A decline in biodiversity in agricultural landscapes has been reported for decades. One of the most important habitats for biodiversity in agricultural landscapes is semi-natural grasslands. To preserve species-rich semi-natural grasslands, research need to focus both on local habitat quality and on landscape composition at various scales. In the current study I examined how nectar sources, host plants and other environmental parameters affected the occurrence of adults of four burnet moth species, residents of semi-natural grasslands. Also, I investigated the effects of landscape composition at 34 spatial scales. The results in general showed positive effects of host plants cover, nectar sources abundance, dry soil, sun exposure, tall sward height, small herb and grass cover. At the landscape level all the species responded negatively to the amount of arable land and positively to the amount of forests at scales up to 10000 m. Two species were negatively affected by the amount of artificial surfaces and pasture at some scales. Though the amount of semi-natural grasslands and patch area did not affect the occurrence of the species, at the local scale semi-natural grasslands were still important for the species. Thus, my results suggest that management should be focused both on local and landscape levels. Focus should be to preserve sunny and open areas of high-quality semi-natural grasslands rich in burnet moths’ host plants and especially nectar sources that seem to be more important than host plants for adults. Semi-natural grasslands should be preferably adjacent to forests and not arable land.

Coastal Sage Scrub (CSS) is a low scrubland plant community native to the coasts of California, housing many threatened and endangered species. Due to the invasion of non-native plants, many areas of CSS have type converted to annual grasslands and the fire frequency has accelerated; fire in turn, may facilitate further invasion, leading to a loss of biodiversity. While many studies document post-fire succession in these communities, pre-fire data are rarely available for comparison, especially data on seedling emergence. I analyzed post-fire recovery of a type-converted grassland community, comparing seedling emergence data for the first and third year after fire to the three years preceding the fire. Non-native species abundances declined more after the fire than did native abundances. This pattern was still present in 2015, three years post-fire. Two native species, Amsinckia menziesii var. intermedia (Amsinckia) and Phacelia distans (Phacelia), were subjects of seed addition treatments pre-fire, but I found no evidence that past seeding increased their abundances post-fire. Amsincki did recover to its pre-fire density three years after the fire, while the density of Phacelia declined over 75% in both the year immediately post-fire and three years after the fire. However, a third native species, Lupinus bicolor (Lupinus), was both much more abundant and also more spatially widespread both immediately after the fire and two years later. This supports the hypothesis that Lupinus is stored in the soil seed bank and the fire may have given this species the opportunity to recover by lowering abundances of non-native competitors. This analysis will inform future conservation efforts by improving our understanding of how seed banks impact the post-fire recovery of native species.

We reconstructed historical fire regimes of montane forest-grassland ecotones in the ~40,000 ha Valles Caldera National Preserve, New Mexico. We used a targeted approach to sample ancient fire-scarred trees along the ecotone, and compared variations in historical fire occurrence within and among valles in the grassland-forest. The resulting tree-ring record extends from 1240-2008 C.E., comprised of 2,443 fire scars from 330 trees representing 238 fire years during the period of analysis, 1601-1902 C.E. Our results confirm pre-1900 historical occurrence of high-frequency, low-severity surface fires over multiple centuries in the ecotone. Mean fire intervals for all fires were 5.5-22.5 years (~6-123 ha) at individual sites, 2.7-10 years (~67-4955 ha) in individual valles, and 1.6 years (~10 386 ha) across the landscape. Synchronous fires burned extensively and occurred at ~10 year intervals during years with significantly low PDSI. Results will be useful in planning forest/grassland restoration actions and reinstituting fire regimes.

In order to predict how terrestrial ecosystems will respond to global change, there is growing recognition that we need to better understand linkages between plant and soil processes. Previously the factors and processes with potential to influence the terrestrial carbon (C) cycle have been investigated in isolation from each other. This study investigated the interactions of nutrient availability and warming in controlling the soil carbon dynamics, with regards to the fate of already sequestered carbon in soil, under conditions of increasing atmospheric temperatures. The project objectives were addressed by three independent experiments designed to explain specific components of the carbon-nutrient cycle interactions, and the findings brought together to describe the implications for future soil carbon storage. The main measurements collected throughout this project included soil carbon dioxide (CO2) fluxes, partitioned into autotrophic and heterotrophic components, net ecosystem exchange and respiration fluxes, and background soil moisture and temperature data, backed by gas, soil and biomass analyses. In the two field experiments, these measurements were taken from plots with or without any inorganic nutrient additions or in the presence or absence of legumes providing biological nitrogen addition to the ecosystem. In the laboratory, temperature and nutrient availability were manipulated within the ecosystem. The reduction in decomposition rates, without reduction of productivity as a result of inorganic nutrient additions, indicated the potential for increasing C storage. There was also evidence that nutrient availability controls the strength of the link between plant and soil processes in semi-natural grasslands. The yields, decomposition rates and soil C fluxes recorded in the presence and absence of legumes provided some evidence of N2 fixation, improving ecosystem productivity and soil properties while reducing soil C effluxes, in a managed grassland. In the laboratory, the warming of soils from lysimeters with and without plants, receiving or not receiving fertiliser, supported the findings from field experiments regarding the importance of the soil-plant link in controlling C fluxes. However, C stocks and δ13C analyses showed that over a year’s worth of warming and nutrient manipulations made little difference to the amount of C stored in the soil, indicating that edaphic factors have greater control over the response of C dynamics to increased temperatures.

Grazing management is very popular nowadays and number of sites with grazing animals with the aim of restoring the sites is steadily growing. All the effects of grazing on grazed grasslands are not elucidated yet. This thesis focused mainly on the contribution of zoochory to restoration of species rich grasslands on stands cleared from Robinia pseudoacacia and stands cleared from Prunus spinosa brushwood. For the purpose of monitoring changes in vegetation, permanent plots have been established. In order to identify sources of new species occurring in permanent plots, inventory of species growing in neighbourhood of the permanent plots has been done, samples of soil seed bank, sheep buttons and seeds from sheep wool have been germinated in a greenhouse. Monitoring of permanent plots showed reduced regrowth of R. pseudoacacia and P. spinosa. We have also found that greater changes in species composition occurred in more degraded stands than in stands better-preserved. Germinating experiments proved soil seed bank being mainly the image of aboveground vegetation with minor importance to restoration of species rich grasslands. On the other hand sheep seem to be of great use for dispersal of seeds both by epizoochory and endozoochory when walking between different stands. This finding is of great importance for...

Master of Science / Department of Geography / Douglas Goodin / Hyperspectral data has been widely used for estimates of canopy biochemical content over the past decades. Most of these studies were conducted in forests or crops with relatively uniform canopies. Feasibility of the use of hyperspectral analysis in heterogeneous canopies with diverse plant species and canopy structures remains uncertain. Spectral data at the canopy level, with mixed background noise, canopy biochemical and biophysical properties create more problems in spectral analysis than that at the leaf level. Complications of heterogeneous canopies make biochemical retrieval through remote sensing even more difficult due to more uneven spatial distribution of biochemical constituents. The objective of my research was to map canopy nitrogen content in tallgrass prairie with mixed canopies by means of hyperspectral data from in-situ and airborne measurements. Research efforts were divided into three steps: (1) the green leaf area index (LAI) retrieval, given LAI is an important parameter in scaling nitrogen content from leaves to canopies; (2) canopy nitrogen modeling from analysis of in-situ hyperspectral data; and (3) canopy nitrogen mapping based on aerial hyperspectral imagery. Research results revealed that a fine chlorophyll absorption feature in the green-yellow region at wavelengths of 562 – 600 nm was sensitive to canopy nitrogen status. Specific spectral features from the normalized spectral data by the first derivative or continuum removal in this narrow spectral region could be selected by multivariate regression for nitrogen modeling. The optimal nitrogen models with high predictive accuracy measured as low values of root-mean-square error (RMSE) were applied to the aerial hyperspectral imagery for canopy nitrogen mapping during the growth seasons from May to September. These maps would be of great value in studies on the interactions between canopy vegetation quality and grazing patterns of large herbivores in tallgrass prairie.

Doctor of Philosophy / Division of Biology / Walter K. Dodds / Tallgrass prairie evolved with fire-grazer interactions. Fire and grazing are vital processes for maintaining grasslands and cattle production, and therefore will be continued as land management schemes. The effects of fire and grazers on prairie streams are understudied, but may significantly influence stream ecology. This dissertation examined how prescribed burning, bison grazing, and patch-burn grazing (by cattle) influence water quality, stream biota, and riparian amphibians and reptiles at Konza Prairie, Kansas, or Osage Prairie, Missouri. Using Global Positioning System, we monitored bison and cattle distribution throughout watersheds. The immediate effects of prescribed burning were examined at both Konza and Osage Prairies. The impacts of bison on water quality were determined by using a long-term dataset from Konza Prairie and compared watersheds with and without bison. Amphibian and reptile assemblages were monitored for two years at Osage, and assemblage data were analyzed using redundancy analysis, permuted analysis of variance, and occupancy modeling. A patch-burn grazing experiment occurred for 5 years at Osage (2 years pretreatment data and 3 years of treatments) and was analyzed using a before-after, control-impact design. Prescribed burning had minimal effects on water chemistry. At Konza Prairie, bison did not alter water quality likely because they spent negligible time (<5%) in streams. Contrarily, cattle at Osage Prairie significantly increased stream concentrations of total suspended solids, nutrients, Escherichia coli bacteria, algal biomass, and primary production. Unlike bison, cattle spent significant time (~21%) in streams if allowed access to riparian zones. In watersheds with cattle excluded from streams by riparian fencing, water quality contaminant concentrations increased significantly, but not to the magnitude of unfenced streams. Amphibian abundance and richness were not different among patch types; instead, they were restricted to specific basins. However, reptiles displayed preference for certain patch-types, and had the highest abundance and richness in watersheds with fire and grazing. These results have implications for natural resource management. Riparian fencing of cattle may be a useful practice in areas where water resource protection is the priority. However, overland flow may alter water quality in watersheds with grazers despite fencing. Land managers will need to define management objectives and accept trade-offs in water quality, amphibian and reptile habitat, and cattle production.

Master of Science / Biology / Alice Boyle / Dispersal is a behavior common to virtually all taxa with important consequences for gene flow, demography, and conservation. Mobile animals such as birds frequently engage in breeding dispersal, but the factors shaping this behavior are not well understood. In mid-continental grasslands, preliminary evidence suggested that Grasshopper Sparrows (Ammodramus savannarum) engaged in breeding dispersal within a single breeding season. This intriguing pattern shaped my research questions: (1) what are the patterns of within-season breeding dispersal in Grasshopper Sparrows? (2) why do some individual sparrows disperse, and others do not? and (3) what shapes settlement decisions following dispersal? I tested hypotheses based on spatial and temporal variation in nest predation, nest parasitism, and food availability. I studied Grasshopper Sparrows at 18 experimentally-managed watersheds with different fire and grazing regimes at Konza Prairie Biological Station during 2013-2015. To describe patterns, I combined re-sighting of 779 marked individuals, territory mapping, and radio-telemetry to quantify territory densities, turnover, and dispersal distances. To determine if nest predation or nest parasitism affected dispersal, I monitored the fate of 223 nests. I estimated food intake rates using plasma metabolites, and prey availability using sweep sampling. Densities of territorial Grasshopper Sparrows varied seasonally in management-specific ways. Turnover was remarkably high, with over half of territorial males being replaced each month. Over a third of males changed territories within-season, dispersing 0.1-9 km between breeding attempts. Dispersal decisions were related to past nest predation, but not nest parasitism. Dispersal likely yields fitness benefits, as sparrows that dispersed increased their chances of nest survival by 23% relative to site-faithful individuals. However, food availability did not affect settlement decisions. My study provides the first evidence of within-season breeding dispersal in Grasshopper Sparrows, and represents one of few tests of alternative hypotheses explaining dispersal decisions of songbirds. My results are consistent with a growing literature on the role of predation shaping dispersal, but suggest, somewhat surprisingly, that food is not important in post-dispersal habitat selection. High dispersal capacity coupled with adaptability to temporal and spatial change may be typical of grassland songbirds, implying that demographic studies and management decisions must consider their mobility for conservation.

Thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy. School of Animal, Plant and Environmental Science, University of the Witwatersrand, Johannesburg, South Africa. October 2016. / Mining has negative impacts on the environment in many different ways. One method developed to quantify some of these impacts is Landscape Function Analysis (LFA) and this has been accepted by some mining companies and regulators. In brief, LFA aims at quantifying the organization of vegetative and landscape components in a landscape into patches along a transect and quantifying, in a relative manner, three basic processes important to landscape functioning, namely: soil stability or susceptibility to erosion, infiltration or runoff, and nutrient cycling or organic matter decomposition. However, LFA is limited in large heterogeneous environments, such as those around mining operations, due to its localized nature, and the man hours required to collect a representative set of measurements for such large and complex environments. Remote sensing using satellite-acquired data can overcome these limitations by sampling the entire environment in a rapid and objective manner. What is required is a method of connecting these satellite-based measurements to LFA measurements and then being able to extrapolate these measurements across the entire mine surface. The aim of this research was to develop a method to use satellite-based hyperspectral imagery to predict landscape function analysis (LFA) using partial least squares regression (PLSR). This was broken down into three objectives: (1) Collection of the LFA data in the field and validation of the LFA indices against other environmental variables collected at the same time, (2) validation of PLSR models predicting LFA indices and various environmental variables from ground-based spectra, and (3) production of risk maps based on predicting LFA indices and above-ground biomass using PLSR models and Hyperion satellite-based hyperspectral imagery. Although the study was based in grasslands at two mining regions, West Wits and Vaal River, a suitable Hyperion image was only available for Vaal River. A minimum of 374 points were sampled for LFA indices, ground-based spectra, above-ground biomass and soil cores along 2880 m of LFA transect from both mine sites. Soil cores were weighed fresh before sieving with a 2 mm sieve to separate root and stone fractions. The sieved soil fraction was tested for pH, EC, SOM, and for the West Wits samples, organic nitrogen and total extractable inorganic nitrogen. There was one modification to the LFA method where grass patches were collapsed into homogenous units as it was deemed not feasible to sample 180 m transects at grass tuft scales of 10 – 30 cm, but other patch definitions followed the LFA manual (Tongway and Hindley, 2004). Evidence suggested that some of the different patch types, in particular the bare/biological soil crust – bare grass – sparse grass patch types, represented successional stages in a continuum although this was not conclusive. There also was evidence that the presence or absence of cattle play a role in some processes active in these grasslands and erosion is mainly through deflation, rain splash and sheet wash. Generally the environmental variables supported the LFA indices although the nutrient cycling index was representative of above-ground nutrient cycling but not below-ground nutrient cycling. Models derived with PLSR to predict the LFA indices from ground-based spectral measurements were strong at both mine sites (West Wits: LFA stability r2 = 0.63, P < 0.0001; LFA infiltration r2 = 0.75, P < 0.0001; LFA nutrient cycling r2 = 0.73, P < 0.0001; Vaal River: LFA stability r2 = 0.39, P < 0.0001, LFA infiltration r2 = 0.72, P < 0.0001, LFA nutrient cycling r2 = 0.54, P < 0.0001), as were PLSR models predicting above-ground biomass (West Wits above-ground biomass r2 = 0.55, P = 0.0003; Vaal River above-ground biomass r2 = 0.79, P < 0.0001) and soil moisture (West Wits soil moisture r2 = 0.45, P = 0.0017; Vaal River soil moisture r2 = 0.68, P < 0.0001). However, for soil organic matter (r2 = 0.50, P < 0.0001) and EC (r2 = 0.63, P < 0.0001), Vaal River had strong prediction models while West Wits had weak models for these variables (r2 = 0.31, P = 0.019 and r2 = 0.10 and P < 0.18, respectively). For EC, the wide range of soil values at Vaal River in association with gypsum crusts, and low values throughout West Wits explained these model results but for soil organic matter, no clear explanation for these site differences was identified. Patch-based models could accurately discriminate between spectrally well-defined patch types such S. plumosum patches but were less successful with patch types that were spectrally similar such as the bare/biological soil crust – bare grass – sparse grass patch continuum. Clustering similar patch types together before PLSR modelling did improve these patch-based spectral models. To test the method proposed to predict LFA indices from satellite-based hyperspectral imagery, a Hyperion image matching 6 transects at Vaal River was acquired by NASA’s EO-1 satellite and downloaded from the USGS Glovis website. LFA transects were partitioned to match and extract pixel spectra from the Hyperion data cube. Thirty-one spectra were separated into calibration (20) and validation (11) data. PLSR models were derived from the calibration data, tested with validation data to select the optimum model, and then applied to the entire Hyperion data cube to produce prediction maps for five LFA indices and above-ground biomass. The patch area index (PAI) produced particularly strong models (r2 = 0.79, P = 0.0003, n =11) with validation data, whereas the landscape organization index (LOI) produced weak models. It is argued that this difference between these two essentially similar indices is related to the fact that the PAI is a 2-dimensional index and the LOI is a 1-dimensional index. This difference in these two indices allowed the PAI to compensate for some burned pixels on the transects by “seeing” the density pattern of grass tufts and patches whereas the linear nature of the LOI was more susceptible to the changing dimensions of patch structure due to the effects of fire. Although validation models for the three LFA indices of soil stability, infiltration and nutrient cycling were strong (r2 = 0.72, P = 0.004; r2 = 0.66, P = 0.008; r2 = 0.70, P = 0.005, n = 9 respectively), prediction maps were confounded by the presence of fire on some transects. The poor quality of the Hyperion imagery also meant great care had to be taken in the selection of models to avoid poor quality prediction maps. The 31 bands from the VNIR (478 – 885 nm) portion of the Hyperion spectra were generally the best for PLSR modelling and prediction maps, presumably because of better signal-to-noise ratios due to higher energy in the shorter wavelengths. With two satellite-based hyperspectral sensors already operational, namely the US Hyperion and the Chinese HJ-1A HSI, and a number expected to be launched by various space agencies in the next few years, this research presents a method to use the strengths of LFA and hyperspectral imagery to model and predict LFA index values and thereby produce risk maps of large, heterogeneous landscapes such as mining environments. As this research documents a method of partitioning the landscape rather than the pixel spectra into pure endmembers, it makes a valuable contribution to the fields of landscape ecology and hyperspectral remote sensing. / LG2017

New farming needs and innovations have, over time, led to changes in land use. Arable fields have been turned into pastures and semi-natural pastures into arable fields. Due to the ecological value of continually grazed semi-natural pastures in Sweden the aim of this study is to find out how former arable fields that are permanently grazed differ from semi-natural pastures. In this study I selected appropriate semi-natural pastures from a national monitoring program on seminatural- pastures and meadows. Plots continuously grazed were compared with grazed plots on former arable fields. Pair-wise differences in the Ellenberg indicator values Light (L), Soil moisture (F), soil pH (R) and soil nitrogen (N), vertical coverage of trees, bushes and vegetation, species richness and species composition were tested. Except for species composition there were only small differences between former arable fields that are permanently grazed and semi-natural pastures. That means former arable fields can with time and grazing gain many of the values continuously grazed semi-natural pastures have.