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.     

Photosynthetic and Respiratory Acclimation to Experimental Warming for Four Species in a Tallgrass Prairie Ecosystem

Global temperature has been Increased by 0.6 ℃ over the past century and is predicted to Increase by 1.4-5.8 ℃ by the end of this century. It is unclear what impacts global warming will have on tallgrass species. In the present study, we examined leaf net photosynthetic rate （P.） and leaf respiration rate in darkness （Rd） of Aster erlcoldes （L.） Nesom, Ambrosia psllostachya DC., Helianthus mollis Lam., and Sorghastrum nutans （L.） Nash In response to experimental warming in a tallgrass prairie ecosystem of the Great Plains, USA, in the autumn （fall） of 2000 and through 2001. Warming has been Implemented with infrared heaters since 21 November 1999. The P. increased significantly In spring, decreased in early fall, and did not change in summer and late fall in the four species under warming compared with control. The Rd of the four species increased significantly until mid-summer and then did not change under warming. Measured temperature-response curves of P. showed that warming Increased the optimum temperature of P. （Topt） by 2.32 and 4.59 ℃ for H. mollis and S. nutans, respectively, in August, whereas there were no changes in May and September, and A. ericoldes and A. psllostachya also showed no changes in any of the 3 months. However, P. at optimum temperature （Popt） showed downregulation in September and no regulation in May and August for all four species. The temperature-response curves of Rd Illustrate that the temperature sensitivity of Rd, Q10, was lower in the warmed plots compared with the control plots, except for A. ericoides in August, whereas there were no changes In May and September for all four species. The results of the present study indicate that photosynthetic and respiratory acclimation varies with species and among seasons, occurring In the mid-growing season and not in the early and late growing seasons.     

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.     

Effects of grazing intensity on growth,reproduction, and abundance of three palatable forbs in Kansas tallgrass prairie

The effects of different intensities of cattle grazing on theaboveground growth, reproduction, and abundances of three palatable forbs werestudied in native tallgrass prairie. Populations of Asterericoides, Ruellia humilis, andAmorpha canescens were sampled at peak flowering duringthe1993–1995 growing seasons in four annually-burned sites varying incattle stocking density [ungrazed, low, moderate, high]. The threeforbs exhibited reduced shoot growth and/or reproduction under moderate toheavy grazing, and in no case did grazing increase any measure of plantperformance. Ruellia showed reduced shoot height andbiomass, percentage of stems flowering, and reproductive biomass in response tograzing. Aster showed decreases in shoot biomass andheightwith grazing. Amorpha showed no change in shoot orreproductive biomass, but a decrease in percent of flowering stems and inreproductive allocation with grazing. Patterns in the percentage of stemsgrazedindicated generally high but variable palatability among these species. Bycontrast, the three species showed inconsistent population response to grazing.Abundance (frequency) of all three species indicated no short-term changebetween years in response to grazing intensity. Responses of these speciesdiffer considerably from those of most other perennial tallgrass prairie forbsthat are unpalatable, unconsumed, and increase in performance (e.g. size,abundance) due to release from competition from the dominant grasses underungulate grazing. The results demonstrate that immediate aboveground growth andreproductive responses of established adults to grazing are not good predictorsof grazer effects on population abundances in tallgrass prairie.     

Differential responses to defoliation frequency in little bluestem (<Emphasis Type="Italic">Schizachyrium scoparium</Emphasis>) in tallgrass prairie: implications for herbivory tolerance and avoidance

Plant responses to herbivory are complex. In grasses, relative growth rate (RGR), seed, and vegetative reproduction, resource allocation, and architecture vary differentially and often nonlinearly with grazing intensity. High grazing tolerance may be achieved through compensatory photosynthesis and leaf growth, or through demographic mechanisms such as activation of a belowground dormant bud bank. This study assessed the relationship between grazing frequency and responses of Schizachyrium scoparium (little bluestem) in a tallgrass prairie, and examined the roles of tiller growth, reproduction, and bud (meristem) populations in its persistence under grazing. Genets were subjected to varying simulated grazing frequencies for a period of 2 years. Strong differential responses were observed among plant traits. RGR, biomass, and flowering showed strong nonlinear reductions in response to increasing clipping frequency, with no evidence of threshold effects. However, meristem density was unaffected, and plants maintained a large bud bank across all clipping treatments. Tiller natality decreased initially, but increased with >4 clippings, suggesting that declines in tiller RGR are partially offset by increasing tiller natality, and that variation in genet size is driven more by demography than by variation in individual tiller growth. Increased grazing frequency also resulted in differential activation of buds at different positions (emerging within vs. outside the subtending leaf sheath), explaining the shift to a more prostrate growth form observed in many caespitose grasses under persistent grazing. Thus, although this grass species lacks the capacity for compensatory foliage re-growth, the maintenance of a large dormant bud bank and the differential activation of buds in different positions contribute to its grazing tolerance and avoidance, respectively, and its long-term persistence in grazed grasslands.     

Effects of mycorrhizal symbiosis on tallgrass prairie plant–herbivore interactions

Complex relationships occur among plants, mycorrhizal fungi, and herbivores. By altering plant nutrient status, mycorrhizas may alter herbivory or plant tolerance to herbivory via compensatory regrowth. We examined these interactions by assessing grasshopper preference and plant growth and fungal colonization responses to herbivory under mycorrhizal and non‐mycorrhizal conditions within tallgrass prairie microcosms. Mycorrhizal symbiosis increased plant regrowth following defoliation, and some strongly mycotrophic plant species showed overcompensation in response to herbivory when they were mycorrhizal. Although grasshoppers spent more time on mycorrhizal plants, herbivory intensity did not differ between mycorrhizal and non‐mycorrhizal plants. Aboveground herbivory by grasshoppers significantly increased mycorrhizal fungal colonization of plant roots. Thus mycorrhizas may greatly benefit plants subjected to herbivory by stimulating compensatory growth, and herbivores, in turn, may increase the development of the symbiosis. Our results also indicate strong interspecific differences among tallgrass prairie plant species in their responses to the interaction of aboveground herbivores and mycorrhizal symbionts.     

A rodent plague on prairie diversity

Selective vole ( Microtus pennsylvanicus ) suppression of prairie grasses and forbs in experimental restorations suggests why many of the plants are likely to be uncommon in nature. Vole herbivory reduced densities of legumes and grasses and increased unpalatable forbs in replicated plantings in Illinois: six otherwise common species ( Dalea purpurea , Desmanthus illinoensis , Elymus canadensis , Panicum virgatum , Phalaris arundinacea , Sorghastrum nutans ) declined 27–89% in abundance, whereas two species ( Echinacea purpurea and Rudbeckia hirta ) increased by 61% and 1023%. Species number dropped by 19% and plant diversity (Simpson's D) by 37% in one treatment to which voles had access. Plots were planted with 18 prairie species of the region, but in even distributions of 35 or 350 seeds species⁻¹ m⁻², rather than skewed in favour of large C₄ grasses common in native remnants. Manipulation of plant composition and vole access revealed what are likely to be formative effects of rodent herbivory on vegetative composition. These experimental tallgrass communities appear to be assembling from plant species that voles prefer not to eat.     

Persistence of Native C4 Grasses under High-Intensity, Short-Duration Summer Bison Grazing in the Eastern Tallgrass Prairie

Disturbances such as burning or grazing maintain the herbaceous nature of eastern tallgrass prairie. These disturbances are also known to affect the relative abundance of warm-season (C4) and cool-season (C3) grasses in native prairie. Although burning is a commonly used tool, the utility of livestock grazing to manage restored prairie is less understood. We established five monocultures and one mixture of C4 grass species of the eastern tallgrass prairie in southern Wisconsin. To examine their persistence under high-intensity, short-duration summer grazing, we estimated cover of several functional groups and C4 species over a 6-year period (2000 through 2006) in a randomized complete block design. After a 2-year establishment phase (1998–1999), bison were rotated through paddocks two or three times annually during late June, July, or early August. All C4 grasses declined over time but at different rates depending on the species. Switchgrass ( Panicum virgatum ) decreased at the lowest rate, whereas Little bluestem ( Schizachyrium scoparium ) cover declined faster than Big bluestem ( Andropogon gerardii ), Indiangrass ( Sorghastrum nutans ), and Sideoats grama ( Bouteloua curtipendula ), whose rates of decline were not significantly different from each other. Succession followed a predictable trajectory with annual grasses initially colonizing interstitial space among C4 grasses, followed by legumes, which ultimately gave way to exotic C3 forage grasses. The focal C4 grasses remained the dominant functional group 8 years postseeding, but recolonization by non-native C3 grasses increased over the study period.     

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 bison grazing on Andropogon gerardii and Panicum virgatum in burned and unburned tallgrass paririe

Summary Responses to clipping and bison grazing in different environmental contexts were examined in two perennial grass species, Andropogon gerardii and Panicum virgatum, on the Konza Prairie in northeastern Kansas. Grazed tillers had lower relative growth rates (RGR) than clipped tillers following defoliation but this difference was transient and final biomass was not affected by mode of defoliation. Grazed tillers of both species had higher RGR throughout the season than ungrazed tillers, resulting in exact compensation for tissue lost to defoliation. However, A. gerardii tillers which had been grazed repeatedly the previous year (1988) had reduced relative growth rates, tiller biomass and tiller survival in 1989. This suggests that the short-term increase in aboveground relative growth rates after defoliation had a cost to future plant growth and tiller survival.In general, the two species had similar responses to defoliation but their responses were altered differentially by fire. The increase in RGR following defoliation of A. gerardii was relatively greater on unburned than burned prairie, and was influenced by topographic position. P. virgatum responses to defoliation were similar in burned and unburned prairie. Thus grazing, fire, and topographical position all interact to influence tiller growth dynamics and these two species respond differently to the fire and grazing interaction. In addition, fire may interact with grazing pattern to influence a plants' grazing history and thus its long-term performance.     

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.     

Overall grazing tolerance index (overall GTI) is not an ideal predictor for describing a single‐species tolerance to grazing

Plants' pattern of compensatory growth is often used to intuitively estimate their grazing tolerance. However, this tolerance is sometimes measured by the overall grazing tolerance index (overall GTI), which assumes that tolerance is a multivariate linear function of various underlying mechanisms. Because the interaction among mechanisms is not independent, the grazing tolerance expression based on overall GTI may be inconsistent with that based on compensatory growth. Through a manipulative field experiment from 2007 to 2012, we measured the responses of 12 traits of Elymus nutans to clipping under different resource availabilities in an alpine meadow and explored the compensatory aboveground biomass and the overall GTI to assess the possible differences between the two expressions of tolerance. Our results showed that these two expressions of tolerance were completely opposite. The expression based on overall GTI was over‐compensatory and did not vary with clipping and resource availability, while the expression based on compensatory aboveground biomass was under‐compensatory and altered to over‐compensation after fertilization. The over‐expression of highly variable traits with extremely high negative mean GTI to defoliation damage, the influence of random errors contained in traits considered, and the doubling weight of functional redundant traits greatly inflated the overall GTI, which leads to the inconsistency of the two tolerance expressions. This inconsistency is also associated with the different determining mechanisms of the two tolerance expressions. Our data suggest that plants' grazing tolerance is not a multivariate linear function of traits or mechanisms that determine grazing tolerance; the overall GTI is only a measure of traits' variability to defoliation damage. Our findings highlight that the tolerance of E. nutans mainly depends on the response of traits with lower variability to defoliation, and the overall GTI is not an ideal predictor for describing a single‐species tolerance to grazing.     

The effects of fire frequency and grazing on tallgrass prairie productivity and plant composition are mediated through bud bank demography

Periodic fire, grazing, and a variable climate are considered the most important drivers of tallgrass prairie ecosystems, having large impacts on the component species and on ecosystem structure and function. We used long-term experiments at Konza Prairie Biological Station to explore the underlying demographic mechanisms responsible for tallgrass prairie responses to two key ecological drivers: fire and grazing. Our data indicate that belowground bud banks (populations of meristems associated with rhizomes or other perennating organs) mediate tallgrass prairie plant response. Fire and grazing altered rates of belowground bud natality, tiller emergence from the bud bank, and both short-term (fire cycle) and long-term (>15 year) changes in bud bank density. Annual burning increased grass bud banks by 25% and decreased forb bud banks by 125% compared to burning every 4 years. Grazing increased the rate of emergence from the grass bud bank resulting in increased grass stem densities while decreasing grass bud banks compared to ungrazed prairie. By contrast, grazing increased both bud and stem density of forbs in annually burned prairie but grazing had no effect on forb bud or stem density in the 4-year burn frequency treatment. Lastly, the size of the reserve grass bud bank is an excellent predictor of long-term ANPP in tallgrass prairie and also of short-term interannual variation in ANPP associated with fire cycles, supporting our hypothesis that ANPP is strongly regulated by belowground demographic processes. Meristem limitation due to management practices such as different fire frequencies or grazing regimes may constrain tallgrass prairie responses to interannual changes in resource availability. An important consequence is that grasslands with a large bud bank may be the most responsive to future climatic change or other global change phenomena such as nutrient enrichment, and may be most resistant to exotic species invasions.     

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.     

Effect of defoliation intensity on aboveground and belowground relative growth rates

According to a simple growth model, grazed and ungrazed plants may have equal absolute growth rates provided that the relative growth rate (RGR) of grazed plants increases exponentially with grazing intensity (proportion of biomass removed). This paper reports results from an experiment designed to determine whether plants of two grass species subjected to a gradient of defoliation intensities, from 0 to 100% aboveground biomass removal, showed such a response. The relationship between aboveground RGR and defoliation intensity was exponential and closely matched the theoretical relationship of equal absolute growth rate. Thus, plants showed the same aboveground growth regardless of defoliation intensity thanks to an exponential stimulation of RGR by defoliation. Belowground RGR was depressed by defoliation of more than 20% of the above-ground biomass. In spite of the drastic modification imposed by the treatments on the relative proportions of different plant parts, after a 42-day recovery period basic allometric relationships, such as root:shoot and leafarea: weight ratios, were not affected by defoliation intensity. Exponential aboveground compensatory responses represent a key feedback process resulting in constant aboveground growth regardless of defoliation intensity and appear to be a simple consequence of strong commitments to certain allometric relationships.     

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.     

Relative herbivory tolerance and competitive ability in two dominant : subordinate pairs of perennial grasses in a native grassland

We evaluated herbivory tolerance and competitive ability within twodominant : subordinate pairs of C₄, perennial grasses at each of twosites to determine the contribution of these processes to herbivore-inducedspecies replacement. Herbivory tolerance was assessed by cumulative regrowthfrom defoliated plants of each species and competitive ability was evaluated byrelative uptake of a ¹⁵N isotope placed into the soil between pairedspecies in the field. Herbivory tolerance was similar for the dominant andsubordinate species in both plant pairs and defoliation intensity had a greaterinfluence on herbivory tolerance than did defoliation pattern. Both specieswithin the Sorghastrum nutans : Schizachyriumscoparium pairs exhibited comparable nitrogen acquisition from a¹⁵N enriched pulse with or without defoliation. In contrast,S. scoparium acquired more ¹⁵N than did itssubordinate neighbor, Bothriochloa laguroides when thisspecies pair was undefoliated. Uniform defoliation of this species pair at adefoliation intensity removing 70% of the shoot mass accentuated this responsefurther demonstrating the greater competitive ability of the dominant comparedto the subordinate species. Although the 90% defoliation intensity reducednitrogen acquisition by the dominant relative to the subordinate species,B. laguroides, it did not reduce nitrogen acquisition bythe dominant below that of the subordinate neighbor. The occurrence of similarherbivory tolerance among dominant and subordinate species indicates thatselective herbivory suppressed the greater competitive ability, rather than thegreater herbivory tolerance, of the dominant grasses in this experimentaldesign. These data suggest that interspecific competitive ability may be ofequal or greater importance than herbivory tolerance in mediatingherbivore-induced species replacement in mesic grasslands and savannas.     

Effect of stress and time for recovery on the amount of compensatory growth after grazing

Summary We tested the hypothesis that the amount of compensatory growth after defoliation is affected by the level of stress at which plants grow when defoliated and by the length of time for recovery. Growth response to defoliation went from partial compensation when plants were growing at high relative growth rates (RGR) to overcompensation when plants were more stressed and growing at low RGR. Defoliation released plants from the limitation imposed by the accumulation of old and dead tissue and this release overrode the negative effect of biomass loss. Compensatory growth resulted from a higher RGR aboveground that was not associated with a reduction in RGR belowground. Time available for recovery had a major impact on the outcome of defoliation. With a short time for recovery, RGR was decreased by defoliation because an immediate increase in net assimilation rate was overridden by a reduction in the ratio of leaf area to plant weight. After defoliation, this ratio increased quickly due to a larger allocation to leaf growth and lower leaf specific weights, resulting in higher RGR. We conclude that the compensatory response to grazing depends on the type and level of stress limiting growth. Allocation and physiological responses to stress may positively or negatively affect the response to grazing and, simultaneously, grazing may alleviate or aggravate the effects of different types of stress.     

Responses of Shrub Midstory and Herbaceous Layers to Managed Grazing and Fire in a North American Savanna (Oak Woodland) and Prairie Landscape

Worldwide, savanna remnants are losing acreage due to species replacement with shade-tolerant midstory forest species as a response to decades of fire suppression. Because canopy closes grasses and other easily ignitable fuels decline, therefore, fire, when reintroduced after years of absence, is not always effective at restoring the open structure original to these communities. Our study sought to determine if managed grazing is an alternative tool for reducing shrub densities and restoring savanna structure without the impacts on soils and native vegetation observed with unmanaged grazing. We compared effects of fire and managed grazing on shrub and herb composition within degraded oak savanna and tallgrass prairie of the U.S. Upper Midwest using a randomized complete block design. The vegetation response to treatments differed by species and by vegetation type. Total shrub stem densities declined 44% in grazed and 68% in burned paddocks within savanna and by 33% for both treatments within prairie. Within savanna, cattle reduced stem densities of Rubus spp. 97%, whereas fire reduced Ribes missouriense stems 96%. Both fire and grazing were effective at reducing stem numbers for several other shrub species but not to the same degree. Native forbs were suppressed in grazed savanna paddocks, as were native grasses in grazed prairie paddocks along with a minor increase of exotic forbs. We did not observe changes in soil bulk density. We conclude that managed grazing can serve as a valuable supplement but not as a replacement to fire for controlling shrubs in these systems.     

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