Master of Science / Department of Biology / Kimberly A. With / Organisms exist within complex landscapes, and landscape features may influence multiple aspects of a species’ distribution within the landscape, including patch occupancy, abundance within patches, and population genetic diversity at a local or regional scale. We took two approaches to identify the relative importance of landscape context for populations of the Eastern Collared Lizard (Crotaphytus collaris collaris) in the northern Flint Hills of Kansas. First, we conducted surveys at limestone outcrops in experimental watersheds managed under different burning and grazing practices. Habitat occupancy and lizard abundance were estimated by constructing models that incorporated aspects of the environment at multiple scales. Both abundance and occupancy were higher on rock ledges that had more crevices, greater complexity in vegetation, covered a larger area, and were more prominent than available habitat. Abundance and occupancy were also higher in watersheds that were burned frequently (1–2 year intervals), but grazing only had a significant effect in less frequently burned (four–year burn interval) watersheds. Our second approach was to measure genetic diversity and population genetic differentiation and relate these measures to differences in landscape context. We sampled collared lizard DNA at four locations (sample sites < 45 km apart) and analyzed trends in 10 polymorphic microsatellite loci. We found evidence for low genetic variability and moderate population differentiation among our sample sites relative to estimates reported in the literature at the core of the species’ range. Differences in migration rates and ancestry among sampling locations also appear to correspond to differences in landscape resistance based on land cover and rock availability. Thus, it appears that habitat management may influence the suitability of habitat patches at the local scale, and that differences in land cover and rock availability may influence the connectivity of populations at the landscape scale.

Doctor of Philosophy / Department of Biology / Walter K. Dodds / Streams exist in a state of dynamic equilibrium with frequent floods and drought. The frequency and intensity of stream disturbances are projected to increase with greater water withdrawal for agriculture and biofuel production, watershed development, and altered climate. Changes in the hydrologic regime may alter stream ecosystems. I studied how stream communities return after disturbances and how nutrients, consumers, and substrata heterogeneity influence recovery trajectories. Large consumers were excluded from pools following a severe drought to assess how community structure and function returned in their absence. Large consumers reduced algal biomass, primary productivity, and nutrient uptake rates, and delayed macroinvertebrate recolonization. However, grazer effects were temporary and their influence weakened after five weeks. In a second experiment, I assessed the relative influence of grazer density and nutrient loadings on algal recovery from flood. Nutrients had a stronger effect on recovery than grazers, but the strength of each varied temporally. Grazer control decreased and nutrient control increased over time. A third experiment addressed the physical properties of stream substrata on algal development. The relationship among algal accumulation and substrata surface topography was assessed by growing algae on substrata with varying orientation and roughness. Total algal biomass decreased on surfaces with angles > 45 degrees, and peaked at an intermediate roughness (pit depth of [similar to]17 [Mu]m). Rougher surfaces collected more tightly attached (grazer resistant) forms and less loosely attached (grazer susceptible) forms. Individual algal forms responded differently to grazing pressure, nutrient availability, and surface features. I developed a method using Fourier-transform infrared microspectroscopy to measure single-cell physiological responses in benthic algae. Nutrients and consumers were strong regulators of ecosystem succession following disturbance, but nutrient influence was stronger. The influence of nutrients and consumers were context dependent, and changed over the course of recovery. Rougher surfaces increase algal growth and shifted algal assemblages to more grazer resistant forms, which may decrease the influence of large consumers on stream function. Altering the severity and frequency of disturbances can change the trajectory of stream recovery and ultimately change community composition and stream metabolic activity, which may alter ecosystem services such as water purification and recreation.

Doctor of Philosophy / Department of Biology / Jack F. Cully, Jr. / Black-tailed prairie dogs (Cynomys ludovicianus) are epizootic hosts for plague (Yersinia pestis); however, alternate enzootic hosts are important for the maintenance of the pathogen. We determined small rodents and prairie dog associations and quantified rodent and flea relationships in the presence and absence of prairie dog colonies and plague. We identified potential alternate hosts and flea vectors for the maintenance and transmission of plague in the prairie ecosystem. This is the first multi-year study to investigate associations between prairie dogs, rodents and fleas across the range of the black-tailed prairie dog. Few rodent species associated with black-tailed prairie dogs and were found to be highly abundant on colonies. Rodent species implicated in plague were present at study areas with and without plague. Peromyscus maniculatus and Onychomus leucogaster, two widely occurring species, were more abundant in areas with a recent history of plague. Flea community characteristics varied within each study area in the presence and absence of prairie dogs. Based on flea diversity on rodents, and the role of rodents and fleas in plague, we identified P. maniculatus and O. leucogaster and their associated fleas, Aetheca wagneri, Malareus telchinus, Orchopeas leucopus, Peromyscopsylla hesperomys, and Pleochaetis exilis to be important for the dynamics of sylvatic plague in our study areas. Peromyscus maniculatus and O. leucogaster were consistently infected with Bartonella spp., another blood parasite. Presence of prairie dog fleas on other rodents at both off and on prairie dog colonies suggests the potential for intra and interspecific transmission of fleas between rodent hosts, and between other small rodents and prairie dogs.

Doctor of Philosophy / Department of Biology / David C. Hartnett / In perennial grasslands, the belowground population of meristems (the bud bank) plays a fundamental role in local plant population structure and dynamics. I tested the “meristem limitation hypothesis” prediction that bud banks increase along an increasing precipitation/productivity gradient in North American grasslands. I sampled bud populations quarterly at six sites across a 1,100 km gradient in central North America. Bud banks increased with average annual precipitation, which explained 80% of variability in bud banks among sites. Seasonal changes in grass bud banks were surprisingly similar across a 2.5-fold range in precipitation and a 4-fold range of aboveground net primary productivity (ANPP). Secondly, I tested the hypothesis that tallgrass prairie plants respond to increases in a limiting resource (nitrogen) through demographic effects on the bud bank. I parameterized matrix models for individual genets, considering each genet as a population of plant parts (buds and stems). Nitrogen addition significantly impacted bud bank demography of both Sporobolus heterolepis and Koeleria macrantha. In 2005, emergence from the bud bank and growth rates (λ) of the tiller population were significantly higher in S. heterolepis genets that received nitrogen. In contrast, nitrogen addition decreased λ in K. macrantha. Both prospective and retrospective analyses indicated that bud bank dynamics are the key demographic processes driving genet responses to nutrient availability. Lastly, I tested the hypothesis that the effects of fire and grazing on plant species composition and ANPP are mediated principally through demographic effects on bud banks. I found that plants respond to fire and grazing with altered rates of belowground bud natality, bud emergence, and both short-term (fire cycle) and long-term changes in bud density. The size of the bud bank is an excellent predictor of long-term ANPP, supporting my hypothesis that ANPP is strongly regulated by belowground demographic processes. Meristem limitation due to water or nutrient availability or management practices such as fire and grazing may constrain grassland responses to inter-annual 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 phenomena such as nutrient enrichment, and may be most resistant to exotic species invasions.

Master of Science / Department of Biology / Brett K. Sandercock / Translocations are a useful management tool for restoring wildlife species to their native ranges, but require post-release monitoring to determine project success. We report results of a 4-year effort to reestablish a breeding population of Evermann's Rock Ptarmigan (Lagopus mutus evermanni) on Agattu Island in the Aleutian Archipelago, Alaska. This endemic subspecies of ptarmigan was extirpated from most of the Near Islands by introductions of arctic fox by fur traders, and natural recolonizations did not occur after fox eradication. All females surviving the 2-week post-release period attempted to nest but initiated clutches later in the season and laid fewer eggs than resident females. Nest success was similar for resident and translocated females. Brood survival was greater for translocated than resident females and differed significantly; however, brood survival varied among years and was reduced by adverse weather conditions in 2006. Seasonal survival of radio-marked birds during the breeding season was 100% for translocated and resident ptarmigan. Over-winter mortality resulted in a low return rate in 2006. Returning birds in 2006 showed strong site fidelity and nest locations in two consecutive years were closely spaced. We examined nest site selection and determined brood movements and home range size of recently translocated and resident females. Nest sites of translocated females averaged 4.2 km from their respective release location and were not different from nest locations of resident females with regard to topographical features. Female nest site selection was influenced by percent composition of rock and forb coverage but was unaffected by slope, aspect, or general habitat. Broods of both resident and translocated females made movements to higher elevations after hatching. While size of brood home range was similar for resident and translocated females, distances traveled between the nest site and the arithmetic center of the brood home range were greater for translocated females. Overall, we conclude that translocations are an effective technique for reestablishing island populations of Rock Ptarmigan. Our study provides successful methods which may benefit future projects to reestablish endemic populations of ptarmigan and landbirds elsewhere in the Aleutian Islands.

Master of Science / Department of Biology / David C. Hartnett / Herbivory is one of the major biotic interactions shaping the structure and dynamics of grassland plant populations and community structure. The two major grass growth forms, rhizomatous and caespitose species, may show different grazing tolerance and short-term overcompensation may offset long-term reductions in plant performance and fitness. The objectives of this study were to assess 1) the effects of long-term ungulate grazing on plant architecture, population structure, and life history traits of the caespitose perennial grass, Schizachyrium scoparium (little bluestem), and the rhizomatous Bouteloua curtipendula (sideoats grama) in tallgrass prairie, and 2) the effects of grazing intensity (frequency of defoliation) on growth responses of little bluestem. Long-term bison grazing decreased the cover, frequency, tiller height, and proportion of tillers producing seed in little bluestem, but caused no changes in tiller density and total genet size. Grazed little bluestem plants maintained a significantly larger below ground bud bank. Bison grazing had no long-term effect on the abundance, bud bank densities, or rhizome growth of the rhizomatous side-oats grama grass. Biomass, tiller density, relative growth rates, and proportion of tillers flowering in little bluestem all decreased with increasing frequency of defoliation. However, even an intense grazing regime (9 defoliations over a 12 month period) caused no plant mortality and no changes in new tiller emergence rates, or bud bank densities. Increasing defoliation frequency did result in shifts in plant architecture, as an increasing proportion of extravaginal tillers led to a more lateral, decumbent growth form. These results show that the rhizomatous side-oats grama grass has a significantly higher grazing tolerance than does little bluestem, and/or that bison selectively graze little bluestem. Results from responses to the experimental defoliation regimes suggest that little bluestem shows lower tolerance to high frequency of defoliation, and its maintenance of a reserve below ground bud bank may be its primary tolerance mechanism, allowing tiller populations to recover following severe defoliation.

Doctor of Philosophy / Department of Biology / Anthony Joern / Aboveground and belowground food webs are linked by plants, but their reciprocal influences are seldom studied. Because phosphorus (P) is the primary nutrient associated with arbuscular mycorrhizal (AM) symbiosis, and evidence suggests it may be more limiting than nitrogen (N) for some insect herbivores, assessing carbon (C):N:P stoichiometry will enhance my ability to discern trophic interactions. The objective of this research was to investigate functional linkages between aboveground and belowground invertebrate populations and communities and to identify potential mechanisms regulating these interactions using a C:N:P stoichiometric framework. Specifically, I examine (1) long-term grasshopper community responses to three large-scale drivers of grassland ecosystem dynamics, (2) food selection by the mixed-feeding grasshopper Melanoplus bivittatus, (3) the mechanisms for nutrient regulation by M. bivittatus, (4) food selection by fungivorous Collembola, and (5) the effects of C:N:P on invertebrate community composition and aboveground-belowground food web linkages. In my analysis of grasshopper community responses to fire, bison grazing, and weather over 25 years, I found that all three drivers affected grasshopper community dynamics, most likely acting indirectly through effects on plant community structure, composition and nutritional quality. In a field study, the diet of M. bivittatus was dominated by forbs with grasses constituting only a minor fraction of their diet under ambient soil conditions, but grass consumption approximately doubled as a result of changes in grass C:N:P. M. bivittatus was found to rely primarily on selective consumption of foods with varying nutritional quality, rather than compensatory feeding or altering post-ingestive processes, to maintain C:N homeostasis in a laboratory experiment. In a soil-based mesocosm study, I show that Collembola feed on both saprophytic and AM fungi, in some cases exhibiting a slight preference for AM fungi. In the final study, although I did not find the expected indirect relationship between soil Collembola and aboveground herbivory as mediated through host plant quality, there were significant effects of root C:N and AM colonization on Collembola density and of plant C:N on aboveground herbivory. Overall, this research shows that host plant C:N:P stoichiometry can influence both above- and belowground invertebrate population, community, and food web dynamics.

Master of Science / Department of Biology / Keith B. Gido / Stream networks provide complex habitats for fish assemblages that can vary gradually along a gradient of stream size or abruptly at transition zones between large rivers and their tributaries. We evaluated the relative importance of these gradual and abrupt habitat transitions in regulating stream fish assemblages by quantifying roles of stream size and spatial position within a drainage network as a determinant of fish assemblage structure within the Kansas River basin, KS. We predicted fish assemblage structure to generally be dependent on stream size and that smaller streams would be influenced by their connectance to larger mainstem rivers. Fishes in the Kansas River basin varied along a gradient of stream size and longitude, and after controlling for these effects, there was evidence that connectivity to a larger river influenced species richness and assemblage structure. In 1st order streams there was an increase in species richness with increasing distance from a mainstem confluence and species composition in larger tributaries (i.e., 4th order streams) varied with proximity to the mainstem river. We also found an increase in species richness at sites located on smaller tributaries connected to a larger downstream mainstem. Species composition in 1st and 4th order streams also varied with connectance to the mainstem river. Within three intensively sampled tributaries, there was an abrupt change in fish fauna between the Kansas River and sample sites above the confluence, but only gradual change in assemblage structure within each tributary with a high degree of seasonal variation. In the first 20 stream km of these three mainstem tributaries adult fishes were more structured along a gradient away from the mainstem river than juveniles, potentially suggesting more generalized habitat needs of juvenile fishes. At the spatial and temporal scale of our analysis, it appeared the effects of large rivers on tributary streams were generally localized. However, the documented influence of spatial position suggests movements between habitats could regulate community level dynamics as well as individual species over longer temporal scales.

Master of Science / Department of Biology / Brett K. Sandercock / Monitoring programs indicate that numerous shorebird populations are subject to on-going declines. The U.S. Shorebird Conservation Plan lists twenty-seven shorebird species as species of high concern and seven as highly imperiled, including the Buff-breasted Sandpiper (Tryngites subruficollis). One hypothesis for ongoing population declines is exposure to toxic chemicals and pollutants. The purpose of this project was to characterize plasma cholinesterases (ChEs) of migratory shorebirds and address potential exposure to organophosphorus (OP) and carbamate (CB) pesticides. Consumption or contact with these pesticides can cause mortality and a variety of sub-lethal effects. Buff-breasted Sandpipers and other upland shorebirds are particularly likely to encounter agrochemicals due to their habitat use at the non-breeding grounds. I sampled migratory shorebirds over three seasons, during north- and southbound migration in 2006 and 2007 in Texas, Kansas, and Nebraska and during the non-breeding season in 2007 in Argentina, Uruguay, and Paraguay. I collected blood samples and footwashings from reference sites, where OP and CB pesticides were not used, and agricultural sites, where these two insecticides were recommended for control of crop pests. I assessed several variables known to affect plasma ChE activity including body size, date of capture, time of capture, condition, sex, and region. Small-bodied species had higher levels of ChE activity in plasma than large-bodied species. Plasma ChE activities varied with date of capture in 3 of 5 species sampled in North America. Sex differences were significant in 1 of 4 species tested. Plasma acetylcholinesterase (AChE) activity was higher among White-rumped Sandpipers sampled in North America but there was no difference between regions among Buff-breasted Sandpipers. Time of capture and individual condition did not affect plasma ChE activity. Estimates of exposure to ChE inhibitors were addressed in five species. Plasma AChE and butyrylcholinesterase (BChE) activities of Buff-breasted Sandpipers were lower at agricultural sites in South America but BChE activity was higher at agricultural sites in North America. There were no differences between sites in four other species tested. A meta-analysis across all species indicated that in 4 of 6 comparisons habitat type had a negative effect on AChE activity consistent with exposure to ChE inhibitors but there was a regional positive effect of agricultural habitat on BChE activity in North America. Comparison of body mass between sites suggested that use of habitats with potential pesticide application did not affect mass gain. Project results suggest that 1 of 5 shorebird species tested was exposed to ChE-inhibiting pesticides at the non-breeding grounds and future monitoring is necessary to assess potential effects at the population level. This study highlights the importance of complete sampling and addressing variability in plasma ChEs before making estimates of exposure to OP and CB pesticides. It provides the first estimates of migratory shorebird exposure to OP and CB pesticides, a potential conservation issue. Future research should include continued monitoring of Buff-breasted Sandpiper ChE levels and habitat use. Other sources of anthropogenic declines such as habitat loss and illegal hunting should be investigated for species that did not show evidence of exposure.

Doctor of Philosophy / Department of Biology / John M. Blair / Approximately 96% of native tallgrass prairie in North America has been lost, which accentuates the need for effective methods to restore the structure and function of these degraded ecosystems. Many prairie restorations aim to restore grass and forb species in proportions reflecting plant species diversity in native prairie. A target grass-forb species mixture is typically chosen at the onset of restoration, but often, grasses become excessively dominant and forbs are underrepresented as the community develops. Several studies have examined the potential for increasing forb cover and diversity in newly restored grasslands, but few studies have assessed factors limiting forb cover and diversity in well-established grass-dominated prairie restorations. The primary objective of this research was to assess the potential for enhancing plant species diversity and productivity in an established grass-dominated prairie restoration by selective removals of dominant grass species, and by manipulating resources (soil nutrients, light availability) or mycorrhizal interactions. A 7-year old grass-dominated restoration was used to evaluate plant and soil responses to manipulations in three separate studies. The first study examined the potential suppressive effects of dominant grasses on plant diversity by reducing the cover and biomass of two dominant grass species, Andropogon gerardii and Panicum virgatum. After 3 years, the removal of A. gerardii increased species richness and diversity, which was correlated with increased light availability, but not changes in soil resources. The second study examined the responses of restored grassland communities to long-term manipulation of soil resources (nutrient availability or soil depth), and to aboveground biomass removal via mowing. The long-term manipulation of soil resources did not alter plant species diversity, but nitrogen and light availability were important factors regulating plant productivity. The third study assessed the effects of manipulating arbuscular mycorrhizal (AM) fungi, through the use of either commercial inoculum or fungicide, on plant communities in restored prairie. Mycorrhizal suppression reduced grass productivity, suggesting that fungicide may be useful for enhancing diversity of restored prairies that are dominated by obligate mycotrophic grasses. In total, these studies suggest that competition between dominant grasses and subordinate forbs limits plant diversity in restored tallgrass prairie.