interactions. We investigated the potential for phosphorus transfer between the dominant grass Andropogon gerardii and several neighboring plant species in tallgrass prairie via a field ³²PO₄ labelling experiment. The mean amount of ³²P received from donor shoots differed significantly among neighboring species and decreased with increasing distance from the donor. In general, forbs and cool-season C₃ grasses received more labelled ³²P than warm-season C₄ grasses. Phosphorus transfer occurred over distances up to 0.5 m. The effects of species and distance on movement of phosphorus changed with increasing time after labelling. The relative mass of receiver and donor shoots did not affect amounts of ³²P transfer. A benomyl fungicide treatment, applied to suppress mycorrhizal activity, likely did not affect existing vegetative hyphae and did not affect the amount of ³²P transferred. These studies demonstrate that: (1) phosphorus is transferred among neighboring species in tallgrass prairie plant communities, (2) phosphorus may be transferred over significantly greater distances than reported in other grasslands, and (3) there is differential transfer among co-occurring species. Hypothesized mechanisms accounting for these patterns in tallgrass prairie include mycorrhizal hyphal interconnections and/or extensive and differential root and rhizosphere overlap among neighboring species.     

Interacting influence of mycorrhizal symbiosis and competition on plant diversity in tallgrass prairie

In tallgrass prairie, plant species interactions regulated by their associated mycorrhizal fungi may be important forces that influence species coexistence and community structure; however, the mechanisms and magnitude of these interactions remain unknown. The objective of this study was to determine how interspecific competition, mycorrhizal symbiosis, and their interactions influence plant community structure. We conducted a factorial experiment, which incorporated manipulations of abundance of dominant competitors, Andropogon gerardii and Sorghastrum nutans, and suppression of mycorrhizal symbiosis using the fungicide benomyl under two fire regimes (annual and 4-year burn intervals). Removal of the two dominant C₄ grass species altered the community structure, increased plant species richness, diversity, and evenness, and increased abundance of subdominant graminoid and forb species. Suppression of mycorrhizal fungi resulted in smaller shifts in community structure, although plant species richness and diversity increased. Responses of individual plant species were associated with their degree of mycorrhizal responsiveness: highly mycorrhizal responsive species decreased in abundance and less mycorrhizal responsive species increased in abundance. The combination of dominant-grass removal and mycorrhizal suppression treatments interacted to increase synergistically the abundance of several species, indicating that both processes influence species interactions and community organization in tallgrass prairie. These results provide evidence that mycorrhizal fungi affect plant communities indirectly by influencing the pattern and strength of plant competitive interactions. Burning strongly influenced the outcome of these interactions, which suggests that plant species diversity in tallgrass prairie is influenced by a complex array of interacting processes, including both competition and mycorrhizal symbiosis. Received: 7 April 1999 / Accepted: 30 July 1999     

Effects of mycorrhizae on growth and demography of tallgrass prairie forbs

The effects of arbuscular mycorrhizal (AM) symbiosis on ramet and genet densities, vegetative growth rates, and flowering of three forb species were studied in native tallgrass prairie in northeastern Kansas. Mycorrhizal activity was experimentally suppressed for six growing seasons on replicate plots in an annually burned and an infrequently burned watershed with the fungicide benomyl. Benomyl reduced mycorrhizal root colonization to an average of 4.2%, approximately a two-thirds reduction relative to controls (13.7% colonization). Mycorrhizae influenced the population structure of these forbs. Although mycorrhizal suppression had no long-term effect on genet densities and no effect on ramet survivorship throughout the growing season, the number of ramets per individual was significantly increased such that ramet densities of all three species were approximately doubled in response to long-term mycorrhizal suppression. Effects of mycorrhizae on ramet growth and reproduction varied among species. Ramet growth rates, biomass, and flowering of Salvia azurea were greater in plots with active mycorrhizal symbiosis, whereas mycorrhizae reduced ramet growth rates and biomass of Artemesia ludoviciana. Aster sericeus ramet growth rates and biomass were unaffected by the fungicide applications, but its flowering was reduced.The pattern of responses of these three species to mycorrhizae differed considerably between the two sites of contrasting fire regime, indicating that the interaction of fire-induced shifts in resource availability and mycorrhizal symbiosis together modulates plant responses and the intensity and patterns of interspecific competition between and among tallgrass prairie grass and forb species. Further, the results indicate that effects of mycorrhizae on community structure are a result of interspecific differences in the balance between direct positive effects of the symbiosis on host plant performance and indirect negative effects mediated through altered competitive interactions.     

Dominant Grasses Suppress Local Diversity in Restored Tallgrass Prairie

Warm‐season (C₄) grasses commonly dominate tallgrass prairie restorations, often at the expense of subordinate grasses and forbs that contribute most to diversity in this ecosystem. To assess whether the cover and abundance of dominant grass species constrain plant diversity, we removed 0, 50, or 100% of tillers of two dominant species (Andropogon gerardii or Panicum virgatum) in a 7‐year‐old prairie restoration. Removing 100% of the most abundant species, A. gerardii, significantly increased light availability, forb productivity, forb cover, species richness, species evenness, and species diversity. Removal of a less abundant but very common species, P. virgatum, did not significantly affect resource availability or the local plant community. We observed no effect of removal treatments on critical belowground resources, including inorganic soil N or soil moisture. Species richness was inversely correlated with total grass productivity and percent grass cover and positively correlated with light availability at the soil surface. These relationships suggest that differential species richness among removal treatments resulted from treatment induced differences in aboveground resources rather than the belowground resources. Selective removal of the dominant species A. gerardii provided an opportunity for seeded forb species to become established leading to an increase in species richness and diversity. Therefore, management practices that target reductions in cover or biomass of the dominant species may enhance diversity in established and grass‐dominated mesic grassland restorations.     

Septate endophyte colonization and host responses of grasses and forbs native to a tallgrass prairie

Native tallgrass prairies support distinct dark septate endophyte (DSE) communities exemplified by Periconia macrospinosa and Microdochium sp. that were recently identified as common root symbionts in this system. Since these DSE fungi were repeatedly isolated from grasses and forbs, we aimed to test their abilities to colonize different hosts. One Microdochium and three Periconia strains were screened for colonization and growth responses using five native grasses and six forbs in an in vitro system. Previously published data for an additional grass (Andropogon gerardii) were included and reanalyzed. Presence of indicative inter- and intracellular structures (melanized hyphae, microsclerotia, and chlamydospores) demonstrated that all plant species were colonized by the DSE isolates albeit to varying degrees. Microscopic observations suggested that, compared to forbs, grasses were colonized to a greater degree in vitro. Host biomass responses varied among the host species. In broad comparisons, more grass species than forbs tended to respond positively to colonization, whereas more forb species tended to be non-responsive. Based on the suspected differences in the levels of colonization, we predicted that tallgrass prairie grasses would support greater DSE colonization than forbs in the field. A survey of field-collected roots from 15 native species supported this hypothesis. Our study supports the “broad host range” of DSE fungi, although the differences in the rates of colonization in the laboratory and in the field suggest a greater compatibility between grasses and DSE fungi. Furthermore, host responses to DSE range from mutualism to parasitism, suggesting a genotype-level interplay between the fungi and their hosts that determines the outcome of this symbiosis.     

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.     

Determinants of C3 forb growth and production in a C4 dominated grassland

Forbs are the most abundant species within the vascular flora of tallgrass prairie and they make the greatest contribution to biodiversity of all growth forms. However, little is known about the factors that determine their productivity and growth rates. The objective of this study was to assess the controls of forb growth (absolute and relative) and production in tallgrass prairie from a long-term burning experiment at the Konza Prairie in NE Kansas. Over the 14-year study, forbs comprised 9% of the total biomass production on sites with a high fire frequency vs. 29% on the low fire frequency site, with gramminoids accounting for the remainder. Although interannual variations in peak biomass of the grasses was strongly correlated with environmental variables related to water availability, there were no similar relationships for forbs, suggesting that production of forbs and grasses responded to interannual variations in climate in different ways. Multivariate analysis of climatic controls on growth rates of grasses and forbs yielded similar results. Although forbs had low biomass and absolute growth per unit ground area in frequently burned prairie, their relative growth rates were highest in such sites. Thus, it appears that reduced growth rates of individual forbs per se do not limit forb success in annually burned prairie. Instead, direct negative effects of fire on forbs (increased mortality) may be responsible. Determinants of forb growth and productivity in unburned prairie remain unresolved.     

Effects of mycorrhizae on plant growth and dynamics in experimental tall grass prairie microcosms

Experimental microcosms (40 X 52 X 32 cm) containing an assemblage of eight tallgrass prairie grass and forb species in native prairie soil were maintained under mycorrhizal (untreated control) or mycorrhizal-suppressed (fungicide-treated) conditions to examine plant growth, demographic, and community responses to mycorrhizal symbiosis. The fungicide benomyl successfully reduced mycorrhizal root colonization in the fungicide-treated microcosms to only 6.4% (an 83% reduction relative to mycorrhizal controls). Suppression of mycorrhizas resulted in a 31% reduction in total net aboveground plant production and changes in the relative production of C4 and C3 plants. The C4 tallgrasses Andropogon gerardi and Sorghastrum nutans produced less plant biomass in the fungicide-treated microcosms, and had a greater ratio of reproductive to vegetative biomass. Cool-season C3 grasses, Koeleria pyramidata and Poa pratensis accumulated more biomass and were a significantly greater proportion of total community biomass in mycorrhizal-suppressed microcosms. Forbs showed variable responses to mycorrhizal suppression. The two legumes Amorpha canescens and Dalea purpurea had significantly lower survivorship in the fungicide-treated microcosms, relative to the controls. The results confirm the high mycorrhizal dependency and growth responsiveness of dominant prairie grasses, and indicate that differential growth and demographic responses to mycorrhizal colonization among species may significantly affect plant productivity and species relative abundances in tallgrass prairie.     

Controls of nitrogen limitation in tallgrass prairie

Summary The relationship between fire frequency and N limitation to foliage production in tallgrass prairie was studied with a series of fire and N addition experiments. Results indicated that fire history affected the magnitude of the vegetation response to fire and to N additions. Sites not burned for over 15 years averaged only a 9% increase in foliage biomass in response to N enrichment. In contrast, foliage production increased an average of 68% in response to N additions on annually burned sites, while infrequently burned sites, burned in the year of the study, averaged a 45% increase. These findings are consistent with reports indicating that reduced plant growth on unburned prairie is due to shading and lower soil temperatures, while foliage production on frequently burned areas is constrained by N availability. Infrequent burning of unfertilized prairie therefore results in a maximum production response in the year of burning relative to either annually burned or long-term unburned sites.Foliage biomass of tallgrass prairie is dominated by C₄ grasses; however, forb species exhibited stronger production responses to nitrogen additions than did the grasses. After four years of annual N additions, forb biomass exceeded that of grass biomass on unburned plots, and grasses exhibited a negative response to fertilizer, probably due to competition from the forbs. The dominant C₄ grasses may out-compete forbs under frequent fire conditions not only because they are better adapted to direct effects of burning, but because they can grow better under low available N regimes created by frequent fire.     

Degraded soil increases the performance of a dominant grass, <Emphasis Type="Italic">Andropogon gerardii</Emphasis> (Big bluestem)

Dominant grasses can suppress subordinate species in grassland restorations. Examining factors that influence performance of a dominant grass when interacting with subordinate forbs may provide insights for maintaining plant community diversity. The objective of our study was to determine how soils of different restoration ages and functionally different forbs influence the performance (using biomass and tillering rate as proxies) of a dominant grass: Andropogon gerardii. Sites included a cultivated field and two restored prairies (4 or 16 years after restoration) at Konza Prairie (NE Kansas). We hypothesized A. gerardii performance would be greater in more degraded soils and when interacting with legumes. Soil structure, nutrient status, and microbial biomass were measured in soil that was used to conduct the plant interaction study. Andropogon gerardii performance was measured during an 18-week greenhouse experiment using the relative yield index calculated from net absolute tillering rate and final biomass measurements in three soil restoration age treatments combined with four interacting forb treatments. Restoration improved soil structure, reduced plant-available nutrients, and increased microbial biomass. Relative yield index values of A. gerardii were greater with non-legumes than legumes. Andropogon gerardii performed best in degraded soils, which may explain the difficulty in restoring tallgrass prairie diversity in long-term cultivated soil. Results from this study suggest practices that promote soil aggregation and fungal biomass, coupled with including a high abundance of legumes in seed mixes could reduce dominance of A. gerardii and likely increase plant diversity in tallgrass prairie restorations.     

Plant–microbe interactions change along a tallgrass prairie restoration chronosequence

Soil microbial communities are critical in determining the performance and density of species in plant communities. However, their role in regulating the success of restorations is much less clear. This study assessed the ability of soil microbial communities to regulate the growth and performance of two potentially dominant grasses and two common forbs in tallgrass prairie restorations. Specifically, we examined the effects of soil microbial communities along a restoration chronosequence from agricultural fields to remnant prairies using experimentally inoculated soils. The two grass species, Andropogon gerardii and Sorghastrum nutans, grew best with the agricultural inoculates and experienced a decline in performance in later stages of the chronosequence, indicating that the microbial community shifted from being beneficial to grasses in the early stages to inhibiting grasses in the later stages of restoration. Growth of the forb, Silphium terebinthinaceum, varied little with inoculation or position along the restoration chronosequence. Growth of Baptisia leucantha, a legume, appeared limited by nodule formation in agricultural soils, peaked in young restoration soils along with nodule formation, but decreased in older soils as the microbial community became more antagonistic. Overall, negative feedbacks tended to be less important early in restoration, but appeared important in remnant and older restored prairies. Our results provide evidence that it may be advantageous for management practices to take negative soil feedbacks into consideration when trying to recreate the diversity of tallgrass prairies.     

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.     

Using prairie restoration to curtail invasion of Canada thistle: the importance of limiting similarity and seed mix richness

Theory has predicted, and many experimental studies have confirmed, that resident plant species richness is inversely related to invisibility. Likewise, potential invaders that are functionally similar to resident plant species are less likely to invade than are those from different functional groups. Neither of these ideas has been tested in the context of an operational prairie restoration. Here, we tested the hypotheses that within tallgrass prairie restorations (1) as seed mix species richness increased, cover of the invasive perennial forb, Canada thistle (Cirsium arvense) would decline; and (2) guilds (both planted and arising from the seedbank) most similar to Canada thistle would have a larger negative effect on it than less similar guilds. Each hypothesis was tested on six former agricultural fields restored to tallgrass prairie in 2005; all were within the tallgrass prairie biome in Minnesota, USA. A mixed-model with repeated measures (years) in a randomized block (fields) design indicated that seed mix richness had no effect on cover of Canada thistle. Structural equation models assessing effects of cover of each planted and non-planted guild on cover of Canada thistle in 2006, 2007, and 2010 revealed that planted Asteraceae never had a negative effect on Canada thistle. In contrast, planted cool-season grasses and non-Asteraceae forbs, and many non-planted guilds had negative effects on Canada thistle cover. We conclude that early, robust establishment of native species, regardless of guild, is of greater importance in resistance to Canada thistle than is similarity of guilds in new prairie restorations.     

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.     

Effects of plant species and soil history on root morphology,arbuscular mycorrhizal colonization of roots,and biomass in four tallgrass prairie species

Although dominant C₄ grasses in tallgrass prairie are highly mycotrophic, for many non-dominant species neither extent of mycorrhizal colonization nor root morphology effects on plant–soil feedback interactions are known. In a laboratory study conducted from November 2013 to February 2014 at Governors State University (University Park, IL), we grew plants of a dominant C₄ grass (Andropogon gerardii) and three non-dominant forbs (Symphyotrichum laeve var. laeve, Symphyotrichum novae-angliae and Parthenium integrifolium) individually in soil collected in the field beneath a conspecific, collected beneath another study species, or in sterilized soil. The study addressed the following questions: (1) Is extent of mycorrhizal colonization of roots related to root structure? (2) How does soil history interact with plant root traits to influence plant–soil feedback? (3) How might plant–soil feedback patterns influence competitive interactions among study species? We found that proportion arbuscular mycorrhizal fungi (AMF) colonization decreased with increasing specific root length. Soil history had a stronger influence than plant species on total biomass of plants, with all species having highest total biomass when grown in Andropogon soil. Consequently, net, or heterospecific, feedback did not differ among pairwise species combinations, and was not different from 0. While these results suggest that no study species should have a competitive advantage in the field, Andropogon might still have an advantage through mechanisms such as competition for light. Future work in the field and including less mycotrophic species is needed to better understand AMF effects on competitive interactions.

     

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 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.     

Effects of bison and cattle on growth, reproduction, and abundances of five tallgrass prairie forbs

Forb populations were sampled on Kansas tallgrass prairie to examine the effects of native (bison) and domestic (cattle) ungulates on plant growth, reproduction, and species abundances. Five locally and regionally abundant native tallgrass prairie perennials, Baptisia bracteata, Oenothera speciosa, Vernonia baldwinii, Solidago missouriensis, and Salvia azurea, were selected for study. Replicate watershed-level treatments included three grazing regimes (ungrazed, grazed by cattle, and grazed by bison), and two spring fire frequencies (annually burned and burned at 4-yr intervals). The results show that forb responses to ungulates in tallgrass prairie are complex and vary significantly among plant species, ungulate species, fire regimes, and plant life history stages. Some forbs (e.g., B. bracteata, O. speciosa, and V. baldwinii) increased in growth and reproduction in grazed sites, indicating competitive release in response to selective grazing of the dominant warm-season matrix grasses. Forbs that reduced performance in grazed sites are likely negatively affected by disturbances generated by ungulate nongrazing activities, because none of the forbs studied were directly consumed by bison or cattle. Large grazers had no detectable effect on the frequency of plant damage by other herbivores or pathogens. Significant effects of grazers on patterns of flowering and seed production were not congruent with their effects on population densities, indicating that variation in sexual reproduction plays a minor role in regulating local population abundances. Furthermore, the native and domestic ungulates differ significantly in their effects on forb growth and reproduction.