Moderate patchiness optimizes heterogeneity, stability, and beta diversity in mesic grassland.

Heterogeneous disturbance patterns are fundamental to rangeland conservation and management because heterogeneity creates patchy vegetation, broadens niche availability, increases compositional dissimilarity, and enhances temporal stability of aboveground biomass production. Pyrodiversity is a popular concept for how variability in fire as an ecological disturbance can enhance heterogeneity, but mechanistic understanding of factors that drive heterogeneity is lacking. Mesic grasslands are examples of ecosystems in which pyrodiversity is linked strongly to broad ecological processes such as trophic interactions because grazers are attracted to recently burned areas, creating a unique ecological disturbance referred to as the fire-grazing interaction, or pyric herbivory. But several questions about the application of pyric herbivory remain: What proportion of a grazed landscape must burn, or how many patches are required, to create sufficient spatial heterogeneity and reduce temporal variability? How frequently should patches burn? Does season of fire matter? To bring theory into applied practice, we studied a gradient of grazed tallgrass prairie landscapes created by different sizes, seasons, and frequencies of fire, and used analyses sensitive to nonlinear trends. The greatest spatial heterogeneity and lowest temporal variability in aboveground plant biomass, and greatest plant functional group beta diversity, occurred in landscapes with three to four patches (25%-33% of area burned) and three- to four-year fire return intervals. Beta diversity had a positive association with spatial heterogeneity and negative relationship with temporal variability. Rather than prescribing that these results constitute best management practices, we emphasize the flexibility offered by interactions between patch number and fire frequency for matching rangeland productivity and offtake to specific management goals. As we observed no differences across season of fire, we recommend future research focus on fire frequency within a moderate proportion of the landscape burned, and consider a wider seasonal burn window.

Factors Affecting Public Preferences for Grassland Landscape Heterogeneity in the Great Plains.

Agricultural intensification has fragmented rangelands in the Great Plains, which has contributed to uniform and homogeneous landscapes and decreased biodiversity. Alternative land management practices involving fire-grazing interactions can help maintain biodiversity without affecting livestock productivity. A survey was designed to understand the factors that influence preferences among the general population towards grassland landscape heterogeneity. Given the ordinal nature of survey responses, requisite data were analyzed using a generalized ordinal logit model. Results suggested that respondents who valued open space and those who recognized a need for a varying mix of uniform grasses and grasslands preferred landscape heterogeneity. Female respondents were about two times as likely to prefer heterogeneous landscapes compared to male respondents. In contrast, population groups that preferred wildlife habitat did not desire heterogeneous landscapes. Results suggest the need for extension and outreach activities to educate certain segments of the general population regarding benefits of alternative management practices that support landscape heterogeneity in the Great Plains.

Using Regional Climate Projections to Guide Grassland Community Restoration in the Face of Climate Change.

Grassland loss has been extensive worldwide, endangering the associated biodiversity and human well-being that are both dependent on these ecosystems. Ecologists have developed approaches to restore grassland communities and many have been successful, particularly where soils are rich, precipitation is abundant, and seeds of native plant species can be obtained. However, climate change adds a new filter needed in planning grassland restoration efforts. Potential responses of species to future climate conditions must also be considered in planning for long-term resilience. We demonstrate this methodology using a site-specific model and a maximum entropy approach to predict changes in habitat suitability for 33 grassland plant species in the tallgrass prairie region of the U.S. using the Intergovernmental Panel on Climate Change scenarios A1B and A2. The A1B scenario predicts an increase in temperature from 1.4 to 6.4°C, whereas the A2 scenario predicts temperature increases from 2 to 5.4°C and much greater CO2 emissions than the A1B scenario. Both scenarios predict these changes to occur by the year 2100. Model projections for 2040 under the A1B scenario predict that all but three modeled species will lose ~90% of their suitable habitat. Then by 2080, all species except for one will lose ~90% of their suitable habitat. Models run using the A2 scenario predict declines in habitat for just four species by 2040, but models predict that by 2080, habitat suitability will decline for all species. The A2 scenario appears based on our results to be the less severe climate change scenario for our species. Our results demonstrate that many common species, including grasses, forbs, and shrubs, are sensitive to climate change. Thus, grassland restoration alternatives should be evaluated based upon the long-term viability in the context of climate change projections and risk of plant species loss.

Climate Extremes, Vegetation Change, and Decoupling of Interactive Fire-Grazing Processes Exacerbate Fly Parasitism of Cattle.

We assessed local horn fly (Haematobia irritans L.) and face fly (Musca autumnalis De Geer) communities on cattle in 2012 and 2013 relative to vegetation and climate data to understand how parasitism of cattle is influenced by change in climate and vegetation structure. We compared heterogeneity management using spatially and temporally discrete fires (i.e., patch-burning one-third of a pasture annually) to homogeneity management (i.e., burning entire pasture in 2012 then no burning in 2013), with cattle grazing all years in both treatments. Predicted emergence of horn flies and face flies was 24 and 34 d earlier in 2012 associated with earlier spring warming, a significant deviation from the five-year mean. Intraannual horn fly dynamics were explained by concurrent high ambient air temperature the day of observations, but face flies were explained by low ambient air temperatures and dry conditions 3 wk before observations. Importance values of information for the theoretic models including fire treatments ranged from 0.89 to 1, indicating that both horn flies and face flies are sensitive to habitat alterations and fire-driven animal movements. Ordination indicates herds on unburned pastures were dissimilar to herds on pastures burned with patchy fires or pastures burned completely and species-specific fly responses to different vegetation structure metrics. For example, horn flies were correlated with vegetation visual obstruction, and face flies were correlated with woody plant cover. Vegetation structure may be as important as climate in driving the dynamics of fly parasites of cattle.

Exotic-Dominated Grasslands Show Signs of Recovery with Cattle Grazing and Fire.

In grasslands, overgrazing by domestic livestock, fertilization, and introduction of exotic forage species leads to plant communities consisting of a mixture of native and exotic species. These degraded grasslands present a problem for land managers, farmers, and restoration ecologists concerned with improving biodiversity while continuing to use the land for livestock production. Here we assessed the response of butterfly and plant community composition to the use of fire and moderate grazing by domestic cattle on degraded grasslands dominated by exotic plants. We evaluated change by comparing experimental pastures to two reference sites that were grasslands dominated by native plants. We used two burning and grazing treatments: 1) patch-burn graze, a heterogeneously managed treatment, where one third of the pasture is burned each year and cattle have free access to the entire pasture, and 2) graze-and-burn, a homogenously managed treatment, where the entire pasture is grazed each year and burned in its entirety every three years. We tested for change in the butterfly and plant community composition over seven years using Bray-Curtis dissimilarity measures. Over the course of seven years, degraded pastures in both treatments became more similar to reference sites with respect to the butterfly and plant communities. Only two butterfly species and two plant functional guilds exhibited significant linear trends over time, with varying responses. Compositional changes in both the butterfly and plant communities indicate that the use of moderate grazing and fire may shift butterfly and plant communities of exotic-dominated grasslands to be more similar to reference tallgrass prairies over time.

Temporal variability in aboveground plant biomass decreases as spatial variability increases.

Ecological theory predicts that diversity decreases variability in ecosystem function. We predict that, at the landscape scale, spatial variability created by a mosaic of contrasting patches that differ in time since disturbance will decrease temporal variability in aboveground plant biomass. Using data from a multi-year study of seven grazed tallgrass prairie landscapes, each experimentally managed for one to eight patches, we show that increased spatial variability driven by spatially patchy fire and herbivory reduces temporal variability in aboveground plant biomass. This pattern is associated with statistical evidence for the portfolio effect and a positive relationship between temporal variability and functional group synchrony as predicted by metacommunity variability theory. As disturbance from fire and grazing interact to create a shifting mosaic of spatially heterogeneous patches within a landscape, temporal variability in aboveground plant biomass can be dampened. These results suggest that spatially heterogeneous disturbance regimes contribute to a portfolio of ecosystem functions provided by biodiversity, including wildlife habitat, fuel, and forage. We discuss how spatial patterns of disturbance drive variability within and among patches.

Bee Abundance and Nutritional Status in Relation to Grassland Management Practices in an Agricultural Landscape.

Grasslands provide important resources for pollinators in agricultural landscapes. Managing grasslands with fire and grazing has the potential to benefit plant and pollinator communities, though there is uncertainty about the ideal approach. We examined the relationships among burning and grazing regimes, plant communities, and Bombus species and Apis mellifera L. abundance and nutritional indicators at the Grand River Grasslands in southern Iowa and northern Missouri. Treatment regimes included burn-only, grazed-and-burned, and patch-burn graze (pastures subdivided into three temporally distinct fire patches with free access by cattle). The premise of the experimental design was that patch-burn grazing would increase habitat heterogeneity, thereby providing more diverse and abundant floral resources for pollinators. We predicted that both bee abundance and individual bee nutritional indicators (bee size and lipid content) would be positively correlated with floral resource abundance. There were no significant differences among treatments with respect to bee abundance. However, some of the specific characteristics of the plant community showed significant relationships with bee response variables. Pastures with greater abundance of floral resources had greater bee abundance but lower bee nutritional indicators. Bee nutritional variables were positively correlated with vegetation height, but, in some cases, negatively correlated with stocking rate. These results suggest grassland site characteristics such as floral resource abundance and stocking rate are of potential importance to bee pollinators and suggest avenues for further research to untangle the complex interactions between grassland management, plant responses, and bee health.

Predator identity influences the effect of habitat management on nest predation.

Predation is the leading cause of nest failure for many passerines and considerable effort is devoted to identifying the habitat characteristics and management practices that influence nest loss. The habitat components associated with nest loss are strongly influenced by the ecology of nest predators and differ among predator species as a result. Nevertheless, there is a tendency to generalize about the effects of habitat features and management on nest failure without considering how resulting patterns are influenced by nest predators. We examined how predator-specific patterns of nest loss differed among predators and in response to grassland management with fire and grazing by cattle (Bos taurus). We used video cameras to monitor and identify predators at nests of the Grasshopper Sparrow (Ammodramus savannarum), a species of conservation concern throughout its range. We observed predation by 15 different species that differed in their response to management and the habitat characteristics associated with nests they preyed on. Losses to mammals and snakes were more likely at nests with greater amounts of litter cover and tall fescue (Schedonorus phoenix). Mammals were less likely to prey on nests surrounded by greater forb cover. Nest predation by snakes was lower in burned areas, whereas predation by mammals and Brown-headed Cowbirds (Molothrus ater) was unaffected by the use of fire. Neither vegetation density at the nest, nor landscape context was related to nest loss by any predator taxon. Although there were many similarities, we identified important differences in the species composing the nest predator community between our. study and other published research. These differences are likely to be responsible for geographic variation in the influence of habitat features and management actions on nest success. Our results demonstrate the need for natural resource managers to incorporate knowledge of local nest predators and their ecology when developing management prescriptions aimed at enhancing the reproductive success of songbirds.

Spatial heterogeneity increases diversity and stability in grassland bird communities.

Grasslands are inherently dynamic in space and time, evolving with frequent disturbance from fire and herbivores. As a consequence of human actions, many remaining grasslands have become homogenous, which has led to reduced ecosystem function, biodiversity loss, and decreased ecological services. Previous research has shown that restoring inherent heterogeneity to grasslands can increase avian diversity, but the amount of heterogeneity (i.e., number of patches or fire return interval) and the impact on avian community stability have yet to be investigated. We used a unique landscape-level design to examine avian response to interacting fire and grazing across multiple experimental landscapes that represented a gradient of fire- and grazing-dependent heterogeneity. We used seven landscapes (430-980 ha; x = 627 ha) with varying levels of patchiness ranging from annually burned (one single patch) with spring-only fires to a four-year fire return interval with spring and summer fires (eight patches). This design created a range of heterogeneity as a result of pyric herbivory, an ecological process in which fire and grazing are allowed to interact in space and time. We found that greater heterogeneity across experimental landscapes resulted in increased avian diversity and stability over time. An index of bird community change, quantified as the sum of the range of detrended correspondence analysis axis site scores, was nearly four times greater in the most homogenous experimental landscape when compared to the most heterogeneous experimental landscape. Species responses were consistently positively associated with increased heterogeneity at the landscape scale, and within-experimental-landscape responses were most often related to litter cover, litter accumulation, and vegetation height. We conclude that increased fire- and grazig-dependent heterogeneity can result in high variability in the bird community at finer, transect scales, but increased diversity and stability at broad landscape scales. We recommend that future management efforts in rangelands focus on restored disturbance processes to increase heterogeneity and improve grassland bird conservation.

Knowledge and tools to enhance resilience of beef grazing systems for sustainable animal protein production.

Ruminant livestock provides meat and dairy products that sustain health and livelihood for much of the world's population. Grazing lands that support ruminant livestock provide numerous ecosystem services, including provision of food, water, and genetic resources; climate and water regulation; support of soil formation; nutrient cycling; and cultural services. In the U.S. southern Great Plains, beef production on pastures, rangelands, and hay is a major economic activity. The region's climate is characterized by extremes of heat and cold and extremes of drought and flooding. Grazing lands occupy a large portion of the region's land, significantly affecting carbon, nitrogen, and water budgets. To understand vulnerabilities and enhance resilience of beef production, a multi-institutional Coordinated Agricultural Project (CAP), the "grazing CAP," was established. Integrative research and extension spanning biophysical, socioeconomic, and agricultural disciplines address management effects on productivity and environmental footprints of production systems. Knowledge and tools being developed will allow farmers and ranchers to evaluate risks and increase resilience to dynamic conditions. The knowledge and tools developed will also have relevance to grazing lands in semiarid and subhumid regions of the world.

Connecting soil organic carbon and root biomass with land-use and vegetation in temperate grassland.

Soils contain much of Earth's terrestrial organic carbon but are sensitive to land-use. Rangelands are important to carbon dynamics and are among ecosystems most widely impacted by land-use. While common practices like grazing, fire, and tillage affect soil properties directly related to soil carbon dynamics, their magnitude and direction of change vary among ecosystems and with intensity of disturbance. We describe variability in soil organic carbon (SOC) and root biomass--sampled from 0-170 cm and 0-100 cm, respectively--in terms of soil properties, land-use history, current management, and plant community composition using linear regression and multivariate ordination. Despite consistency in average values of SOC and root biomass between our data and data from rangelands worldwide, broad ranges in root biomass and SOC in our data suggest these variables are affected by other site-specific factors. Pastures with a recent history of severe grazing had reduced root biomass and greater bulk density. Ordination suggests greater exotic species richness is associated with lower root biomass but the relationship was not apparent when an invasive species of management concern was specifically tested. We discuss how unexplained variability in belowground properties can complicate measurement and prediction of ecosystem processes such as carbon sequestration.

Fire induced reproductive mechanisms of a Symphoricarpos (Caprifoliaceae) shrub after dormant season burning.

BACKGROUND: Symphoricarpos, a genus of the Caprifoliaceae family, consists of about 15 species of clonal deciduous shrubs in North America and 1 species endemic to China. In North American tallgrass prairie, Symphoricarpos orbiculatus (buckbrush) is the dominant shrub often forming large colonies via sexual and asexual reproductive mechanisms. Symphoricarpos shrubs, in particular S. orbiculatus, use a unique sexual reproductive mechanism known as layering where vertical stems droop and the tips root upon contact with the soil. Because of conflicting societal values of S. orbiculatus for conservation and agriculture and the current attempt to restore historical fire regimes, there is a need for basic research on the biological response of S. orbiculatus to anthropogenic burning regimes. RESULTS: From 2007 through 2013 we applied prescribed fires in the late dormant season on grazed pastures in the Grand River Grasslands of Iowa. From 2011 to 2013, we measured how S. orbiculatus basal resprouting and layering stems were affected by patchy fires on grazed pastures, complete pasture fires on grazed pastures or fire exclusion without grazing for more than three years. We measured ramet height, ramet canopy diameter, stems per ramet, ramets per 100 m2, and probability of new layering stems 120 days after fire. Height in burned plots was lower than unburned plots but S. orbiculatus reached ~ 84% of pre-burn height 120 days after fire. Stems per ramet were 2x greater in the most recently burned plots due to basal re-sprouting. Canopy diameter and density of ramets was not affected by time since fire, but burned pastures had marginally lower densities than plots excluded from fire (P = 0.07). Fire triggered new layering stems and no new layering stems were found in plots excluded from fire. CONCLUSIONS: The mechanisms of both basal sprouting and aerial layering after fire suggest S. orbiculatus is tolerant to dormant season fires. Furthermore, dormant season fires, regardless if they were patchy fires or complete pasture fires, did not result in mortality of S. orbiculatus. Dormant season fires can reduce S. orbiculatus structural dominance and maintain lower ramet densities but also trigger basal resprouting and layering.

Conservation implications of native and introduced ungulates in a changing climate.

In many grasslands, grazing by large native or introduced ungulates drives ecosystem structure and function. The behavior of these animals is important as it directs the spatial effects of grazing. To the degree that temperature drives spatial components of foraging, understanding how changes in climate alter grazing behavior will provide guidance for the conservation of ecosystem goods and services. We determined the behavioral response of native bison (Bison bison) and introduced cattle (Bos taurus) to temperature in tallgrass prairie within the Great Plains, USA. We described the thermal environment by measuring operative temperature (the temperature perceived by animals) through space and time. Site selection preferences of ungulates were quantified using resource selection functions. Woody vegetation in tallgrass prairie provided a cooler thermal environment for large ungulates, decreasing operative temperature up to 16 °C in the heat of the summer. Cattle began to seek thermal refugia at lower air temperatures (24 °C) by selecting areas closer to woody vegetation and water sources. Bison, however, sought refugia within wooded areas at higher air temperatures (36 °C), which occurred much less frequently. Both species became more attracted to riparian areas as air temperature increased, with preferences increasing tenfold during the hottest periods. As predicted warming occurs across the Great Plains and other grasslands, grazing behavior and subsequent grazing effects will be altered. Riparian areas, particularly those with both water and woody vegetation, will receive greater utilization and selection by large ungulates. The use of native grazers for conservation or livestock production may mitigate negative effects caused by increased temperatures.

Experimental tree removal in tallgrass prairie: variable responses of flora and fauna along a woody cover gradient.

Woody plant encroachment is a worldwide phenomenon in grassland and savanna systems whose consequence is often the development of an alternate woodland state. Theoretically, an alternate state may be associated with changes in system state variables (e.g., species composition) or abiotic parameter shifts (e.g., nutrient availability). When state-variable changes are cumulative, such as in woody plant encroachment, the probability of parameter shifts increases as system feedbacks intensify over time. Using a Before-After Control-Impact (BACI) design, we studied eight pairs of grassland sites undergoing various levels of eastern redcedar (Juniperus virginiana) encroachment to determine whether responses of flora and fauna to experimental redcedar removal differed according to the level of pretreatment redcedar cover. In the first year after removal, herbaceous plant species diversity and evenness, woody plant evenness, and invertebrate family richness increased linearly with pretreatment redcedar cover, whereas increases in small-mammal diversity and evenness were described by logarithmic trends. In contrast, increases in woody plant diversity and total biomass of terrestrial invertebrates were accentuated at levels of higher pretreatment cover. Tree removal also shifted small-mammal species composition toward a more grassland-associated assemblage. During the second year postremoval, increases in herbaceous plant diversity followed a polynomial trend, but increases in most other metrics did not vary along the pretreatment cover gradient. These changes were accompanied by extremely high growing-season precipitation, which may have homogenized floral and faunal responses to removal. Our results demonstrate that tree removal increases important community metrics among grassland flora and fauna within two years, with some responses to removal being strongly influenced by the stage of initial encroachment and modulated by climatic variability. Our results underscore the importance of decisive management for reversing the effects of woody plant encroachment in imperiled grassland ecosystems.

Ungulate preference for burned patches reveals strength of fire-grazing interaction.

The interactions between fire and grazing are widespread throughout fire-dependent landscapes. The utilization of burned areas by grazing animals establishes the fire-grazing interaction, but the preference for recently burned areas relative to other influences (water, topography, etc.) is unknown. In this study, we determine the strength of the fire-grazing interaction by quantifying the influence of fire on ungulate site selection. We compare the preference for recently burned patches relative to the influence of other environmental factors that contribute to site selection; compare that preference between native and introduced ungulates; test relationships between area burned and herbivore preference; and determine forage quality and quantity as mechanisms of site selection. We used two large ungulate species at two grassland locations within the southern Great Plains, USA. At each location, spatially distinct patches were burned within larger areas through time, allowing animals to select among burned and unburned areas. Using fine scale ungulate location data, we estimated resource selection functions to examine environmental factors in site selection. Ungulates preferred recently burned areas and avoided areas with greater time since fire, regardless of the size of landscape, herbivore species, or proportion of area burned. Forage quality was inversely related to time since fire, while forage quantity was positively related. We show that fire is an important component of large ungulate behavior with a strong influence on site selection that drives the fire-grazing interaction. This interaction is an ecosystem process that supersedes fire and grazing as separate factors, shaping grassland landscapes. Inclusion of the fire-grazing interaction into ecological studies and conservation practices of fire-prone systems will aid in better understanding and managing these systems.

Pyric herbivory: rewilding landscapes through the recoupling of fire and grazing.

Our understanding of fire and grazing is largely based on small-scale experimental studies in which treatments are uniformly applied to experimental units that are considered homogenous. Any discussion of an interaction between fire and grazing is usually based on a statistical approach that ignores the spatial and temporal interactions on complex landscapes. We propose a new focus on the ecological interaction of fire and grazing in which each disturbance is spatially and temporally dependent on the other and results in a landscape where disturbance is best described as a shifting mosaic (a landscape with patches that vary with time since disturbance) that is critical to ecological structure and function of many ecosystems. We call this spatiotemporal interaction pyric herbivory (literal interpretation means grazing driven by fire). Pyric herbivory is the spatial and temporal interaction of fire and grazing, where positive and negative feedbacks promote a shifting pattern of disturbance across the landscape. We present data we collected from the Tallgrass Prairie Preserve in the southern Great Plains of North America that demonstrates that the interaction between free-roaming bison (Bison bison) and random fires promotes heterogeneity and provides the foundation for biological diversity and ecosystem function of North American and African grasslands. This study is different from other studies of fire and grazing because the fires we examined were random and grazing animals were free to roam and select from burned and unburned patches. For ecosystems across the globe with a long history of fire and grazing, pyric herbivory with any grazing herbivore is likely more effective at restoring evolutionary disturbance patterns than a focus on restoring any large vertebrate while ignoring the interaction with fire and other disturbances.

Vulnerability of rehabilitated agricultural production systems to invasion by nontarget plant species.

Vast areas of arable land have been retired from crop production and "rehabilitated" to improved system states through landowner incentive programs in the United States (e.g., Conservation and Wetland Reserve Programs), as well as Europe (i.e., Agri-Environment Schemes). Our review of studies conducted on invasion of rehabilitated agricultural production systems by nontarget species elucidates several factors that may increase the vulnerability of these systems to invasion. These systems often exist in highly fragmented and agriculturally dominated landscapes, where propagule sources of target species for colonization may be limited, and are established under conditions where legacies of past disturbance persist and prevent target species from persisting. Furthermore, rehabilitation approaches often do not include or successfully attain all target species or historical ecological processes (e.g., hydrology, grazing, and/or fire cycles) key to resisting invasion. Uncertainty surrounds ways in which nontarget species may compromise long term goals of improving biodiversity and ecosystem services through rehabilitation efforts on former agricultural production lands. This review demonstrates that more studies are needed on the extent and ecological impacts of nontarget species as related to the goals of rehabilitation efforts to secure current and future environmental benefits arising from this widespread conservation practice.

Small mammals as indicators of short-term and long-term disturbance in mixed prairie.

Disturbance by military maneuvers over short and long time scales may have differential effects on grassland communities. We assessed small mammals as indicators of disturbance by military maneuvers in a mixed prairie in southern Oklahoma USA. We examined sites on two soil series, Foard and Lawton, across a gradient of disturbance intensity. A MANOVA showed that abundance of small mammals was associated (p = 0.03) with short-term (cover of vehicle tracks) disturbance but was not associated (p = 0.12) with long-term (loss of soil organic carbon, SOC) disturbance intensity. At the individual species level, Sigmodon hispidus (cotton rat) and Peromyscus maniculatus (deer mouse) occurred across all levels of disturbance and in both soil types. Only P. maniculatus abundance changed (p < 0.01) with short-term disturbance and increased by about one individual per 5% of additional track-cover. Abundance of P. maniculatus also increased (p = 0.04) by about three individuals per 1% increase in soil carbon. Chaetodipus hispidus (hispid pocket mouse) and Reithrodontomys fulvescens (fulvous harvest mouse) only occurred in single soil types limiting their potential as more general indicators. Abundance of P. maniculatus was positively related to shifts in plant species composition and likely reflected changes in vegetation structure (i.e. litter depth) and forage availability resulting from disturbance. Peromyscus maniculatus may be a useful biological indicator of ecosystem change because it responded predictably to both long-term and short-term disturbance and, when coupled with soil, plant, and disturbance history variables, can reveal land condition trends.

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.

Development of a grassland integrity index based on breeding bird assemblages.

We utilized landscape and breeding bird assemblage data from three Breeding Bird Survey (BBS) routes sampled from 1965-1995 to develop and test a grassland integrity index (GII) in a mixed-grass prairie area of Oklahoma. The overall study region is extensively fragmented from long-term agricultural activity, and native habitat remnants have been degraded by recent encroachment of woody vegetation, namely eastern red cedar (Juniperus virginiana L.). The 50 individual bird survey points along the BBS routes, known as stops, were used as sample sites. Our process first focused on developing a grassland disturbance index (GDI) as a measure of cumulative landscape disturbances for these sites. The GDI was based on five key landscape variables identified in an earlier species-level study of long-term avian community dynamics: total tree, shrub, and herbaceous vegetation cover indices, overall mean landscape patch size, and grassland patch core size. The GII was then developed based on breeding bird assemblage data. Assemblages were based on commonly used response guilds reflective of five avian life history parameters: foraging mode/location, nesting location, habitat specificity, migratory pattern, and dietary guild. We tested the response of 78 candidate assemblage metrics to the GDI, and eliminated those with no or poor response or with high correlations (redundant), resulting in 13 metrics for use in the final index. Individual metric scores were scaled to fall between 0 and 10, and the cumulative index to range from 0 to 100. Although broader application and refinement are possible, the avian-based GII has an advantage over labor-intensive, habitat-based monitoring in that the GII is derived from readily available long-term BBS data. Therefore, the GII shows promise as an inexpensive tool that could easily be applied over other areas to monitor changes in regional grassland conditions.