Tallgrass prairies have some of the deepest and most fertile topsoil on earth. Widespread conversion of these grasslands to agriculture has decreased soil Carbon (C) storage by exacerbating erosion and disrupting aggregates that protect C from decomposition, coupled with lower plant C inputs. Thus, a primary goal of some grassland restorations is to improve soil structure and functioning. Conversion of cultivated systems to perennial grasslands often increases soil C, microbial biomass, and soil aggregate size and stability. A few studies have documented changes in soil microbial community structure after restoration. The objective of this work was two fold: 1) to explore the importance of soil texture and 2) determine plant diversity effects on recovery of soil biotic and abiotic properties. In the first study changes in soil microbial phospholipid fatty acid (PLFA) profiles and soil aggregates were examined in two 0-19 year chronosequences of restored grasslands in Nebraska on soils differing in texture, silty clay loam (SCL) and loamy fine sand (LFS), and compared them to native prairie. Soil was collected from the 0-10 cm soil depth at each site in May of 2007 and 2008. The SCL chronosequence exhibited increases in total PLFA biomass P<0.05, r2=0.29), PLFA richness (P<0.0001, r2=0.25), fungi (P<0.0001, r2=0.65), fungal:bacterial ratio (P<0.0001, r2=0.67), Gram (+) bacteria (P=0.02, r2=0.22), Gram (-) bacteria (P=0.05, r2=0.16), and actinomycetes (P=0.02, r2=0.23). Average soil aggregate diameter also increased (p=0.0002, r2=0.52). However, LFS sites showed no change across the chronosequence for any parameter. Total PLFA biomass (ANOVA, P<0.0001), richness (P<0.0001), and fungi (P=0.005) were greater on SCL restorations than LFS, but LFS had greater fungal:bacterial ratios (P=0.02). Soil microbial groups and soil aggregates were highly correlated, especially in the LFS choronosequence indicating that structural recovery is key to microbial community recovery. The second study investigated high diversity restorations with low diversity restorations on silty clay loam. In this study, high diversity and low diversity restorations in southeast Nebraska, aged 4 and 8 years were compared. The quantity of forbs seeded was too low and high diversity communities were a mixture of dominant C4 grasses (Andropogon gerardii Vitman, Schizachyrium scoparium (Michx.) Nash, Panicum virgatum L., Bouteloua curtipendula (Michx.) Torr. and Sorghastrum nutans (L.) Nash) and subdominant C3 grasses (Elymus canadensis L., Pascopyrum smithii (Rybd.) A. Löve, and Elymus virginicus L.). Eight year old plantings had greater root biomass, root C storage, root C:N ratio (P<0.05 for all), microbial biomass (low diversity only, PC<0.1, PN<0.05), PLFA richness (low diversity only, P<0.05), mycorrhizal fungi (P<0.05), and C mineralization (low diversity only, P<0.05) than 4 year old plantings. Low diversity plantings, which contained almost exclusively dominant C4 prairie grasses, had greater root C storage (P<0.1), mycorrhizal fungi (8 years only, P<0.1), and C mineralization (8 years only, P<0.05). Thus, C4 grasses and their associated arbuscular mycorrhizal fungi seem to drive recovery of soil C, soil respiration, and soil microbial communities over time. Overall, this work indicates that rates and success of belowground recovery are dependent on both abiotic and biotic factors in restoration. Restored plant communities affected soil recovery as dominant C4 grasses appeared to drive belowground recovery, but recovery depended on soil texture.

Woody vegetation encroachment has become a major threat to tallgrass prairie streams mainly because of fire suppression. This process converts prairie streams from open to closed canopy systems. The effects of these riparian changes are poorly understood, but the relative importance of basal resources presumably shifts from primarily autochthonous to allochthonous with increasing canopy cover, potentially altering macroinvertebrate functional structure and production. To assess the effects of woody vegetation encroachment on stream ecosystem structure and function, riparian trees were removed from two headwater stream reaches on the Konza Prairie Biological Station (KPBS) in eastern Kansas. Experimental stream reaches were compared to streams with naturally open and closed canopies before and after the manipulation. Benthic organic matter and macroinvertebrates were collected monthly from each reach for one year before and one year after woody vegetation removal. Total community production in canopy removal reaches ranged from 8.9-10.2 g AFDM m-2 y-1 before riparian removal, and this increased significantly to 13.4-14.5 g AFDM m-2 y-1 after riparian removal. Scraper production in canopy removal reaches was 2.8-3.9 g AFDM m-2 y-1 before riparian removal, and increased significantly to 6.0-8.7 g AFDM m-2 y-1 after riparian removal, presumably due to enhanced food availability. Total community production in naturally open reaches ranged from 7.6-12.6 g AFDM m-2 y-1 before riparian removal and decreased to 6.5-9.8 g AFDM m-2 y-1 after riparian removal. Riparian forest removal altered macroinvertebrate production and functional structure, but higher macroinvertebrate production in canopy removal reaches compared to naturally open reaches suggested natural conditions were not restored one year after riparian removal. However, macroinvertebrate communities in naturally open and canopy removal reaches became more similar after riparian removal. Functional structure, based on production, in naturally open and canopy removal reaches after riparian removal was dominated by scrapers (45-60% of total production), with similar proportions of collector-gatherers (12-26%) and predators (15-25%). Collector-filterers and shredders contributed < 9% of total production in naturally open and canopy removal reaches after riparian removal. Results demonstrate that woody vegetation encroachment and riparian forest removal significantly influence tallgrass prairie stream structure and function. Information from this study can help inform and guide management, restoration, and conservation of remaining tallgrass prairie streams.

Tallgrass prairie in North America has been severely degraded over the past century due to anthropogenic changes and is a subject of many restoration projects. Using these restoration projects, it is possible to examine potential drivers that influence community assembly. The environmental heterogeneity hypothesis provides a basis for enhanced diversity as function of resource partitioning and coexistence of potentially competing species. In essence, an area with higher levels of resource heterogeneity would be able to support a higher number of potentially competing species in contrast to an area with lower levels of resource heterogeneity (e.g. agricultural fields). The tallgrass prairie is naturally heterogeneous in abiotic resources such as soil depth and soil nitrogen, native prairie species both drive and exploit this heterogeneity and assemble a highly biodiverse community. Chapter 2 attempted to elucidate the effect of soil resource heterogeneity on plant community assembly, niche availability, and dimensionality. Chapter 3 attempted to examine the indirect influence of soil resources on aboveground Orthoptera herbivores. Both studies were conducted in a 16-year tallgrass prairie restoration experiment over a two-year period. There were no differences in plant community composition on a whole plot level. However, on a subplot level, shallow soil generally resulted in higher species richness and diversity. In contrast to previous studies, I found nitrogen addition increased forb richness and nitrogen reduction reduced forb diversity. As expected, the dominant grass Andropogon gerardii was positively influenced by high nitrogen regardless of soil depth. The multivariate analysis indicated the new species added to the experiment had unique trait spaces. Further analysis indicated niche availability and dimensionality were highest in treatments with nitrogen addition. This study suggests though fine scale spatial heterogeneity influences plant community composition, coarse scale spatial heterogeneity does not. This study also suggests that soil nitrogen may be a poor indicator of plant species diversity in the tallgrass prairie community. Orthoptera richness and biomass were higher in maximum heterogeneity treatments relative to control. The influence of high resource heterogeneity was highest on the richness of mixed-feeder grasshoppers and katydids. This effect, however, was inconsistent between years. Grass-feeder biomass was higher in the maximum heterogeneity treatment than control both years. This was attributed to nitrogen addition resulting in patches of higher quality forage in the maximum heterogeneity plots. Orthopterans are also influenced by the structural complexity of the plant community mediated by varying levels of soil resources. The maximum heterogeneity treatment contained higher variation in the cover and ANPP of a dominant grass, Andropogon gerardii. The positive relationship between plant richness and Orthoptera diversity suggests that maintaining plant richness in restored areas is important for maintaining diversity of higher trophic levels. The negative relationship between light interceptions and Orthoptera abundance suggests the dense vegetation from dominant tallgrass species may impede recruitment of some species. These results suggest suppression of dominant grasses can positively influence the plant community composition and Orthoptera herbivores. Understanding how soil resources influence plant community composition and higher trophic levels can aid our understanding of the community assembly process. Plant species benefited from higher variation in soil resources, particularly soil depth and soil nitrogen, while insect herbivores that depended on these plant species were also indirectly benefited. This study suggests soil heterogeneity is important for the assemblage of species on a multi-trophic level and this knowledge can assist land managers in restoration projects to achieve desired goals.

Master of Science / Department of Animal Sciences and Industry / K C Olson / The largest intact remnant of the tallgrass prairie, the Flint Hills ecoregion, is currently under threat from the invasive weed sericea lespedeza (Lespedeza cuneata [Dumont] G. Don; SL). The objectives of this research were to evaluate the efficacy of late-season prescribed burning and fall herbicide application, alone and in concert, for comprehensive control of sericea lespedeza and to assess their broader treatment impacts on native plant communities. A 31-ha native tallgrass pasture with a light to moderate infestation of SL was divided into 16 subunits for this experiment. Each subunit was randomly assigned to 1 of 4 treatments: negative control, spray-only, burn-only, or burn-plus-spray. A prescribed burn was conducted on burn-only and burn-plus-spray subunits in early September 2016. Following the re-emergence of SL, spray-only and burn-plus-spray subunits received a broadcast application of metsulfuron methyl (Escort XP, DuPont, Wilmington, DE) at a rate of 70.1 g ˖ ha⁻¹ in late September. Frequency and vigor of SL, total forage biomass, soil cover, and plant species composition were measured along permanent 100-m transects in each subunit prior to treatment application and again 12 mo later, in 2017 (i.e., 1 YAT). In 30 x 30-cm plots at 1-m intervals along each transect, the presence or absence of SL was noted. Where SL was present, crown maturity and maximum stem length of the SL plant nearest to the transect were recorded. Presence of multiple stems in plots was also recorded. Prior to treatment application, SL comprised 1 ± 2.0% of total basal cover and was not different between treatments (P = 0.38). One YAT, SL was more abundant (P ≤ 0.02) in negative control subunits than in spray-only, burn-only, or burn-plus-spray subunits, which were not different (P ≥ 0.95) from one another. Aerial frequency of SL, abundance of mature SL crowns, and incidence of plots with multiple SL stems were greatest (P ≤ 0.03) for negative controls, although not different (P ≥ 0.50) between the other 3 treatments. The change in forage biomass production 1 YAT did not differ (P = 0.16) between treatments. A tendency (P = 0.06) for a shift from litter cover to bare soil was noted when the spray-only, burn-only, and burn-plus-spray treatments were compared to the negative control. Graminoid basal cover was greater (P < 0.01) in the spray-only and burn-plus-spray treatments than in the negative-control and burn-only treatments 1 YAT. Conversely, forb basal cover was less (P = 0.01) in spray-only and burn-plus-spray treatments than in negative-control and burn-only treatments. The evenness component of diversity decreased in the burn-plus-spray treatment relative to the negative control (P ≤ 0.01). These data indicate that each of these strategies were effective in reducing SL populations. Although late-summer prescribed burning produced no detected negative responses within the native plant community, fall herbicide application, alone or in conjunction with prescribed burning, resulted in collateral damage to forb populations. A late-summer prescribed burn alone is recommended for low-cost comprehensive control of a light to moderate sericea lespedeza infestation.

The overall objective of this study was to quantify the effects of dominant grass propagule source (i.e., cultivar vs. non-cultivar) and seeded diversity of propagules on community structure and ecosystem function during prairie restoration. Two field experiments, and two chronosequences were used to investigate this main objective. The two field experiments were established at the same latitude separated by 620 km (corresponding to a precipitation gradient from eastern Kansas to western Illinois), and consisted of a split plot design, with dominant grass source as the whole-plot factor (2 levels) and seeded dominance of grasses as the subplot factor (5 levels). Percent cover of each species in each treatment combination was quantified during the first five years of restoration. Total plant species richness and diversity were not adversely affected by cultivars in Kansas or Illinois. The effect of the dominant grass population source on the cover of focal grasses, planted species, and volunteer species were contingent upon location. By the fifth year of restoration, diversity and richness were greatest, and cover of volunteer species was lowest in the low grass dominance (i.e., high diversity) treatment. ANPP, as well as total, microbial, and mineralizable pools of C and N were measured to quantify ecosystem function in these two field experiments. Changes in ecosystem function in Kansas and Illinois were primarily driven by time and regional abiotic differences, not propagule source or seeded diversity. The effect of plant species diversity on ecosystem function was further investigated at a landscape scale by developing and sampling two chronosequences of high (HDC; n=20) and low diversity (LDC; n=15) prairies spanning over two decades of restoration in northwestern Illinois. In general most metrics of ecosystem function in both chronosequences moved towards levels measured in remnant prairies. While the constituent prairies of the HDC had higher species richness, diversity, and more rapidly increasing root biomass than the fields of the LDC, recovery of other important ecosystem functions including aboveground net primary productivity, total, microbial, and mineralizable soil C, and soil aggregate mean weighted diameter were achieved equally well with either high or low diversity prairie plantings.

Doctor of Philosophy / Department of Geography / Douglas G. Goodin / Prescribed burning in tallgrass prairie affects a wide range of human and natural systems. Consequently, managing this biome based on sound science, and with the concerns of all stakeholders taken into account, requires a method for mapping burned areas. In order to devise such a method, many different spectral ranges and spectral indices were tested for their ability to differentiate burned from unburned areas at both the field and satellite scales. Those bands and/or indices that performed well, as well as two different classification techniques and two different satellite-based sensors, were tested in order to come up with the best combination of band/index, classification technique, and sensor for mapping burned areas in tallgrass prairie. The ideal method used both the red and near-infrared spectral regions, used imagery at a spatial resolution of at least 250 m, used satellite imagery with daily temporal resolution, and used pixel-based classification techniques rather than object-based techniques. Using this method, burned area maps were generated for the Flint Hills for every year from 2000-2010, creating a fire history of the region during that time period. These maps were compared to active fire and burned area products, and these products were found to underestimate burned areas in tallgrass prairie.

Doctor of Philosophy / Division of Biology / Anthony Joern / John Blair / Ecological communities are comprised both of species and their interactions. The importance of species interactions is embraced by ecological network analysis, a framework used to identify non-random patterns in species interactions, and the consequences of these patterns for maintaining species diversity. Here, I investigated environmental drivers of the structure of plant-pollinator and plant-herbivore networks. Specifically, I asked: (1) Do global-scale climate gradients shape mutualistic and antagonistic networks? (2) At a landscape scale (within a 3,487 ha research site), how do contrasting regimes of major grassland disturbances - fire frequency and grazing by bison (Bison bison) - shape plant-pollinator network structure? (3) How do fire and grazing affect plant-grasshopper network structure? And, (4) What is the role of plant species diversity in determining plant-herbivore network structure? At the global scale, variability in temperature was the key climatic factor regulating both antagonistic and mutualistic network structural properties. At the landscape scale, fire and grazing had major consequences for plant-pollinator and plant-herbivore communities. In particular, bison grazing increased network complexity and resistance to species loss for both plant-pollinator and plant-herbivore systems. Results from an experimental grassland restoration that manipulated plant diversity suggest that plant diversity directly affects plant-herbivore structure and increases network stability. Collectively, these results suggest that environmental gradients and plant species diversity regulate the network structure of ecological communities. Determining how the structure of ecological interactions change with environmental conditions and species diversity improves our ability to identify vulnerable communities, and to predict responses of biodiversity to global change.

Master of Science / Division of Biology / David A. Haukos / Sericea lespedeza (Lespedeza cuneata) is an invasive forb that reduces native grass and forb abundance in tall-grass prairie by up to 92%. Controlling invasions is difficult because traditional land management tools used in the Flint Hills, broad spectrum herbicides, spring prescribed fire, and cattle grazing, are ineffective against sericea. Recent research has demonstrated, however, that mid- and late summer prescribed fire and spring fire with early season grazing by steers followed by late season grazing by sheep are effective at reducing sericea whole plant mass, number of seeds produced, and seed mass. Field results were from two separate experiments conducted in tall-grass prairie study sites in the Flint Hills. On a Geary County, Kansas, study site, the utility of 1) spring fire (control), 2) mid-summer fire, and 3) late summer fire on sericea control were compared. On a Woodson County, Kansas, study site, the utility of 1) spring fire with early season steer grazing followed by rest (control) and 2) spring fire with early season steer grazing and late season sheep grazing on sericea control were compared. At the same study sites, I measured responses by the native wildlife community to use of summer fire and sheep grazing, relative to their controls, to manage sericea lespedeza. Specifically, my objectives were to compare grassland songbird density, grassland songbird nest survival, and grassland butterfly species composition and density among treatments at both study sites. I also related patterns in the vegetation community of each treatment for each study site to respective patterns in grassland bird and butterfly communities. Within study sites, density, nest density, and nest success of grassland bird communities responded similarly to treatments and controls, with the exception that densities of Grasshopper Sparrows (Ammodramus savanarrum) were 3.4- and 2.2-fold greater in mid- and late summer fire plots than spring fire plots, respectively, in the Geary County study site. Species compositions of butterfly communities were similar across treatments within experiments, but grassland specialist species comprised only 8.6 and 1.2% of all butterfly observations in the Geary County and Woodson County experiments, respectively. Grassland specialist butterfly species may benefit from summer fire, as their nectar sources were more abundant in Summer Fire plots than Spring Fire plots. Overall, within each experiment, grassland bird and butterfly communities were similar across treatments, suggesting that treatments did not negatively affect grassland songbird and butterfly communities. I additionally demonstrated that Dickcissel (Spiza americana) nest sites contain a lower proportion of sericea than random points, the first evidence that the invasion is detrimental to grassland songbird species. Lacking control, the continued sericea invasion will out compete cumulatively more forb plants resulting in declining quality of grassland bird nesting habitat on the landscape. Controlling sericea lespedeza invasions will allow native forb species to increase in abundance and improve the condition of grasslands for native wildlife and livestock producers. Therefore, I advocate use of summer fire or spring fire with a combination of cattle and sheep grazing to control sericea lespedeza with the long-term goal of tall-grass prairie restoration.

Eighty-three species of vascular plants were inventoried on the prairie relict during peak dickcissel nesting. Based on foliar cover and occurrence frequency, the five dominant plants were heath aster (Aster ericoides), eastern gammagrass (Tripsacum dactyloides), little bluestem (Schizachyrium scoparium), sensitive briar (Schrankia roemeriana) and meadow dropseed (Sporobolus asper). Sixty-one percent of dickcissel nests were constructed on or immediately next to three plant species: eastern gammagrass, sensitive briar and green milkweed.

Master of Science / Division of Biology / David A. Haukos / The regal fritillary (Speyeria idalia) was once an abundant butterfly species of North American prairie communities. Despite its once broad geographic distribution, populations have declined by ~99% in the prairie region for reasons that are poorly understood. The rapid, range-wide declines and persistent threats to extant populations from habitat loss and mismanagement prompted the U.S. Fish and Wildlife Service to initiate a species status review of the regal fritillary as a potential candidate for listing under the endangered species act in September 2015. Due to the uncertain status and contention regarding the effects of management practices (i.e., burning, grazing, and haying) on regal fritillary, my research objectives were to assess the effects of management practices and habitat features on the distribution and density of regal fritillary and their preferred larval host plant for the Midwest, prairie violet (Viola pedatifida). I generated species distribution models (SDM) of prairie violet to readily identify potential areas across the landscape containing patches of host plants and subsequently facilitate the location of regal fritillary larvae. The SDM produced maps of the probabilistic occurrence distribution of prairie violet throughout my study area and highlighted habitat features and management practices important to the occurrence of prairie violet. The seven final variables used to create the SDM and identified as important to the occurrence of prairie violet were elevation, slope, hillshade, slope position, land cover type, soil type, and average fire frequency. Using the SDM for prairie violet, I located eight areas to conduct surveys for regal fritillary larvae that were managed using various management (grazing and haying) regimes and fire-return intervals (low ≥ 10 years, moderate 3-5 years, and high 1-2 years). I used a binomial generalized linear model to determine the effects of management, host plant density, months since burn, and the interaction between months since burn and management on the occurrence distribution of regal fritillary larvae. My results indicate that greater host plant density and short fire-return intervals are important to the occurrence of regal fritillary larvae and, despite current management recommendations, larvae may be negatively impacted by a lack of fire. Finally, I surveyed tracts of prairie with my study area using a distance sampling approach along line transects stratified by overall management (burned, grazed, and hayed) and fire-return interval (low ≥10 years, moderate 3-5 years, and high 1-2 years) for adult regal fritillary. My results indicated that adult density was at least 84% greater in areas that received moderate fire-return intervals and greatest in areas that were grazed and burned on a moderate fire-return interval. However, density estimates of adult regal fritillary did not differ among overall management practices (i.e., burned grazed, hayed). Additionally, adult density increased as percent cover of grass, litter, and prairie violets increased. In contrast, adult density decreased as percent cover of woody vegetation and forbs increased. These results support the use of prescribed fire in a shifting mosaic or patch-burning practice as a viable management strategy for maintaining and conserving regal fritillary populations within the Flint Hills region.