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.     

Experimental Studies on the Competitive Balance Between two Central European Roadside Grasses With Different Growth Forms

The weakly-competitive grass, Puccinellia distans (Jacq.) Parl., and the highly competitive grass, Elymus repens (L.), coexist on roadsides in Central Europe which are regularly mowed. The effect of mowing on this existence was evaluated in situ at the roadside and in experimental field plots under non-limiting water and nutrient conditions. In four experimental garden plots, tussocks of P. distans were transplanted into monocultures of E. repens early in the growing season. After establishment, two of the plots were mowed and two left unmowed. In the unmowed plots, P. distans was quickly overtopped and died, while in mowed plots, P. distans was able to persist throughout the growing season. Despite persistence in the mowed plots, the number of tillers of P. distans declined throughout the growing season as numbers of tillers of E. repens increased. It was concluded that mowing could enhance coexistence in situations of unlimited nutrients and water where the faster growing and aggressive E. repens would exclude P. distans without mowing. At the roadside, six plots were established at a site containing both species early in the growing season. Weekly mowing was performed on three of the plots while the others were left unmowed. In mid-July, when the grasses were beginning to senesce and had produced seed-heads, foliage area for P. distans was significantly higher in the mowed than the unmowed plots while the reverse was true for E. repens. While P. distans had higher foliage area in the mowed plots, it was able to persist to seedhead production in the unmowed plots. Simulations conducted with a multispecies canopy photosynthesis model indicated that reductions in carbon gain for P. distans due to shading by foliage of E. repens did not correspond well with foliage area for P. distans at the start of experiment or in the mowed and unmowed plots in mid-July. In both the mowed and unmowed plots, the portion of foliage consisting of P. distans increased with closeness to the roadway and corresponded inversely with soil depth. At soil depths of greater than 15 cm, P. distans did not occur. More effective exploitation of shallow soil may shift the competitive balance toward P. distans and be a significant factor in the coexistence of these two species in the shallow soils at the roadside. Differences in leaf temperatures at the roadside that might result from different leaf widths were also evaluated. The narrower-leaved P. distans was hypothesized to have lower midday leaf temperatures at sites close to the asphalt and perhaps be closer to the temperature optimum for photosynthesis during warm summer days. In situ leaf temperature measurements made with small thermocouples attached to intact leaves, however, were not significantly different for the two species, and coexistence was not likely to be affected by leaf temperatures.     

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.     

Interactive Effects of Fire, Elevated Carbon Dioxide, Nitrogen Deposition, and precipitation on a California Annual Grassland

Although it is widely accepted that elevated atmospheric carbon dioxide (CO₂), nitrogen (N) deposition, and climate change will alter ecosystem productivity and function in the coming decades, the combined effects of these environmental changes may be nonadditive, and their interactions may be altered by disturbances, such as fire. We examined the influence of a summer wildfire on the interactive effects of elevated CO₂, N deposition, and increased precipitation in a full-factorial experiment conducted in a California annual grassland. In unburned plots, primary production was suppressed under elevated CO₂. Burning alone did not significantly affect production, but it increased total production in combination with nitrate additions and removed the suppressive effect of elevated CO₂. Increased production in response to nitrate in burned plots occurred as a result of the enhanced aboveground production of annual grasses and forbs, whereas the removal of the suppressive effect of elevated CO₂ occurred as a result of increased aboveground forb production in burned, CO₂-treated plots and decreased root production in burned plots under ambient CO₂.The tissue nitrogen–phosphorus ratio, which was assessed for annual grass shoots, decreased with burning and increased with nitrate addition. Burning removed surface litter from plots, resulting in an increase in maximum daily soil temperatures and a decrease in soil moisture both early and late in the growing season. Measures of vegetation greenness, based on canopy spectral reflectance, showed that plants in burned plots grew rapidly early in the season but senesced early. Overall, these results indicate that fire can alter the effects of elevated CO₂ and N addition on productivity in the short term, possibly by promoting increased phosphorus availability.     

Fire and Litter Effects on Seedling Establishment in Western Oregon Upland Prairies

Prescribed burning is an important tool for managing and restoring prairies and other ecosystems. One effect of fire is plant litter removal, which can influence seedling establishment. Four experimental treatments (burned, clipped and raked to remove litter, burned with litter reapplied, and unmanipulated) were applied to 2 × 2.5–m plots in three western Oregon, United States, upland prairies to determine how burning affects seedling establishment. Seeds of common exotic and native prairie species were sowed into the experimental plots after treatments. Seedlings were censused the following spring. The experiment was repeated on each of the three sites, representing three common types of prairie vegetation: an Annual Exotic Grass site, a Perennial Exotic Grass site, and a Native Bunchgrass site. In both the Annual Exotic Grass and the Perennial Exotic Grass sites, burning significantly improved native, but not exotic, seedling establishment over those on unburned plots. Litter removal was a significant component of this burn effect, particularly on the Perennial Exotic Grass site. In these winter‐moist systems, the net effect of litter is to inhibit seedling establishment. Burning treatments on the Native Bunchgrass site significantly increased seedling establishment only of short‐lived exotic species. These results suggest that in prairie ecosystems similar to the Annual and Perennial Exotic Grass sites, prescribed burning followed by sowing native seeds can be an effective restoration technique. Burning alone or sowing alone would be counter‐productive, in the first case because increased establishment would come from exotic species and in the second case because establishment rates are low in unburned plots.     

Complex Effects of Prescribed Fire on Restoring the Soil Water Content in a High-Elevation Riparian Meadow, Arizona

One of the largest and rarest Bebb willow (Salix bebbiana) communities in the United States occurs at Hart Prairie, Arizona. Low recruitment of the willow over the past several decades has been linked to inadequate soil water content for seed germination and seedling establishment. We tested a hypothesis that a prescribed burn would reduce biomass of and evapotranspiration by herbaceous plants, thereby increasing soil water content. Three treatments (unburned control, early‐growing season burned, late‐growing season burned) were applied in year 2001 to replicated plots in fern‐ and grass‐dominated herbaceous communities. Soil water content (0–30 cm) was measured weekly in plots during the 2001, 2002, and 2003 growing seasons. Both early‐ and late‐season burning reduced herbaceous biomass in the fern‐dominated community in 2002 and 2003 and reduced biomass in the grass‐dominated community in 2002 but not in 2003. Soil water content increased for approximately four weeks in 2001 following the early‐season burn, but the early‐season and late‐season burns reduced soil water content in both communities over much of the 2002 and 2003 growing seasons. Thus, early‐season burning may benefit willow seed germination by increasing soil water content immediately following burning but be detrimental to germination in the second and third growing seasons after burning because of drier soil. Large temporal variation in the effect of prescribed burning on soil water content will complicate the use of fire as a restoration tool to manage soil water available for threatened plants such as Bebb willow and for recharge of groundwater.     

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.     

Restorative mowing on an abandoned semi‐natural meadow: short‐term and predicted long‐term effects

Abstract. When management, in the form of cattle grazing and mowing, ceases the abundance of competitively superior plant species tends to increase in abandoned semi‐natural meadows. Litter accumulation elevates the soil nutrient levels and hinders seedling recruitment. We surveyed changes in plant cover and species composition of a formerly grazed meadow in permanent plots for six years. Some plots were unmown, while others were mown and raked annually in August. The cover of grasses decreased and herb cover remained unchanged regardless of the treatment. Mowing and raking significantly reduced litter accumulation and increased the number of ground layer species. The expected long‐term effects of abandonment and restorative mowing were studied by calculating the transition probabilities for unmown and mown plots and simulating the course of succession as projected by the transition matrices. During a simulation period of 30 yr, abandonment led to (1) a decrease in the cover of small herbs, (2) a slight increase in the cover of tall herbs and (3) a slight decrease in the cover of grasses. In contrast, the cover of small herbs on the mown plots remained unchanged or slightly increased during the course of simulation. These results suggest that mowing late in the season is primarily a management tool for the maintenance of the existing species diversity and composition. However, it may not be an effective restorative tool to induce overall changes in the resident vegetation of abandoned grass‐dominated meadows. Grazing or mowing early in the season may be more effective in this respect. Consequently, mowing early or, alternatively, late in the season may provide management strategies for the maintenance and restoration of species diversity, respectively.     

POST-BURN DIFFERENCES IN SOLAR RADIATION,LEAF TEMPERATURE AND WATER STRESS INFLUENCING PRODUCTION IN A LOWLAND TALLGRASS PRAIRIE

The influence of standing dead biomass on available solar radiation, leaf temperature (T_leaf) and leaf water potential (ѱ_leaf) of Andropogon gerardii in unburned tallgrass prairie was compared to burned prairie in eastern Kansas. The standing dead reduced photosynthetically active radiation incident on emerging shoots by 58.8% in unburned compared to burned prairie during the initial 30 days of the growing season. Aboveground production in unburned prairie was similarly reduced during this period (55.4%) compared to burned prairie. Leaf temperatures in A. gerardii were greater in unburned prairie than in burned early in the season, but were nearly equal by the end of the growing season. The maximum elevation of T_leaf in unburned prairie above burned was 9.5 C. The maximum unburned T_leaf measured was 41.5 C compared to 39.4 C in burned prairie. Lower windspeed adjacent to leaves in unburned prairie resulting in reduced convective cooling may have caused higher T_leaf in unburned prairie. Leaf water potential was significantly lower in unburned prairie than in burned prairie early in the season but was higher in unburned prairie by late season. The seasonal minimum ѱ_leaf in burned prairie was — 1.60 MPa compared to —1.45 MPa in unburned prairie. The combined effect of these post-burn differences in solar radiation, T_leaf and ѱ_leaf may be significant in contributing to the lower production in unburned compared to burned tallgrass prairie.     

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.     

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.     

To Burn or Not to Burn? Effect of Management Strategy on North American Prairie Vegetation for Public Urban Areas in Germany

North American prairie vegetation has been a role model for designing highly attractive plantings for German urban green spaces for the past decade. In combination with gravel mulch top layers on planting sites and non-selective maintenance techniques like mowing or burning, prairie plantings are considered to be cost-effective and low-maintenance. This study was undertaken to assess the impact of different maintenance strategies and especially the necessity of fire management on the development success of ornamental prairie plantings in central Europe. A four factorial split-plot-block design was set up for investigation of different mixtures of prairie species under varying management conditions (mow-only, mowing plus selective weeding, mowing plus weeding and burning) on two differing soil types (in-situ topsoil and in-situ topsoil with a graywacke gravel mulch top layer) over three years. Significant effects of maintenance strategy on mortality rates and vitality were documented for a number of target species, which responded species specifically, either being slightly affected by the burning or thriving on it. Those effects were mostly restricted to topsoil sites. A strong impact on weed species presence and abundance and resulting maintenance times was found on both soil types. On topsoil sites, mow-only treatment resulted in a short-term loss of the original planting due to extensive weed growth. Corresponding gravel mulch sites were generally less colonised and visually dominated by weeds. Differences between weeded and weeded plus burned sites were minor. Unexpectedly, weed species populations were mostly unaffected by the additional burning treatment, while maintenance times and costs increased. No overall benefit of fire management for the establishment of prairie plantings was documented. The most effective management combination proved to be mowing plus regular selective weeding measures on gravel mulched planting sites.     

Restoration Efforts for Plant and Bird Communities in Tallgrass Prairies Using Prescribed Burning and Mowing

Recent losses and fragmentation of tallgrass prairie habitat to agriculture and urban development have led to corresponding declines in diversity and abundance of plants and birds associated with such habitat. Mowing and burning are alternative management strategies for restoring and rejuvenating prairies in fragmented landscapes, but their specific, comparative effects are the subjects of ongoing evaluation. We compared the responses of plant and bird communities on four sets of mowed, burned, and untreated sites of small (3–10 ha), fragmented tallgrass prairies at the DeSoto National Wildlife Refuge (DNWR), Iowa, U.S.A., during May–July in 1998 and 1999. Species richness and diversity of plants, resident grassland birds, and communities of birds associated with grassland edges (edge species) were independent of treatment. Although not affecting species richness and diversity in plant communities, mowed sites ranked lower in total plant coverage and total forb coverage than burned sites or untreated sites. In contrast, untreated sites had more coverage by shrubs, suggesting that mowing and burning did retard shrub encroachment. Overall, abundance and diversity of plants and birds were generally insensitive to management strategies. Small, fragmented sites of rare habitat may not respond in the short term to management treatments and may not be capable of supporting highly diverse communities, no matter how intensively manipulated. It is more probable that diversity of native prairie communities can be enhanced and restored only through long‐term efforts, acquisition of large land units capable of supporting stable populations, and deliberate reintroduction of species of high conservation value.     

Production on burned and unburned sand prairies during drought and non-drought years

Biomass production was determined on burned and unburned sand prairie sites in Illinois during the 1986–89 growing seasons. Severe droughts characterized the 1988 and 1989 growing seasons with precipitation being at least 50% less in these two years than in 1985 and 1986. During the growing season following a fire, in drought and non-drought years, burned sites had higher total green biomass (all species), little bluestem (Schizachyrium scoparium) root and shoot biomass, and total grass and forb biomass production than the unburned sites. In 1988 and 1989, production on all sites was frequently significantly lower each month than it was in 1986 or 1987. The differences in productivity between burn and unburned sites were less pronounced during the drought years (1988–1989) than during the non-drought years (1986–87). Grass productivity (all species) declined considerably during drought years. In contrast, forb productivity did not vary significantly among years. Despite the drought, productivity of all species and little bluestem (root and shoot combined) was best predicted by the burn condition of the sites. Unlike many studies where drought alone can have significant effects on productivity, this study shows that on sand prairies the burn condition of sites, or burning regimes, and associated influences, can have a greater substantial impact on productivity than drought. Moisture deficits due to drought may, however, have longer term effects than those detected during the span of this study.     

Symbiosis and the Regulation of Communities

Ecologists have long been interested in factors that controlthe structure of communities and the relative importance oftop-down effects of predators versus bottom-up effects of resources.There is a growing body of evidence that microbial symbiosesare important determinants of plant community structure andindirectly affect herbivore and predator trophic levels. Studieswith mycorrhizal fungi, N-fixing bacteria and endophytes ofgrasses have demonstrated that they can affect competition,coexistence, soil nutrient dynamics and plant-herbivore interactions.Long-term field experiments with one grass/endophyte interactionsuggest that grassland community structure is determined bythe fungus. While total plant productivity of experimental plotswas similar, the composition of the vegetation was altered byendophyte symbiosis. The host grass tall fescue (Festuca arundinacea)dominated plots when infected while other grasses greatly increasedin uninfected plots. Indirect evidence suggests that changesin prairie vole (Microtus ochrogaster) grazing patterns andreproductive physiology may be in part responsible for vegetationalchanges. These results provide evidence that, in addition tobottom-up and top-down forces, microbial symbionts of plantsare important determinants of community structure.     

Effect of cattail (<Emphasis Type="Italic">Typha</Emphasis> spp<Emphasis Type="Italic">.</Emphasis>) mowing on water beetle assemblages: changes of environmental factors and the aerial colonization of aquatic habitats

Some studies found no, or weak evidence that dense monotypic cattail (Typha spp.) stands exclude water beetle species from aquatic habitats, or modify aquatic beetle assemblages. Other studies suggest that cattail may reduce the chance of aerial water beetle colonization, and decreases water temperature; negatively affecting these insects. We examined the response of aquatic beetle assemblages to the mowing of cattail (Typha angustifolia L., T. latifolia L.) in a freshwater marsh. Following removal of cattail at the water level in experimental plots, aquatic beetles were sampled both in mowed and intact (control) plots weekly, through a month in the spring of 2008. Aquatic beetles were more abundant in mowed plots. Species richness was the same, but it showed different patterns in mowed and intact plots. Shannon’s diversity was similar between treatments, while evenness was lower in mowed plots. 29% of the aquatic beetles showed a strong preference for mowed plots, and 15% preferred the control plots. Water temperature was an important factor, with mowed plots having higher water temperatures because of increased solar radiation. Polarization visibility of the water surface was also a factor, since aerially colonizing (flying) aquatic beetles use horizontally polarized light reflected from the water surface to seek potential locations. Using imaging polarimetry, we showed that mowing strongly enhanced the water-reflected polarized light signal, because it reduced the screening effect of cattail leaves, which made the visual detection of water easier. Our results suggest that cattail mowing is a useful method in aquatic beetle conservation: it increases the chance of aerial colonization due to the enhanced polarization visibility of the water surface, and creates a habitat for more abundant assemblages otherwise excluded by the monodominant dense cattail stands. Thus, sustaining hemi-marsh conditions with vegetated and mowed areas is advisable to maximize overall aquatic beetle diversity.     

Growth and nutrient content of perennial grasslands following burning in semi-arid,sub-tropical Australia

Fire often increases the productivity of perennial tussock grasslands inmesic environments but can reduce growth for one or more growing seasons inaridand semi-arid environments. We examined effects of single-burns on growth andnutrient content of grasslands in sub-tropical, northwestern Australia. Thesegrasslands were dominated by Themeda triandra, a speciesoften managed by regular burning in wetter temperate and tropical zones. Burnswere in the late dry season and were replicated using small plots (5 ×5-m) within fenced areas at two sites.Total projective cover and aboveground biomass were significantly less in burntplots relative to controls for 2.5 years after burning despite four growingseasons, including the first summer, of above-average rainfall. Recovery ofburnt plots was hindered by an extended dry period in the second year,demonstrating that rainfall in subsequent seasons can be as important asrainfall in the first season in determining post-burn productivity ofgrasslandsin semi-arid environments. Greater decreases in grass cover in burnt plotsduring the extended dry period may have been due to less standing dead andlitter than controls, and therefore less insulation from extreme summertemperatures, although relationships between cover changes and cover at thestart of the period were weak. With the exception of increased pH near grasstussocks, burning had little effect on chemical characteristics of surfacesoilsin the first week. Concentrations of N, and particularly P, in abovegroundplantmaterial were greater in burnt plots four months after burning, followingsummerrains, but were either less than or similar to those in controls withincreasingly dry conditions. Significantly lower concentrations of P in greenfoliage from burnt plots during dry seasons, when uptake from soil pools wouldbe minimal, indicated that burning decreased P retranslocation from plantstores. However, we found no evidence that single-burns increased nutrientlimitations to growth because plant contents of N and P were comparable inburntand control plots during periods of adequate water supply. Our data supportprevious generalizations that prescribed burning of perennial tussockgrasslandsin semi-arid environments is mostly unnecessary because putative benefits ofincreased productivity and forage quality, characteristic of more mesicenvironments, were not realized.     

Ecological Consequences of Shifting the Timing of Burning Tallgrass Prairie

In the Kansas Flint Hills, grassland burning is conducted during a relatively narrow window because management recommendations for the past 40 years have been to burn only in late spring. Widespread prescribed burning within this restricted time frame frequently creates smoke management issues downwind. A potential remedy for the concentrated smoke production in late spring is to expand burning to times earlier in the year. Yet, previous research suggested that burning in winter or early spring reduces plant productivity and cattle weight gain while increasing the proportion of undesirable plant species. In order to better understand the ecological consequences of burning at different times of the year, plant production and species abundance were measured for 20 years on ungrazed watersheds burned annually in autumn, winter, or spring. We found that there were no significant differences in total grass production among the burns on either upland or lowland topographic positions, although spring burned watersheds had higher grass culm production and lower forb biomass than autumn and winter burned watersheds. Burning in autumn or winter broadened the window of grass productivity response to precipitation, which reduces susceptibility to mid-season drought. Burning in autumn or winter also increased the phenological range of species by promoting cool-season graminoids without a concomitant decrease in warm-season grasses, potentially widening the seasonal window of high-quality forage. Incorporating autumn and winter burns into the overall portfolio of tallgrass prairie management should increase the flexibility in managing grasslands, promote biodiversity, and minimize air quality issues caused by en masse late-spring burning with little negative consequences for cattle production.     

Vegetation responses to different spatial patterns of soil disturbance in burned and unburned tallgrass prairie

Pocket gopher (Geomyidae) disturbances are created in spatiallypredictable patterns. This may influence resource heterogeneity and affectgrassland vegetation in a unique manner. We attempt to determine the extent towhich density and spatial pattern of soil disturbances influence tallgrassprairie plant community structure and determine how these disturbances interactwith fire. To investigate the effects of explicit disturbance patterns we createdsimulated pocket gopher burrows and mounds in various spatial patterns.Simulated burrows were drilled into the soil at different densities inreplicated plots of burned and unburned prairie. Separate plots of simulatedmounds were created in burned and unburned prairie at low, medium, or high mounddensities in clumped, uniform, or random spatial dispersions. In both burned and unburned plots, increased burrow density decreasedgraminoid biomass and increased forb biomass. Total-plant and graminoid biomasswere higher in burned than unburned plots while forb biomass was higher inunburned plots. Total-plant species richness was not significantly affected byburrow density or burning treatments, but graminoid species richness increasedin unburned plots and forb species richness increased in burned plots. Plant species richness was temporarily reduced directly on mounddisturbances compared to undisturbed prairie. Over time and at larger samplingscales, the interaction of fire and mound disturbance patterns significantlyaffected total-plant and graminoid species richness. The principal effect inburned and unburned prairie was decreased total-plant and graminoid speciesrichness with increased mound disturbance intensity. Although species richness at small patch scales was not increased by anyintensity of disturbance and species composition was not altered by theestablishment of a unique guild of disturbance colonizing plants, our studyrevealed that interactions between soil disturbances and fire alter the plantcommunity dominance structure of North American tallgrass prairie primarily viachanges to graminoids. Moreover, these effects become increasingly pronouncedover time and at larger spatial sampling scales.     

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.