North American Tallgrass Prairie Research Articles

Big Bluestem Gene Pools in the Central and Northeastern United States

ABSTRACTBig bluestem (Andropogon gerardii Vitman) is a native grass of the North American tallgrass prairie, used as a forage crop and as a possible biofuel feedstock. Understanding the genetic diversity of big bluestem can enable better classification of accessions and increase the ability to recognize unique genetic variation for use within breeding programs. Previous studies of the genetic diversity of big bluestem have generated mixed conclusions; further study is warranted to provide clarification of the genetic structure. Genetic diversity was measured both among and within three groups of big bluestem populations: natural populations collected in Wisconsin, natural populations from the northeastern United States, and cultivars or accessions available through various gene banks (mostly originating in the Great Plains). A total of 417 amplified fragment length polymorphism loci were used for principal coordinate analysis, analysis of molecular variance, and Mantel tests. Results indicate that germplasm groups represent three distinct genetic pools with overlapping regions. Partitioning of genetic variance for each of the three groups revealed significant variance among ecoregions and hardiness zones and among populations within ecoregions and hardiness zones. Unique big bluestem germplasm is present outside of the Great Plains, providing opportunities to increase genetic variation within breeding germplasm. Efforts should be made to preserve these genetic resources.

Flowering phenology as a functional trait in a tallgrass prairie

• The timing of flowering is a critical component of the ecology of plants and has the potential to structure plant communities. Yet, we know little about how the timing of flowering relates to other functional traits, species abundance, and average environmental conditions. • Here, we assessed first flowering dates (FFDs) in a North American tallgrass prairie (Konza Prairie) for 431 herbaceous species and compared them with a series of other functional traits, environmental metrics, and species abundance across ecological contrasts. • The pattern of FFDs among the species of the Konza grassland was shaped by local climate, can be linked to resource use by species, and patterns of species abundance across the landscape. Peak FFD for the community occurred when soils were typically both warm and wet, while relatively few species began flowering when soils tended to be the driest. Compared with late-flowering species, species that flowered early had lower leaf tissue density and were more abundant on uplands than lowlands. • Flowering phenology can contribute to the structuring of grassland communities, but was largely independent of most functional traits. Therefore, selection for flowering phenology may be independent of general resource strategies.

Natural Hybrids and Gene Flow between Upland and Lowland Switchgrass

ABSTRACTSwitchgrass (Panicum virgatum L.) is a perennial grass native to the North American tallgrass prairie and savanna habitats and is broadly adapted to the central and eastern United States. Upland and lowland ecotypes represent the two major taxa within switchgrass, which have distinct but overlapping distributions. The purpose of this study was to survey a broad array of putative upland and lowland accessions for the possible presence of natural hybrids or hybrid derivatives and evidence of historic gene flow between the two ecotypes. All plants were classified as upland, lowland, or intermediate based on visual assessment of phenotype, using large nurseries of known upland or lowland plants as controls. A total of 480 plants were surveyed for 19 simple sequence repeat (SSR) markers and sequenced using five chloroplast DNA (cpDNA) segments. Genetic structure analysis revealed 21 individuals with strong evidence for intertaxa hybrid origin and another 25 individuals with moderate evidence for intertaxa hybrid origin. All but two of these individuals originated from remnant populations of the central or eastern Gulf Coast or along the Atlantic Seaboard, a region that is populated with significant quantities of both upland and lowland ecotypes. We propose the central and eastern Gulf Coast glacial refuge as the primary center of origin and diversity for switchgrass, with the western Gulf Coast as the secondary center of origin and diversity. Much of this diversity appears to have been preserved along one of the major northward postglacial migration routes, the Atlantic Seaboard.

Phenology-Based Assessment of Perennial Energy Crops in North American Tallgrass Prairie

Biomass is the largest source of renewable energy in the United States, and corn ethanol currently constitutes the vast majority of the country's biofuel. Extended plantation of annual crops for biofuel production, however, has raised concerns about long-term environmental, ecological, and socioeconomic consequences. Switchgrass (Panicum virgatum L.), along with other warm-season grasses, is native to the precolonial tallgrass prairie in North America and is identified as an alternative energy crop for cellulosic feedstocks. This article describes a phenology-based geospatial approach to mapping the geographic distribution of this perennial energy crop in the tallgrass prairie. Time series of Moderate Resolution Imaging Spectroradiometer (MODIS) satellite imagery (500-m resolution, eight-day interval) in 2007 were processed to extract five phenology metrics: end of season, season length, peak season, summer dry-down, and cumulative growth. A multitier decision tree was developed to map major crops, especially native prairie grasses in the region. The geographic context of the 20 million ha of perennial native grasses extracted in this study could be combined with economic and environmental considerations in a geographic information system to assist decision making for energy crop development in the prairie region.

The Effect of Herbicide on the Re-Establishment of Native Grasses in the Blackland Prairie

ABSTRACT: The Blackland Prairie of east-central Texas, the southernmost tip of the once vast North American Tallgrass Prairie, now covers only 1% of its original 6.8 million hectares. Thus, there is considerable interest in restoration and re-establishment of this endangered ecosystem. In a randomized complete block field experiment conducted near San Marcos, Texas, we tested the effectiveness of the post-emergent herbicide imazapic on weed control, establishment, and growth of four C4 warm-season grasses native to the Blackland Prairie (Andropogon gerardii, Sorghastrum nutans, Schizachyrium scoparium, and Bouteloua curtipendula). Treatment with three different levels of imazapic (0.07, 0.092, and 0.138 kg ai/ha) all significantly increased seedling density of native grasses, and low-to-medium levels of imazapic increased seedling shoot growth and flowering relative to water controls. End-of-season aboveground biomass of native grasses also increased with imazapic treatment while that of broadleaf species...

Fire history of a prairie/forest boundary: more than 250 years of frequent fire in a North American tallgrass prairie

Questions: Most modern fire-prone landscapes have experienced disruptions of their historic fire regimes. Are the primary tallgrass prairies of the Flint Hills reflective of a history of continuous fire occurrence? Did fire frequency, severity, size and seasonality change in connection with changes in land use? Has fire occurrence been related to drought conditions? Location: Edges of Cross Timbers forest stands at the Tallgrass Prairie Preserve (TGPP) in the Flint Hills of Osage County, Oklahoma, USA. Methods: Cross-sections of 76 Quercus stellata were collected from Cross Timbers stands at or near the grassland edge in the TGPP. Dendrochronological methods were used to identify years of formation for tree rings and fire scars. Superposed epoch analysis was used to evaluate the effect of drought conditions on fire occurrence. Results: Fires were recorded in 46.6% of the years between 1729 and 2005. In 41 cross-sections at one site, the mean fire interval between 1759 and 2003 was 2.59 years, with fire interval decreasing from a mean fire interval of 3.76 years in the early part of the record to 2.13 years in modern times. No extended periods without fire were recorded in the study area. Drought conditions had no significant effect on fire occurrence. Conclusions: In contrast with many fire-prone landscapes worldwide, the prairies of the Flint Hills have experienced no recent fire suppression or exclusion. Changes in fire frequency mark transitions in land use, primarily from being traditionally used by Native Americans to being managed for cattle production.

Ecohydrological responses to multifactor global change in a tallgrass prairie: A modeling analysis

Relative impacts of multiple global change factors on ecohydrological processes in terrestrial ecosystems have not been carefully studied. In this study, we used a terrestrial ecosystem (TECO) model to examine effects of three global change factors (i.e., climate warming, elevated CO2, and altered precipitation) individually and in combination on runoff, evaporation, transpiration, rooting zone soil moisture content, water use efficiency (WUE), and rain use efficiency (RUE) in a North American tallgrass prairie. We conducted a total of 200 different scenarios with gradual changes of the three factors for 100 years. Our modeling results show strong responses of runoff, evaporation, transpiration, and rooting zone soil moisture to changes in temperature and precipitation, while effects of CO2 changes were relatively minor. For example, runoff decreased by 50% with a 10°C increase in temperature and increased by 250% with doubled precipitation. Ecosystem‐level RUE increased with CO2, decreased with precipitation, and optimized at 4–6°C of warming. In contrast, plant‐level WUE was highest at doubled CO2, doubled precipitation, and ambient temperature. The different response patterns of RUE and WUE signify that processes at different scales responded uniquely to climate change. Combinations of temperature, CO2, and precipitation anomalies interactively affected response magnitude and/or patterns of ecohydrological processes. Our results suggest that ecohydrological processes were considerably affected by global change factors and then likely regulate other ecosystem processes, such as carbon and nitrogen cycling. In particular, substantial changes in runoff to different climate change scenarios could have policy implications because it is a major component to replenishing freshwater. These modeling results should be tested by and could influence design of field experiments on ecohydrological processes.

Hierarchical cues in brood parasite nest selection

Numerous hypotheses have been developed to explain how a brood parasite selects a host nest into which it lays its eggs. Most hypotheses address various aspects of nest placement. We used an extensive dataset to tease apart the relative strength of various hypotheses associated with nest placement and timing. These data were from North American tallgrass prairie and included nearly 2,000 nests of 17 host species known to accept eggs of the brood parasitic Brown-headed cowbird Molothrus ater. Regression tree analyses, with host species as a categorical covariate built around successive logistic regressions, implied that the “edge effect” and “perch proximity” hypotheses, the latter assessed as distance to woody vegetation, received the strongest support. Hypotheses concerning nest height, livestock proximity, habitat density, nest exposure, laying date, and host clutch size received weaker or subsidiary support, the latter meaning that the hypotheses received significant support only after edge effects or distance to woody vegetation were accounted for first. Host species was associated significantly with parasitism rate, but host species was itself correlated with various vegetation and landscape variables that we assessed. Brood parasitism rate and nest height were associated non-linearly. In addition to a clear hierarchical pattern among factors associated with rates of parasitism, several key explanatory variables had marked interactions, such as prairie edge and extent of woody vegetation or nest height and nest exposure. Such interactions, including between host species and certain landscape and vegetation variables, such as nest height and distance from woody vegetation, suggest caution is warranted when assessing the various competing hypotheses, which are not mutually exclusive.

Tallgrass Prairie Plant Community Dynamics Along a Canopy Cover Gradient of Eastern Redcedar (Juniperus virginiana L.)

North American grasslands make up less than 75% of their historic pre-European settlement area, and they continue to be converted to woodlands by woody plant encroachment. Conversion of grassland to woodland alters nutrient cycling, water use, and light penetration, which drives herbaceous plant community dynamics. Because studies examining this relationship among Juniperus species are limited largely to individual trees, we designed a study to examine the relationship between stand-level canopy cover of eastern redcedar ( Juniperus virginiana L.) and the herbaceous plant community. We documented herbaceous plant species composition, abundance, and biomass within a North American tallgrass prairie invaded by eastern redcedar in which canopy cover of eastern redcedar ranged from 0% to 80%. Herbaceous species richness declined as a function of increased canopy cover of eastern redcedar and subsequent loss of open space, but this decrease in species richness closely followed a species–area model. Moreover, composition of C 3 and C 4 grasses and forbs did not change with increasing canopy cover. Herbaceous biomass, which declined with increasing canopy cover, varied most within those plots with intermediate canopy cover. While we found that species richness and biomass declined as canopy cover increased, the decline followed a species–area relationship and was without abrupt change typical of ecological thresholds. We recommend additional research with removal of eastern redcedar trees over a range of canopy cover to assess restoration potential along the encroachment gradient. Las praderas de América del Norte comprenden menos del 75% de su área histórica previa a la ocupación europea, y continúa su conversión a vegetación de monte debido a la expansión de plantas leñosas. La conversión de praderas a montes altera el ciclado de nutrientes, el uso del agua, y la penetración de luz, que determinan la dinámica de las comunidades herbáceas. Dado que los estudios que examinan esta relación entre especies de Juniperus están limitados mayormente a árboles individuales, diseñamos un ensayo para estudiar la relación entre cobertura de canopeo a nivel del monte de Juniperus virginiana L. y la comunidad herbácea. Documentamos la composición florística del estrato herbáceo, abundancia y biomasa en una pradera de pastos altos de América del Norte invadida por J. virginiana en la que la cobertura de dosel de J. virginiana fue de 0% al 80%. La riqueza de especies herbáceas declinó en función de un incremento en la cobertura de dosel de J. virginiana y la pérdida subsecuente de espacio abierto, pero esta reducción en riqueza de especies mostró un alto grado de aproximación a un modelo de área–número de especies. Además, la composición de pastos C 3 y C 4 y de hierbas no varió ante incrementos de la cobertura de dosel. La biomasa herbácea, que declinó a medida que aumentó la cobertura de dosel, mostró la mayor variación en las parcelas con cobertura de dosel intermedias. Si bien encontramos que la riqueza de especies y la biomasa declinaron a medida que aumentó la cobertura de dosel, la reducción se aproximó a una relación de área–número de especies y no mostró cambios abruptos característicos de los umbrales ecológicos. Recomendamos investigación adicional con la remoción de J. virginiana a lo largo de un rango de cobertura de dosel para evaluar el potencial de restauración a lo largo de un gradiente de invasión de esta especie.

Big Bluestem (Andropogon gerardii; Poaceae) Communities in the Carolinas: Composition and Ecological Factors

Andropogon gerardii (Big Bluestem) is a dominant grass of the North American tallgrass prairie. It is also found in remnant populations in the eastern United States, including North and South Carolina, often in association with other species with prairie affinities. We characterized eight A. gerardii population sites across various physiographic regions of North and South Carolina. A total of 362 quadrats (1 m × 1 m) were sampled during the 2006–2008 growing seasons for species occurrence and site and quadrat frequency. Associated species were assigned a commonness index (CI). A Sørensen's Community Coefficient was used to determine floristic similarities among the sites. In addition, soil samples in three quadrats were sampled at each site at three depths (0–10 cm, 11–20 cm, and 21–30 cm) and analyzed for pH; organic C and N contents; extractable P, K, Ca, Mg, Zn, Mn; and CEC (cation exchange capacity). A total of 306 vascular plant species was identified comprising 64 families, including 99 (32%) graminoids. There were 61 (20%) Poaceae and 63 (20%) Asteraceae. Species per quadrat ranged from 1 to 13 with a mean of 5. Andropogon gerardii had the highest CI value (5900), followed by Rubus spp. (1260). Community Coefficient values were < 0.5 for all pairings between sites, indicating high divergence in species composition among even nearby sites. There were 14 rare or watch-listed species identified, including the federal endangered Helianthus schweinitzii at Troy Prairie. A total of 153 (50%) of the species had been previously described as occurring in prairie-like associations. Soil pH values varied from 4.8 to 6.9 among the sites and depths. Ca and Mg nutrient values were also highly variable. Andropogon gerardii was found at high frequencies across all sites, indicating its broad tolerance for a variety of edaphic conditions.

Herbivore and Fungal Pathogen Exclusion Affects the Seed Production of Four Common Grassland Species

Insect herbivores and fungal pathogens can independently affect plant fitness, and may have interactive effects. However, few studies have experimentally quantified the joint effects of insects and fungal pathogens on seed production in non-agricultural populations. We examined the factorial effects of insect herbivore exclusion (via insecticide) and fungal pathogen exclusion (via fungicide) on the population-level seed production of four common graminoid species (Andropogon gerardii, Schizachyrium scoparium, Poa pratensis, and Carex siccata) over two growing seasons in Minnesota, USA. We detected no interactive effects of herbivores and pathogens on seed production. However, the seed production of all four species was affected by either insecticide or fungicide in at least one year of the study. Insecticide consistently doubled the seed production of the historically most common species in the North American tallgrass prairie, A. gerardii (big bluestem). This is the first report of insect removal increasing seed production in this species. Insecticide increased A. gerardii number of seeds per seed head in one year, and mass per seed in both years, suggesting that consumption of flowers and seed embryos contributed to the effect on seed production. One of the primary insect species consuming A. gerardii flowers and seed embryos was likely the Cecidomyiid midge, Contarinia wattsi. Effects on all other plant species varied among years. Herbivores and pathogens likely reduce the dispersal and colonization ability of plants when they reduce seed output. Therefore, impacts on seed production of competitive dominant species may help to explain their relatively poor colonization abilities. Reduced seed output by dominant graminoids may thereby promote coexistence with subdominant species through competition-colonization tradeoffs.

High leaf tissue density grassland species consistently more abundant across topographic and disturbance contrasts in a North American tallgrass prairie

Understanding the coupling between plant functional traits and abundance provides insight into the often hidden forces that structure plant communities. To better understand the coupling between leaf traits and abundance of grassland species in a mesic North American grassland, we measured specific leaf area (SLA) and its two components, tissue density and thickness for 125 grassland species. Plants with high tissue density were more abundant over a 17-year period across a range of environments: uplands, grazed and ungrazed watersheds, and frequently and infrequently burned watersheds. The consistent relationships between leaf tissue density and abundance across ecological contrasts imply that belowground resource availability constrains community composition independent of grazing and burning regimes. Leaf tissue density did not explain species abundance in lowlands, where belowground resources are the highest. Neither did it explain the differential abundance of species between grazing or fire frequency contrasts, suggesting that changes in burning or grazing select for species based on other traits. Relative to leaf tissue density, SLA was a poor predictor of abundance, reinforcing a long-observed—but often ignored—call that measurements of SLA need to be coupled with thickness measurements in order to effectively predict the performance of species. More generally, future research needs to investigate which belowground resources control community composition in the grassland and whether the importance of water or nutrients change with burning and grazing.

High propagule production and reproductive fitness homeostasis contribute to the invasiveness of Lespedeza cuneata (Fabaceae)

Comparative studies of congeneric native and exotic species have proved fruitful in understanding plant traits that foster invasion. Using this approach, we investigate the complex reproductive system of the invasive Lespedeza cuneata (Dum.-Cours.) G. Don in relation to three native congeners in the variable environment of the North American tallgrass prairie. Lespedeza species produce both chasmogamous (CH) and cleistogamous (CL) flowers, and propagate clonally via vegetative buds. Utilizing multiple natural populations over 2 years, we investigated reproductive modes of individuals from bagged and unbagged treatments of each species. We found that L. cuneata produced a mean of five times as many seeds and a significantly greater number of vegetative buds than any native studied, and over twenty times as many CH flowers. Insect visitation significantly affected seed set in CH flowers, though some autonomous CH selfing occurred in all species. The invasive relied relatively less on selfing than the natives and exhibited less variation in reproductive output from both modes of reproduction. We conclude that the diverse reproductive biology and wide regeneration niche of L. cuneata in relation to its native congeners confer a fitness homeostasis that facilitates the successful spread of this invasive under a wide range of conditions.

Genetic Diversity in Switchgrass Collections Assessed by EST-SSR Markers

Switchgrass (Panicum virgatum L.), is a warm season C4 perennial grass, native to North American tall grass prairies. The high biomass production potential of switchgrass with low inputs makes it an excellent choice as a sustainable bioenergy crop. The objective of this study was to determine the genetic variability within and among 31 switchgrass populations obtained from Germplasm Resources Information Network (GRIN). Six plants from each population (186 genotypes) were characterized with 24 conserved grass expressed sequence tag-simple sequence repeats (EST-SSR) and 39 switchgrass EST-SSR markers. The partitioning of variance components based on the analysis of molecular variance (AMOVA) revealed that the variability within population was significantly higher (80%) than among populations (20%). Pair-wise genetic distance estimates based on SSR data revealed dissimilarity coefficients for genotypes ranging from 0.45 to 0.81. The uplands and lowlands were generally grouped in distinct sub-clusters. The genotypes were grouped into the different adaptive zones based on the geographical locations of the collections. Ploidy estimation showed that 20 of the accessions were tetraploid, four of them were octoploids and the remaining seven had mixed ploidy levels. Flow cytometry analysis of genotypes within cultivars collected from commercial seed sources did not always support the ploidy mixtures that were found in GRIN collections. Three genotypes of two accessions that clustered differently from other genotypes of the same accessions also had different ploidy levels.

Influence of grazing and fire frequency on small‐scale plant community structure and resource variability in native tallgrass prairie

In grasslands worldwide, grazing by ungulates and periodic fires are important forces affecting resource availability and plant community structure. It is not clear, however, whether changes in community structure are the direct effects of the disturbance (i.e. fire and grazing) or are mediated indirectly through changes in resource abundance and availability. In North American tallgrass prairies, fire and grazing often have disparate effects on plant resources and plant diversity, yet, little is known about the individual and interactive effects of fire and grazing on resource variability and how that variability relates to heterogeneity in plant community structure, particularly at small scales. We conducted a field study to determine the interactive effects of different long‐term fire regimes (annual vs four‐year fire frequency) and grazing by native ungulates (Bos bison) on small‐scale plant community structure and resource variability (N and light) in native tallgrass prairie. Grazing enhanced light and nitrogen availability, but did not affect small‐scale resource variability. In addition, grazing reduced the dominance of C4grasses which enhanced species richness, diversity and community heterogeneity. In contrast, annual fire increased community dominance and reduced species richness and diversity, particularly in the absence of grazing, but had no effect on community heterogeneity, resource availability and resource variability. Variability in the abundance of resources showed no relationship with community heterogeneity at the scale measured in this study, however we found a relationship between community dominance and heterogeneity. Therefore, we conclude that grazing generated small‐scale community heterogeneity in this mesic grassland by directly affecting plant community dominance, rather than indirectly through changes in resource variability.

Genetic Diversity, Plant Adaptation Regions, and Gene Pools for Switchgrass

Switchgrass (Panicum virgatumL.) is a perennial grass native to the North American tallgrass prairie and broadly adapted to the central and eastern USA. Transfer of germplasm throughout this region creates the potential of contaminating local gene pools with genes that are not native to a locale. The objective of this study was to identify structural patterns and spatial variation for molecular markers of switchgrass populations from the northern and central USA. Forty‐six prairie‐remnant populations and 11 cultivars were analyzed for random amplified polymorphic DNA (RAPD) markers. Although there was significant population differentiation, little of this variation was associated with geographic regions. A small amount of population differentiation was associated with hardiness zones and ecoregions, suggesting that a recent proposal to use these two criteria for defining plant adaptation regions has merit for defining gene pools and seed‐transfer zones of switchgrass. Cultivars of switchgrass cannot be differentiated from prairie‐remnant populations in the northern and central USA on the basis of RAPD markers, indicating that they are still highly representative of natural germplasm. Seed sources of switchgrass can be moved considerable distance within hardiness zones and ecoregions without causing significant contamination, pollution, swamping, or erosion of local gene pools.

Altered Ecosystem Nitrogen Dynamics as a Consequence of Land Cover Change in Tallgrass Prairie

ABSTRACT In recent decades, substantial areas of North American tallgrass prairie have been lost to the establishment and expansion of woodlands and forests, including those dominated by eastern redcedar (Juniperus virginiana). This shift in dominant plant life form, from C4 grasses to coniferous trees, may be accompanied by changes in productivity, standing stocks of biomass and nutrients and biogeochemical cycles. The goal of this study was to quantify and compare major pools and fluxes of nitrogen in recently established (≤ 80 y) redcedar forests and adjacent native grasslands. Three former grassland sites in the Flint Hills region of Kansas that developed closed-canopy redcedar forests in the recent past were paired with adjacent grassland sites on similar soil type and topographic position (n = 3 sites/land cover type), and selected soil and plant nitrogen pools and fluxes were measured in replicate plots (n = 6/site) along transects in each forest or grassland site over a 20-mo period. We found ...

Grasshopper (Orthoptera: Acrididae) communities respond to fire, bison grazing and weather in North American tallgrass prairie: a long-term study

Because both intrinsic and extrinsic factors influence insect population dynamics, operating at a range of temporal and spatial scales, it is difficult to assess their contributions. Long-term studies are ideal for assessing the relative contributions of multiple factors to abundance and community dynamics. Using data spanning 25 years, we investigate the contributions of weather at annual and decadal scales, fire return interval, and grazing by bison to understand the dynamics of abundance and community composition in grasshopper assemblages from North American continental grassland. Each of these three primary drivers of grassland ecosystem dynamics affects grasshopper population and community dynamics. Negative feedbacks in abundances, as expected for regulated populations, were observed for all feeding guilds of grasshoppers. Abundance of grasshoppers did not vary in response to frequency of prescribed burns at the site. Among watersheds that varied with respect to controlled spring burns and grazing by bison, species composition of grasshopper assemblages responded significantly to both after 25 years. However, after more than 20 years of fire and grazing treatments, the number of years since the last fire was more important than the managed long-term fire frequency per se. Yearly shifts in species composition (1983-2005), examined using non-metric multidimensional scaling and fourth-corner analysis, were best explained by local weather events occurring early in grasshopper life cycles. Large-scale patterns were represented by the Palmer Drought Severity Index and the North Atlantic Oscillation (NAO). The NAO was significantly correlated with annual mean frequencies of grasshoppers, especially for forb- and mixed-feeding species. Primary grassland drivers-fire, grazing and weather-contributing both intrinsic and extrinsic influences modulate long-term fluctuations in grasshopper abundances and community taxonomic composition.

Photosynthetic traits in C 3 and C 4 grassland species in mesocosm and field environments

The North American tallgrass prairie is composed of a diverse mix of C 3 and C 4 plant species that are subject to multiple resource limitations. C 4 grasses dominate this ecosystem, purportedly due to greater photosynthetic capacity and resource-use efficiency associated with C 4 photosynthesis. We tested the hypothesis that intrinsic physiological differences between C 3 and C 4 species are consistent with C 4 grass dominance by comparing leaf gas exchange and chlorophyll fluorescence variables for seven C 4 and C 3 herbaceous species (legumes and non-legumes) in two different settings: experimental mesocosms and natural grassland sites. In the mesocosms, C 4 grasses had higher photosynthetic rates, water potentials and water-use efficiency than the C 3 species. These differences were absent in the field, where photosynthetic rates declined similarly among non-leguminous species. Thus, intrinsic photosynthetic advantages for C 4 species measured in resource-rich mesocosms could not explain the dominance of C 4 species in the field. Instead, C 4 dominance in this ecosystem may depend more on the ability of the grasses to grow rapidly when resources are plentiful and to tolerate multiple limitations when resources are scarce.

SHOULD HETEROGENEITY BE THE BASIS FOR CONSERVATION? GRASSLAND BIRD RESPONSE TO FIRE AND GRAZING

In tallgrass prairie, disturbances such as grazing and fire can generate patchiness across the landscape, contributing to a shifting mosaic that presumably enhances biodiversity. Grassland birds evolved within the context of this shifting mosaic, with some species restricted to one or two patch types created under spatially and temporally distinct disturbance regimes. Thus, management-driven reductions in heterogeneity may be partly responsible for declines in numbers of grassland birds. We experimentally altered spatial heterogeneity of vegetation structure within a tallgrass prairie by varying the spatial and temporal extent of fire and by allowing grazing animals to move freely among burned and unburned patches (patch treatment). We contrasted this disturbance regime with traditional agricultural management of the region that promotes homogeneity (traditional treatment). We monitored grassland bird abundance during the breeding seasons of 2001-2003 to determine the influence of altered spatial heterogeneity on the grassland bird community. Focal disturbances of patch burning and grazing that shifted through the landscape over several years resulted in a more heterogeneous pattern of vegetation than uniform application of fire and grazing. Greater spatial heterogeneity in vegetation provided greater variability in the grassland bird community. Some bird species occurred in greatest abundance within focally disturbed patches, while others occurred in relatively undisturbed patches in our patch treatment. Henslow's Sparrow, a declining species, occurred only within the patch treatment. Upland Sandpiper and some other species were more abundant on recently disturbed patches within the same treatment. The patch burn treatment created the entire gradient of vegetation structure required to maintain a suite of grassland bird species that differ in habitat preferences. Our study demonstrated that increasing spatial and temporal heterogeneity of disturbance in grasslands increases variability in vegetation structure that results in greater variability at higher trophic levels. Thus, management that creates a shifting mosaic using spatially and temporally discrete disturbances in grasslands can be a useful tool in conservation. In the case of North American tallgrass prairie, discrete fires that capitalize on preferential grazing behavior of large ungulates promote a shifting mosaic of habitat types that maintain biodiversity and agricultural productivity.