Biomass and density responses in tallgrass prairie legumes to annual fire and topographic position

Annually burned tallgrass prairie is purported to be a nitrogen-limited system, especially when compared to unburned prairie. To test the hypothesis that legumes, potential nitrogen-fixers, would increase in relative abundance in annually burned sites, we assessed their density and biomass for two seasons on upland and lowland soils in annually burned and unburned watersheds. Total legume density was significantly higher in burned (8.0 ± 1.0 [SE] stems/m²) than in unburned watersheds (3.0 ± 0.3 stems/m²). Species with higher (P < 0.05) densities in burned than in unburned prairie included Amorpha canescens, Dalea candida, Dalea purpurea, Lespedeza violacea, Psoralea tenuiflora, and Schrankia nuttallii. Desmodium illinoense was the only legume that responded negatively to annual fire. Total legume biomass did not differ between burned (11.3 ± 1.3 g/m²) and unburned prairie (10.5 ± 0.9 g/m²). Biomass productions of Dalea candida and Psoralea tenuiflora were higher (P < 0.05) in burned than in unburned sites, but biomasses of other legumes were similar between burn treatments. Average individual stem masses of Amorpha canescens and Baptisia bracteata were significantly greater in unburned than in burned prairie. Legumes were affected differentially by topographic location. Total legume density was higher (P < 0.05) on lowland soils (6.6 ± 1.0 stems/m²) than on upland soils (4.3 ± 0.5 stems/m²). However, total legume biomass was not different between lowland soils (12.0 ± 1.2 g/m²) and upland soils (9.9 ± 1.0 g/m²). Densities and biomasses of Amorpha canescens, Desmodium illinoense, and Lespedeza capitata were higher on lowland sites than on upland sites, whereas densities and biomasses of Baptisia bracteata and Dalea purpurea were higher on upland than on lowland soils. Most legume species are either fire tolerant or exhibit a positive response to fire and their persistence in annually burned prairie suggests that they may play an important role in the nitrogen budget of this ecosystem.     

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.     

Long- and short-term effects of fire on nitrogen cycling in tallgrass prairie

Fires in the tallgrass prairie are frequent and significantly alter nutrient cycling processes. We evaluated the short-term changes in plant production and microbial activity due to fire and the long-term consequences of annual burning on soil organic matter (SOM), plant production, and nutrient cycling using a combination of field, laboratory, and modeling studies. In the short-term, fire in the tallgrass prairie enhances microbial activity, increases both above-and belowground plant production, and increases nitrogen use efficiency (NUE). However, repeated annual burning results in greater inputs of lower quality plant residues causing a significant reduction in soil organic N, lower microbial biomass, lower N availability, and higher C:N ratios in SOM. Changes in amount and quality of below-ground inputs increased N immobilization and resulted in no net increases in N availability with burning. This response occurred rapidly (e.g., within two years) and persisted during 50 years of annual burning. Plant production at a long-term burned site was not adversely affected due to shifts in plant NUE and carbon allocation. Modeling results indicate that the tallgrass ecosystem responds to the combined changes in plant resource allocation and NUE. No single factor dominates the impact of fire on tallgrass plant production.     

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.     

Disturbance regimes and mountain plover habitat in shortgrass steppe: Large herbivore grazing does not substitute for prairie dog grazing or fire

Restoring historical disturbance regimes to enhance habitat for grassland birds can conflict with livestock production goals and has been controversial because of uncertainty in the frequency and pattern of different disturbances prior to European settlement. We studied nesting habitat for the mountain plover (Charadrius montanus) in relation to prescribed fire, grazing by large herbivores (cattle), and grazing by black-tailed prairie dogs (Cynomys ludovicianus) in the shortgrass steppe of northeastern Colorado. Breeding mountain plovers primarily occurred on black-tailed prairie dog colonies or areas burned during the previous dormant season. Vegetation surrounding mountain plover nests and foraging locations was characterized by a fine-scale mosaic of prostrate (<4 cm tall) vegetated patches interspersed with >35% bare soil in a given square meter, with this fine-scale pattern distributed over a broad (>100-m radius) area. Mountain plovers rarely occupied grassland lacking prairie dogs or recent fire, but those that did selected sites with similar vegetation height and bare soil exposure as sites on burns and prairie dog colonies. Vegetation structure at mountain plover-occupied sites was also similar to random sites on burns and prairie dog colonies, but differed substantially from sites managed only with cattle. Intensive cattle grazing at twice the recommended stocking rate during spring (Mar–May) or summer (May–Oct) for 6 years produced significantly less bare soil than burns and prairie dog colonies, particularly following years with average or above-average precipitation. Thus, intensive cattle grazing did not substitute for prairie dog grazing or fire in terms of effects on vegetation structure and mountain plover habitat. Both prescribed burning and increased size and distribution of black-tailed prairie dog colonies appear to be effective and complementary means to manage for mountain plover breeding habitat in shortgrass steppe. Provision of mountain plover habitat has tradeoffs with traditional management for livestock production. Thus, managers need to clearly define desired outcomes for management to provide multiple ecosystem goods and services. © 2012 The Wildlife Society.     

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.     

Responses of plant and bird communities to prescribed burning in tallgrass prairies

Historic losses and fragmentation of tallgrass prairie habitat to agriculture and urban development have led to declines in diversity and abundance of plants and birds associated with such habitat. Prescribed burning is a management strategy that has potential for restoring and rejuvenating prairies in fragmented landscapes, and through such restoration, might create habitat for birds dependent upon prairies. To provide improved data for management decision-making regarding the use of prescribed fire in tallgrass prairies, we compared responses of plant and bird communities on five burned and five unburned tallgrass prairie fragments at the DeSoto National Wildlife Refuge, Iowa, USA, from 1995 to 1997. Overall species richness and diversity were unaffected by burning, but individual species of plants and birds were affected by year-treatment interactions, including northern bobwhite (Colinus virginianus) and ring-necked pheasant (Phasianus colchicus), which showed time-delayed increases in density on burned sites. Analyses of species/area relationships indicated that, collectively, many small sites did make significant contributions to plant biodiversity at landscape levels, supporting the overall conservation value of prairie fragments. In contrast, most birds species were present on larger sites. Thus, higher biodiversity in bird communities which contain area-sensitive species might require larger sites able to support larger, more stable populations, greater habitat heterogeneity, and greater opportunity for niche separation.     

Effects of fire and grazing on grasshopper sparrow nest survival

Patch-burn grazing is a management framework designed to promote heterogeneity in grasslands, creating more diverse grassland structure to accommodate the habitat requirements of many grassland species, particularly grassland birds. Published studies on the effects of patch-burn grazing on passerines have been conducted on relatively large (430–980 ha pastures), contiguous grasslands, and only 1 of these studies has investigated the reproductive success of grassland birds. We assessed the effects of the patch-burn grazing and a more traditional treatment on the nesting ecology of grasshopper sparrows (Ammodramus savannarum) in small (<37 ha pastures) grasslands located in southern Iowa from May to August of 2008 and 2009. The study pastures were grazed from May to September and prescribed burns were conducted in the spring. We investigated the effects of treatments on clutch size and modeled grasshopper sparrow nest survival as a function of multiple biological and ecological factors. We found no difference in clutch size between treatments; however, we did find a reduction in clutch size for nests that were parasitized by brown-headed cowbirds (Molothrus ater). Constant daily survival rates were greater in patch-burn grazed pastures than in grazed-and-burned pastures (patch-burn grazed rate and grazed-and-burned rate ). Competitive survival models included year, stage of nest, nest age, and cool-season grass (csg) abundance within 5 m of the nest. Overall, csg abundance had the greatest effect on survival and had a negative influence. Although survival rates were highest in patch-burn grazed pastures, multiple factors influenced grasshopper sparrow survival. Nest survival rates for both treatments were relatively low, and variables other than treatment were more instrumental in predicting grasshopper sparrow survival. We recommend decreasing overall vegetation cover if increasing nesting habitat for grasshopper sparrows is a management goal. In addition, we recommend further investigation of heterogeneity management in fragmented landscapes to better understand how it affects biodiversity in relatively small management units that typify grassland habitats in the Midwest. © 2011 The Wildlife Society.     

Seeding tallgrass prairie in monospecific patches promotes native species establishment and cover

In tallgrass prairie reconstruction, the way desired seeds are arranged on the landscape may affect species establishment, species persistence, and the establishment and persistence of undesired (nonseeded) species from the local propagule pool. To test effects of species seeding pattern on how grasslands develop spatially, we seeded 20—4 × 4–m bare soil plots with 16 tallgrass prairie species. Treatment plots were divided into 16—1 × 1–m subplots, 64—0.5 × 0.5–m subplots, 256—0.25 × 0.25–m subplots, or 1,024—0.125 × 0.125–m subplots. Each species was hand broadcast into separate subplots (1 m² total area/species) within each plot. An additional treatment included uniformly mixing and broadcasting all seeds across a plot. We recorded species cover at the 0.125 × 0.125–m scale within each plot at the beginning of the second and third growing seasons. While species persistence was greatest within plots seeded with larger subplots, plots with smaller subplots were more spatially diverse and less occupied by nonseeded species over time than larger subplot and mixed plots. As is common in reconstruction efforts, establishment was variable among species and seeding with monospecific subplots enhanced colonization of desired rhizomatous species (e.g., Heliopsis helianthoides, Monarda fistulosa, Elymus virginicus) into unoccupied locations at the expense of species from the local propagule pool. Results suggest that seeding species in smaller, monospecific patches could result in grasslands with a more balanced native species composition than those established with a seed mixture approach.     

Grasshopper fecundity responses to grazing and fire in a tallgrass prairie

Grasshopper abundance and diversity vary with management practices such as fire and grazing. Understanding how grasshopper life history traits such as fecundity respond to management practices is key to predicting grasshopper population dynamics in heterogeneous environments. Landscape-level experimental fire and bison grazing treatments at the Konza Prairie Biological Station (Manhattan, KS) provide an opportunity to examine how management affects grasshopper fecundity. Here we report on grasshopper fecundity for nine common species at Konza Prairie. From 2007 to 2009, adult female grasshoppers were collected every 3 wk from eight watersheds that varied in fire and grazing treatments. Fecundity was measured by examining female reproductive tracts, which contain a record of past and current reproductive activity. Body size was a poor predictor of fecundity for all species. Despite large differences in vegetation structure and composition with management regime (grazing and fire interval), we observed little effect of management on grasshopper fecundity. Habitat characteristics (grasshopper density, vegetation biomass, and vegetation quality; measured in 2008 and 2009) were better predictors of past fecundity than current fecundity, with species-specific responses. Fecundity increased throughout the summer, indicating that grasshoppers were able to acquire sufficient nutritional resources for egg production in the early fall when vegetation quality is generally low. Because fecundity did not vary across management treatments, population stage structure may be more important for determining population level reproduction than management regime at Konza Prairie.     

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.     

Soil microbial responses to experimental warming and clipping in a tallgrass prairie

Global surface temperature is predicted to increase by 1.4–5.8°C by the end of this century. However, the impacts of this projected warming on soil C balance and the C budget of terrestrial ecosystems are not clear. One major source of uncertainty stems from warming effects on soil microbes, which exert a dominant influence on the net C balance of terrestrial ecosystems by controlling organic matter decomposition and plant nutrient availability. We, therefore, conducted an experiment in a tallgrass prairie ecosystem at the Great Plain Apiaries (near Norman, OK) to study soil microbial responses to temperature elevation of about 2°C through artificial heating in clipped and unclipped field plots. While warming did not induce significant changes in net N mineralization, soil microbial biomass and respiration rate, it tended to reduce extractable inorganic N during the second and third warming years, likely through increasing plant uptake. In addition, microbial substrate utilization patterns and the profiles of microbial phospholipid fatty acids (PLFAs) showed that warming caused a shift in the soil microbial community structure in unclipped subplots, leading to the relative dominance of fungi as evidenced by the increased ratio of fungal to bacterial PLFAs. However, no warming effect on soil microbial community structure was found in clipped subplots where a similar scale of temperature increase occurred. Clipping also significantly reduced soil microbial biomass and respiration rate in both warmed and unwarmed plots. These results indicated that warming‐led enhancement of plant growth rather than the temperature increase itself may primarily regulate soil microbial response. Our observations show that warming may increase the relative contribution of fungi to the soil microbial community, suggesting that shifts in the microbial community structure may constitute a major mechanism underlying warming acclimatization of soil respiration.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

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.     

Nitrogen and phosphorus dynamics of a tallgrass prairie ecosystem exposed to elevated carbon dioxide

A tallgrass prairie ecosystem was exposed to ambient and twice-ambient CO₂ concentrations in open-top chambers and compared to unchambered ambient CO₂ during the entire growing season from 1989 through 1991. Dominant species were Andropogon gerardii (C₄), A. scoparius (C₄), Sorghastrum nutans (C₄) and Poa pratensis (C₃). Nitrogen and phosphorus concentrations in A. gerardii, P. pratensis and dicotyledonous herbs above ground biomass were estimated by periodic sampling throughout the growing season in 1989 and 1990. In 1991, N and P concentrations in peak biomass were estimated by an early August harvest. N and P concentrations in root production as a function of treatment were estimated using root ingrowth bags that remained in place throughout the growing season. Total N and P in above- and belowground biomass were calculated as products of concentration and peak biomass by species groups. N concentration in A. gerardii and dicotyledonous herb aboveground biomass was lower and total N higher in elevated CO₂ plots than in ambient CO₂ plots. N concentration in P. pratensis aboveground biomass was lower in elevated CO₂ plots than in ambient, but total N did not differ among treatments in 2 out of 3 years. In 1990, N concentration in root ingrowth bag biomass was lower and total N greater in elevated CO₂ than in ambient CO₂ plots. Root ingrowth bag biomass N concentration did not differ among treatments in 1991, but total N was greater in elevated CO₂ plots than in ambient CO₂ plots. P concentration was lower under elevated CO₂ compared to ambient in 1989, but did not differ substantially among treatments in 1990 or 1991. In all years, total P in aboveground A. gerardii and root ingrowth bag biomass was greater under elevated CO₂ than ambient. P concentration and total P in P. pratensis was similar among treatments.     

Plant nitrogen concentration, use efficiency, and contents in a tallgrass prairie ecosystem under experimental warming

Plant nitrogen (N) relationship has the potential to regulate plant and ecosystem responses strongly to global warming but has not been carefully examined under warmed environments. This study was conducted to examine responses of plant N relationship (i.e. leaf N concentration, N use efficiency, and plant N content in this study) to a 4‐year experimental warming in a tallgrass prairie in the central Great Plains in USA. We measured mass‐based N and carbon (C) concentrations of stem, green, and senescent leaves, and calculated N resorption efficiency, N use efficiency, plant N content, and C : N ratios of five dominant species (two C₄ grasses, one C₃ grass, and two C₃ forbs). The results showed that warming decreased N concentration of both green and senescent leaves, and N resorption efficiency for all species. N use efficiencies and C : N ratios were accordingly higher under warming than control. Total plant N content increased under warming because of warming‐induced increases in biomass production that are larger than the warming‐induced decreases in tissue N concentration. The increases in N contents in both green and senescent plant tissues suggest that warming enhanced both plant N uptake and return through litterfall in the tallgrass ecosystem. Our results also suggest that the increased N use efficiency in C₄ grasses is a primary mechanism leading to increased biomass production under warming in the grassland ecosystem.