Long-Term Nitrogen Amendment Alters the Diversity and Assemblage of Soil Bacterial Communities in Tallgrass Prairie

Anthropogenic changes are altering the environmental conditions and the biota of ecosystems worldwide. In many temperate grasslands, such as North American tallgrass prairie, these changes include alteration in historically important disturbance regimes (e.g., frequency of fires) and enhanced availability of potentially limiting nutrients, particularly nitrogen. Such anthropogenically-driven changes in the environment are known to elicit substantial changes in plant and consumer communities aboveground, but much less is known about their effects on soil microbial communities. Due to the high diversity of soil microbes and methodological challenges associated with assessing microbial community composition, relatively few studies have addressed specific taxonomic changes underlying microbial community-level responses to different fire regimes or nutrient amendments in tallgrass prairie. We used deep sequencing of the V3 region of the 16S rRNA gene to explore the effects of contrasting fire regimes and nutrient enrichment on soil bacterial communities in a long-term (20 yrs) experiment in native tallgrass prairie in the eastern Central Plains. We focused on responses to nutrient amendments coupled with two extreme fire regimes (annual prescribed spring burning and complete fire exclusion). The dominant bacterial phyla identified were Proteobacteria, Verrucomicrobia, Bacteriodetes, Acidobacteria, Firmicutes, and Actinobacteria and made up 80% of all taxa quantified. Chronic nitrogen enrichment significantly impacted bacterial community diversity and community structure varied according to nitrogen treatment, but not phosphorus enrichment or fire regime. We also found significant responses of individual bacterial groups including Nitrospira and Gammaproteobacteria to long-term nitrogen enrichment. Our results show that soil nitrogen enrichment can significantly alter bacterial community diversity, structure, and individual taxa abundance, which have important implications for both managed and natural grassland ecosystems.     

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

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

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.     

Tallgrass prairie ants: their species composition,ecological roles,and response to management

Ants are highly influential organisms in terrestrial ecosystems, including the tallgrass prairie, one of the most endangered ecosystems in North America. Through their tunneling, ants affect soil properties and resource availability for animals and plants. Ants also have important ecological roles as consumers of plant tissue and seeds. In the last several decades, various organizations, agencies, and agricultural producers have attempted to create wildlife habitat or reduce soil erosion by seeding thousands of hectares of bare cropland in the central United States with tallgrass prairie seed mixes. Although initially, monitoring of these restorations and of unplowed prairie remnants focused on plants and birds, in recent years the response of invertebrates such as ants has increasingly been the subject of research. An understanding of tallgrass prairie ant communities can help land managers and scientists better monitor the ecological condition of tallgrass prairie and guide management and restoration efforts. Here I review our current knowledge of ant species found within tallgrass prairie, their ecological roles, and their response to management.     

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 C₄ grasses 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.     

Comparison of butterfly communities and abundances between marginal grasslands and conservation lands in the eastern Great Plains

The alteration and fragmentation of native tallgrass prairie in the Midwestern United States has created a need to identify other land types with the ability to support grassland butterfly species. This study examines butterfly usage of marginal grasslands, which consist of semi-natural grasslands existing within in a larger agricultural matrix, compared to grasslands managed for conservation of prairie species. Using generalized linear mixed models we analyzed how land purpose (marginal vs. conservation grasslands) affected butterfly abundance. We found grassland butterfly species to be significantly more common on conservation grasslands, whereas generalist species were significantly more common on marginal grasslands. Results of ordination analyses indicated that while many species used both types of habitats, butterfly species assemblages were distinct between habitat types and that edge to interior ratio and the floristic quality index of sites were important habitat characteristics driving this distinction. Within conservation grasslands we examined the relationship between butterfly abundance and the planting diversity used in restoring each site. We found higher diversity restorations hosted more individuals of butterflies considered habitat generalists, as well as species considered to be of conservation concern.     

Functional consequences of climate change-induced plant species loss in a tallgrass prairie

Future climate change is likely to reduce the floristic diversity of grasslands. Yet the potential consequences of climate-induced plant species losses for the functioning of these ecosystems are poorly understood. We investigated how climate change might alter the functional composition of grasslands for Konza Prairie, a diverse tallgrass prairie in central North America. With species-specific climate envelopes, we show that a reduction in mean annual precipitation would preferentially remove species that are more abundant in the more productive lowland positions at Konza. As such, decreases in precipitation could reduce productivity not only by reducing water availability but by also removing species that inhabit the most productive areas and respond the most to climate variability. In support of this prediction, data on species abundance at Konza over 16 years show that species that are more abundant in lowlands than uplands are preferentially reduced in years with low precipitation. Climate change is likely to also preferentially remove species from particular functional groups and clades. For example, warming is forecast to preferentially remove perennials over annuals as well as Cyperaceae species. Despite these predictions, climate change is unlikely to unilaterally alter the functional composition of the tallgrass prairie flora, as many functional traits such as physiological drought tolerance and maximum photosynthetic rates showed little relationship with climate envelope parameters. In all, although climatic drying would indirectly alter grassland productivity through species loss patterns, the insurance afforded by biodiversity to ecosystem function is likely to be sustained in the face of climate change.     

Plant community responses to bison reintroduction on the Northern Great Plains,United States: a test of the keystone species concept

Keystone species restoration, or the restoration of species whose effect on an ecosystem is much greater than their abundance would suggest, is a central justification for many wildlife reintroduction projects globally. Following restoration, plains bison (Bison bison L.) have been identified as a keystone species in the tallgrass prairie ecoregion, but we know of no research to document similar effects in the mixed‐grass prairie where restoration efforts are ongoing. This study addresses whether Northern Great Plains (NGP) mixed‐grass prairie plant communities exhibit traits consistent with four central keystone effects documented for bison in the tallgrass prairie. We collected species composition, diversity, abundance, bare ground cover, and plant height data in three treatments: where livestock (Bos taurus L.) continuously grazed, livestock were removed for 10 years, and bison have been introduced and resident for 10 years. We observed mixed support for bison acting as keystone species in this system. Supporting the keystone role of bison, we observed higher species richness and compositional heterogeneity (β‐diversity) in the bison treatment than either the livestock retention or livestock removal treatments. However, we observed comparable forb, bare ground, and plant height heterogeneity between bison‐restored sites and sites where livestock were retained, contradicting reported keystone effects in other systems. Our results suggest that after 10 years of being restored, bison partially fulfill their role as a keystone species in the mixed‐grass prairie, and we encourage continued long‐term data collection to evaluate their influence in the NGP.     

The importance of soil microorganisms for maintaining diverse plant communities in tallgrass prairie

? Premise of the study: According to the "Janzen-Connell hypothesis," soil microorganisms have the potential to increase plant community diversity by mediating negative feedback on plant growth. Evidence for such microbe-driven negative feedback has been found in a variety of terrestrial systems. However, it is currently unknown how general this phenomenon is within most plant communities. Also unknown is the role of mutualists in generating such feedback: do they decrease the influence of soil-mediated negative feedback on plant fitness or do they increase its effect by proliferating with plant hosts to which they give the least benefit? ? Methods: We investigated soil-microbe-mediated feedback via a series of reciprocal transplant experiments in the greenhouse using soil from a restored tallgrass prairie and native tallgrass prairie plant species. ? Key results: We found that negative feedback was very common but that mutualists (arbuscular mycorrhizal fungi) influence plant growth in opposition to the overall negative feedback trend. ? Conclusions: Widespread microbially mediated negative feedback indicates that plant community diversity and composition in tallgrass prairie are dependent on soil microorganisms. Native soil microorganisms should be considered in restoration efforts of tallgrass prairie and, potentially, other native plant communities.     

The role of hemiparasitic plants: influencing tallgrass prairie quality,diversity, and structure

Wood betony, Orobanchaceae (Pedicularis canadensis) and bastard toadflax, Santalaceae (Comandra umbellata) are two root‐hemiparasitic plant species found in tallgrass prairie communities. Natural resource managers are interested in utilizing these species as “pseudograzers” in grasslands to reduce competitively dominant grasses and thereby increase ecological diversity and quality in prairie restorations and urban plantings. We performed an observational field study at 5 tallgrass prairie sites to investigate the association of hemiparasite abundance with metrics of phylogenetic and ecological diversity, as well as floristic quality. Although no reduction in C₄ grasses was detected, there was a significant association between hemiparasite abundance and increased floristic quality at all 5 sites. Hemiparasite abundance and species richness were positively correlated at one restoration site. In a greenhouse mesocosm experiment, we investigated response to parasitism by P. canadensis in 6 species representing different plant functional groups of the tallgrass prairie. The annual legume partridge pea, Fabaceae (Chamaecrista fasciculata) had the greatest significant dry biomass reduction among 6 host species, but the C₄ grass big bluestem, Poaceae (Andropogon gerardii) had significantly greater aboveground biomass when grown with the hemiparasite. Overall, host species biomass as a total community was significantly reduced in mesocosms, consistent with other investigations that demonstrate influence on community structure by hemiparasitic plant species. Although hemiparasites were not acting as pseudograzers, they have the potential to influence community structure in grassland restorations and remnants.     

Butterfly, bee and forb community composition and cross-taxon incongruence in tallgrass prairie fragments

Pollinators provide an important class of ecological services for crop plants and native species in many ecosystems, including the tallgrass prairie, and their conservation is essential to sustaining prairie remnants. In Iowa these remnants are typically either block-shaped or long, linear strips along transportation routes. In this study we examined differences in the butterfly, bee, and forb community composition in linear and block prairie remnants, determined correlations between species diversity among butterflies, bees and forbs in the 20 prairie remnants sampled, and examined correlations of community similarity among butterflies, bees and forbs. Correspondence analysis showed that distinct communities exist for butterflies and forbs in block versus linear sites and discriminant analysis showed that the bee and forb communities in block and linear sites can be distinguished on the basis of a few species. Diversity of one group was a poor predictor of diversity in another, except for a significant inverse relationship between bees and butterflies. These two pollinator taxa may be responding very differently to microhabitat components within fragmented ecosystems. Our studies show that there need to be differences in conservation strategies for bees and butterflies to maintain both pollinator communities.     

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.     

Fire Frequency and Mosaic Burning Effects on a Tallgrass Prairie Ground Beetle Assemblage

Fire frequency has significant effects on the biota of tallgrass prairie, including mammals, vascular plants and birds. Recent concern has been expressed that widespread annual burning, sometimes in combination with heavy livestock grazing, negatively impacts the biota of remaining prairie remnants. A common management recommendation, intended to address this problem, is to create a landscape with a mosaic of different burn regimes. Pitfall trapping was used to investigate the impacts of fire pattern on the diversity and species composition of ground beetles (Coleoptera: Carabidae) at Konza Prairie Biological Station in eastern Kansas, USA. Trapping was conducted over three seasons in landscape units burned on average every 1, 4, or 20 years, and in a fourth season across the available range of vegetative structure to assess the variability of the community within the study system. In the fifth season communities were also followed immediately after two fire events to detect within-season effects of fire and to study short-term patterns of post-disturbance community assembly. Fire frequency had comparatively minimal effects on ground beetle diversity measures, and most numerically common species were observed widely across habitat and management types. Fire frequency effects were manifested primarily in changes in abundance of common species. Colonization of burned areas apparently did not occur from juxtaposed non-burned areas, but from underground or from long distances. While these results suggest that widespread annual burning of tallgrass prairie remnants may not have dramatic effects on prairie ground beetles, we urge caution regarding the application of these results to other taxa within tallgrass prairie.     

Assessing the Rate, Mechanisms, and Consequences of the Conversion of Tallgrass Prairie to Juniperus virginiana Forest

We assessed the determinants and consequences of the expansion of Juniperus virginiana L. (red cedar) populations into central US grasslands using historical aerial photos and field measurements of forest extent, tree growth, fire-induced mortality, and responses in herbaceous species diversity and productivity. Photos from northeast Kansas dating back to 1956 indicate that native tallgrass prairie can be converted to closed-canopy red cedar forest in as little as 40 years (a 2.3% increase in forest cover per year). Mean tree density in 21 forested sites ranged from 130 to 3500 trees/ha, with most sites at more than 800 trees/ha. In younger stands, maximum growth rates of individual red cedar trees exceeded 20 cm/y in height. Land management practices were critical to the establishment and growth of red cedar forest. Grazing reduced the fuel loads by more than 30% in tallgrass prairie. Based on measurements of mortality for more than 1800 red cedar trees, fire-induced mortality in grazed areas averaged 31.6% versus more than 90% at ungrazed sites. When tallgrass prairie was converted to red cedar forest, herbaceous species diversity and productivity were drastically reduced, and most grassland species were virtually eliminated. Consequently, community structure shifted from dominance by herbaceous C₄ species to evergreen woody C₃ species; this shift is likely to be accompanied by alterations in carbon storage and other ecosystem processes in a relatively short time period. Here we present a conceptual model that integrates the ecological and socioeconomic factors that underlie the conversion of grassland to red cedar forest. Received 29 August 2001; accepted 5 February 2002.     

Effects of urbanization on site occupancy and density of grassland birds in tallgrass prairie fragments

Tallgrass prairies are among the most threatened ecosystems in the world. Remaining prairies tend to be small and isolated and many are associated with urban and suburban landscapes. We asked how urbanization might impact the conservation value of tallgrass prairie fragments for grassland birds by comparing the densities and the probability of occurrence of Dickcissels (Spiza americana), Grasshopper Sparrows (Ammodramus savannarum), and Eastern Meadowlarks (Sturnella magna) across 28 grasslands surrounded by low, moderate, and high levels of urbanization. We employed a hierarchical model selection approach to ask how variables that describe the vegetation structure, size and shape of grasslands, and urbanization category might explain variation in density and occurrence over two breeding seasons. Occurrence of all three species was explained by a combination of vegetation and patch characteristics, though each species was influenced by different variables and only Eastern Meadowlark occurrence was explained by urbanization. Abundance of all three species was negatively impacted by urbanization, though vegetation variables were also prevalent in the best‐supported models. We found no evidence that vegetation structure or other patch characteristics varied in a systematic way across urbanization categories. Although our results suggest that grassland bird density declines with urbanization, urban tallgrass prairies still retain conservation value for grassland birds because of the limited availability of tallgrass prairie habitat and the limited impact of urbanization on species occurrence.     

Using Auchenorrhyncha (Insecta: Hemiptera) to develop a new insect index in measuring North American tallgrass prairie quality

Auchenorrhyncha (i.e., leafhoppers, treehoppers, spittlebugs, and planthoppers) represent some of the most diverse groups of herbivorous insects in the tallgrass prairie biome, they have close associations with many native prairie grasses and forbs, and respond in predictable ways to changes in native grassland degradation. These attributes make Auchenorrhyncha ideal candidates in the development of a habitat quality index to measure tallgrass prairie quality. In this study we propose the development of a species-based habitat quality index called the Auchenorrhyncha quality index or AQI as a useful method in tracking the condition of tallgrass prairie quality. The AQI is computed by summing six ecological characteristics (i.e., host plant specificity, voltinism, overwintering microhabitat, wing length, habitat fidelity, and origin) for each Auchenorrhynchan insect encountered, yielding coefficient of conservatism (CC) values that range from 0 (habitat generalist/tolerant to disturbance) to 18 (habitat specific/intolerant to disturbance). These CC values are averaged and combined with species richness producing un-weighted by abundance AQI (AQI_w/outN) and weighted by abundance AQI (AQI_w/N). The performance of the AQI was evaluated by examining the effects of sampling intensity on this index using a sweep net and a vacuum apparatus from 10 sites located on the three main North American tallgrass prairie communities, wet-mesic, sand, and loess hill. Scientists and land managers can adequately sample Auchenorrhyncha from four transects using a vacuum. Also, the highest AQI values were found from loess hill and sand prairies, indicating that conservation efforts should focus on these prairie communities. Additional applications of the AQI may include: (1) discriminating prairie quality at various spatial scales; (2) testing hypotheses about the effects of disturbance on prairie habitat (e.g. prescribed burning); (3) using the AQI as a model in developing habitat quality indices based on other diverse groups of grassland insects; and (4) the AQI has the capacity to be readily modified in assessing the quality of other biomes. Ultimately, the AQI should be used in combination with other habitat quality indices based on other diverse groups of organisms, such as plants and other insects, to provide a more complete assessment of native habitat quality.     

Drivers of Variation in Aboveground Net Primary Productivity and Plant Community Composition Differ Across a Broad Precipitation Gradient

Aboveground net primary production (ANPP) is a key integrator of C uptake and energy flow in many terrestrial ecosystems. As such, ecologists have long sought to understand the factors driving variation in this important ecosystem process. Although total annual precipitation has been shown to be a strong predictor of ANPP in grasslands across broad spatial scales, it is often a poor predictor at local scales. Here we examine the amount of variation in ANPP that can be explained by total annual precipitation versus precipitation during specific periods of the year (precipitation periods) and nutrient availability at three sites representing the major grassland types (shortgrass steppe, mixed-grass prairie, and tallgrass prairie) spanning the broad precipitation gradient of the U.S. Central Great Plains. Using observational data, we found that precipitation periods and nutrient availability were much stronger predictors of site-level ANPP than total annual precipitation. However, the specific nutrients and precipitation periods that best predicted ANPP differed among the three sites. These effects were mirrored experimentally at the shortgrass and tallgrass sites, with precipitation and nutrient availability co-limiting ANPP, but not at the mixed-grass site, where nutrient availability determined ANPP exclusive of precipitation effects. Dominant grasses drove the ANPP response to increased nutrient availability at all three sites. However, the relative responses of rare grasses and forbs were greater than those of the dominant grasses to experimental nutrient additions, thus potentially driving species turnover with chronic nutrient additions. This improved understanding of the factors driving variation in ANPP within ecosystems spanning the broad precipitation gradient of the Great Plains will aid predictions of alterations in ANPP under future global change scenarios.     

An assessment of diurnal water uptake in a mesic prairie: evidence for hydraulic lift?

Hydraulic lift, the passive movement of water through plant roots from wet to dry soil, is an important ecohydrological process in a wide range of water-limited ecosystems. This phenomenon may also alter plant functioning, growth, and survival in mesic grasslands, where soil moisture is spatially and temporally variable. Here, we monitored diurnal changes in the isotopic signature of soil and plant xylem water to assess (1) whether hydraulic lift occurs in woody and herbaceous tallgrass prairie species (Rhus glabra, Amorpha canescens, Vernonia baldwinii, and Andropogon gerardii), (2) if nocturnal transpiration or grazing by large ungulates limits hydraulic lift, and (3) if a dominant grass, A. gerardii, utilizes water lifted by other tallgrass prairie species. Broadly, the results shown here suggest that hydraulic lift does not appear to be widespread or common in this system, but isolated instances suggest that this process does occur within tallgrass prairie. The isolated instance of hydraulic lift did not vary by grazing treatment, nor did they result in facilitation for neighboring grasses. We suggest that the topographic complexity of this tallgrass prairie and the high rates of nocturnal transpiration observed in this study likely limit the frequency and occurrence of hydraulic lift. These results suggest that hydraulic lift can be a patchy process, particularly in heterogeneous landscapes.     

Biomass-diversity responses and spatial dependencies in disturbed tallgrass prairies

Monotonic, hump-shaped and zero-correlation productivity-diversity relationships have been found to date in many ecosystems. This diversity of responses has puzzled ecologists in their search for general principles on ecosystem functioning. Some state that the scale of observation is crucial in defining this relationship. We have developed a spatial model of tallgrass prairies where biomass and litter dynamics are defined by uncoupled difference equations. In this system, we periodically apply prescribed fire as a disturbance that propagates through neighboring cells. The model shows percolation thresholds at points where small-scale spatial heterogeneity and large-scale, global correlation coexist, resulting in power-law distributions in available areas for non-dominant species. These points maximize the biomass-diversity relationship. Our results suggest that spatial dependencies and the disturbance heterogeneity hypothesis are the cornerstone processes accounting for unimodality in productivity-diversity relationships.     

Experimental analysis of patch dynamics and community heterogeneity in tallgrass prairie

Effects of fire and small-scale soil disturbances on species richness, community heterogeneity, and microsuccession were investigated in a central Oklahoma tallgrass prairie. In the fall of 1985, 0.2 m² soil disturbances were created on burned and unburned tallgrass prairie. Vegetation on and off disturbances was sampled at monthly intervals over two growing seasons. During the first growing season, the cover of forbs and annuals, and species richness were significantly greater on versus off disturbances, but these differences did not persist through the second year. The variation in species composition among disturbed plots (heterogeneity) was significantly greater compared to undisturbed areas throughout the study. Fire had no consistent effect on richness and heterogeneity of vegetation on soil disturbances but fire reduced heterogeneity on undisturbed vegetation. Rate of succession, based on an increase in cumulative cover of perennial grasses over time, did not differ among treatments during the first growing season. During the second year, rate of succession was significantly greater on burned soil disturbances compared to unburned soil disturbances. These results suggest that while small-scale soil disturbances have primarily short-lived effects on grassland community structure, disturbances do help to maintain spatial and temporal variation in tallgrass prairie communities. Unlike in undisturbed vegetation, however, species richness and heterogeneity on soil disturbances were little effected by fire, but the rate of colonization onto disturbances appeared to be enhanced by fire.