Doctor of Philosophy / Department of Geography / Douglas Goodin / Quantitative remote sensing provides an effective way of estimating and mapping vegetation characteristics over an extensive area. The spatially explicit distribution of canopy vegetative properties from remote sensing imagery can be further used for studies of spatial patterns and processes in grassland systems. My research focused on remote sensing of grassland vegetation characteristics and its applications to spatial analysis of grassland dynamics involving interactions between pyric herbivory and vegetation heterogeneity. In remote sensing of vegetation characteristics, (1) I estimated the foliar pigments and nutritional elements at the leaf level using hyperspectral data. The foliar pigments, chlorophylls and carotenoids, were retrieved by inverting the physical radiative transfer model, PROSPECT. The nutritional elements were modeled empirically using partial least squares (PLS) regression. Correlations were found between the leaf pigments and nutritional elements. This provided insight into the use of pigment-related vegetation indices as a proxy of the plant nutritional quality. (2) At the canopy level, I assessed the use of the broadband vegetation indices, normalized difference vegetation index (NDVI) and green-red vegetation index (GRVI), in detecting vegetation quantity (LAI) and quality (leaf and canopy chlorophyll concentrations). The relationships between vegetation indices and vegetation characteristics were examined in the physical model, PROSAIL, and validated by a field dataset collected from a tallgrass prairie. NDVI showed high correlations with LAI and canopy chlorophylls. GRVI performed even better than NDVI in estimating LAI. A new index GNVI (green-red normalized vegetation index) that combined NDVI and GRVI was proposed to extract leaf chlorophyll concentration. These findings showed the potential of using broadband vegetation indices from multispectral remote sensors to monitor vegetation quantity and quality over a wide spatial extent. In the spatial analysis, I examined interactions between pyric herbivory and grassland heterogeneity at multiple scales from the remote sensing imagery. (3) At a coarse, watershed level, I evaluated effects of fire and large herbivores on the spatial distribution of canopy nitrogen. It was found that the interactive effects of fire and ungulate grazing were present in the watersheds burnt in spring, where a high level of ungulate grazing reduced vegetation density, but promoted canopy heterogeneity. Two grazer species, bison and cattle, were compared. Differences in the vegetation canopy between sites with bison and cattle were observed, which may be related to differences in the grazing intensity, forage behavior and habitat selection between the two grazer species. (4) At a fine, patch level (30 m), bison forage pattern was examined associated with canopy nitrogen heterogeneity. Bison preference for patches with high canopy nitrogen was evident in May. Later in June – September, bison tended to avoid sites with high canopy nitrogen. Vegetation heterogeneity showed significant influences on bison habitat selection in June. Bison preferred sites with low variance in canopy nitrogen, where the patch types were highly aggregated and equitably proportioned.

The aim of this thesis is to assess the impact of farming on grassland and assess the frequency of use of grassland on yield and quality of forage biomass. A part of this thesis is also a recommendation of possible management changes of evaluated grasslands. Botanical inventory surveys were conducted on agricultural lands Bílovské zemědělské a.s., in the northern Plzeň region. There have been selected three sites, two meadows and one pasture vegetation. The comprehensive analysis of the stand composition was created by agrotechnology of meadows. An important part of this thesis is to assess the impact of different grassland management practices with regard to their economic, social and ecological importance.

O Pampa é um bioma do tipo savana exclusivo da região Neotropical, situado na porção sul da América do Sul, cobrindo a Argentina, o Uruguai e o Brasil. O clima é quente temperado e úmido, com temperaturas anuais que variam de 0° C no inverno e mais de 24° C no verão. Os grandes distúrbios no Pampa se iniciaram a cerca de 300 anos com a chegada da colonização Europeia na região e o advento da agricultura, silvicultura, manejo sem controle pelo fogo e a criação de gado, leiteiro e de corte. Apesar de contribuir para a falta de heterogeneidade vegetal e perda na diversidade de fauna, a pecuária ainda é extensivamente indicada como manejo sustentável do Pampa. Com pesquisas e trabalhos por vezes mais focados no aumento da produtividade do rebanho que na manutenção da biodiversidade, a conservação do Pampa vem sendo negligenciada. O monitoramento de diferentes metodologias de pastejo, tradicional, em mosaico e exclusão total do gado, se faz necessário para um maior entendimento de como a diversidade de plantas e animais se comportam nestes sistemas. Para isso, são necessários bons modelos ecológicos para testes de hipóteses e diferentes manejos, o que sustenta a utilização de aranhas para estes estudos. Aranhas são rápidas colonizadoras, ótimas dispersoras, predadoras generalistas, além de estarem estritamente relacionadas à estrutura vegetal. Suscetíveis a impactos bióticos e abióticos, aranhas são ótimas medidoras de qualidade ambiental e, sendo de fácil amostragem, podem ser envolvidas em estudos sobre diversidade de espécies e indivíduos. Para tanto, este estudo foi realizado na APA do Ibirapuitã, na Campanha do Rio Grande do Sul, Brasil. Seis parcelas, de um hectare cada, foram distribuídas em pares em três fazendas. Cada par é constituído de uma parcela com pastejo, demarcadas apenas por estacas, onde o gado tem livre acesso, e uma parcela sem pastejo, cercadas desde Agosto de 2012 onde o pastejo do gado foi excluído por completo. A principal expectativa principal é que a abundancia, riqueza, biomassa e composição de aranhas serão diferentes quando áreas com e sem pastejo são comparadas forem comparados. As amostragens foram realizadas com o uso de armadilhas de queda do tipo “Pitfall traps” nas primaveras de 2011, 2012 e 2013, e sazonalmente no outono (maio), inverno (setembro) e primavera (novembro) de 2015 e verão (fevereiro) de 2016 para a captura de aranhas de hábito epígeo. Para análises estatísticas, Testes T, ANOVA e ANOSIM foram realizadas. Linyphiidae e Lycosidae foram as famílias mais abundantes e ricas em todas as coletas realizadas. Os resultados obtidos em 2011, 2012 e 2013 não demonstraram diferença significativa na fauna de aranhas, abundância e riqueza quando áreas com e sem pastejo foram comparadas. Contudo, os resultados apresentaram valores maiores nas áreas com pastejo quando comparando com áreas sem pastejo. Em 2015/2016 a abundância e a riqueza de aranhas também não foram significativamente diferentes nos dados coletados, contudo, os valores brutos também apresentaram maiores abundância e riqueza em áreas com pastejo do que nas sem pastejo. Analisando-se as estações como um todo, juntando áreas com e sem pastejo, o verão foi a estação com mais riqueza geral. Contudo, analisando as áreas com e sem pastejo separadamente, o inverno foi a estação mais rica e abundante em cada uma destas áreas. Apesar dos resultados anteriores, as espécies mais abundantes demonstraram diferença significativa em suas abundancias em cada estação do ano quando áreas com e sem pastejo foram comparados. A biomassa das aranhas não foi significativamente diferente para todos os anos entre áreas com e sem pastejo, com valores maiores em áreas com pastejo. Entretanto, espécies mais abundantes demonstraram diferenças significativas em sua biomassa sazonalmente nos anos de 2015 e 2016, com valores maiores em áreas com pastejo. Áreas com e sem pastejo ainda se mantém similares mesmo após quase quatro anos de exclusão do gado em termos de abundância total de aranhas. Apenas aranhas mais abundantes demonstram diferença entre áreas com e sem pastejo em termos de abundância e biomassa. / Pampa is a savanna-like biome, exclusively of Neotropical region, lying in southern part o South America, covering Argentina, Uruguay and Brazil. The climate is hot temperate and humid, with annual temperatures ranging of 0° C in winter and higher than 24° C in summer. The increase of disturbance at Pampa began around 300 years after Europian colonization and the advent of agriculture, silviculture, uncontrolled fire management and creation of dairy and beef cattle. Even livestock causing loss of vegetal heterogeneity and fauna diversity, it is a widely recommended methodology for sustainable management of Pampa. With studies more focused in forage properties and beef production than diversity maintenance, the Pampa conservation has been neglected. Monitoring different livestock methodologies are necessary to improve knowledge of how diversity of plants and animals change during this impact. Thus, using great ecological models, like spiders, are necessary to test hypotheses and different managements. Spiders are quick settlers, great dispersers, generalist predators and are much correlated with vegetal structure. Susceptible to biotic and abiotic impacts, spiders are great environmental quality meters, being easy to sample and to measure abundance, diversity and richness. This study was conduct at Ibirapuitã River Environmental Federal Protection Area “Área de Proteção Ambiental – APA do Rio Ibirapuitã” in Rio Grande do Sul state, Brazil. Six plots, of one hectare each, were distributed in pars in three farms. Each pair is constituted of a closed plot, a fenced plots since August 2012, with total exclusion of cattle graze; and a opened plot with only poles in the corners to demarcate the place, and cattle have free access for graze. Our main expectative is that abundance, richness, biomass and composition of arenofauna will be different when grazed and ungrazed areas were compared. Ground dwelling spiders were sampled with Pitfall traps in springs of 2011, 2012 and 2013, and seasonally in autumn (May), winter (September) and spring (November) 2015 and summer (February) 2016. Statistical analyses were performed using Stundents’ Test, ANOVA and ANOSIM. Linyphiidae and Lycosidae were the most abundant and richer families. Results of 2011, 2012 and 2013 were no significant different in spider abundance and richness when grazed and ungrazed plots were compared. However, raw data suggest higher values of abundance in grazed areas than ungrazed ones. In 2015/2016 samples, spider abundance and richness were not significant different between grazed and ungrazed areas. Raw data suggest that grazed areas abundance and richness was higher than in ungrazed though. Although summer was the richer season in general, winter was the richer season separately in grazed and ungrazed areas. Despite the previous results, abundance of the most abundant species were significant different between grazed and ungrazed areas in general and in each season. Spider biomass was not significant different in any year when grazed and ungrazed areas were compared. However, biomass of most abundant species were significant different seasonally and in each season in 2015/2016. Grazed and ungrazed areas are still very similar even after almost four years of cattle exclusion in terms of spider abundance and biomass. Only most abundant species demonstrate a significant difference between grazed and ungrazed areas in terms of abundance and biomass, with higher values in grazed plots.

O conhecimento da fenologia é baseado nas observações de períodos de desenvolvimento da planta. Mudanças intra e interanuais na temperatura e precipitação podem influenciar diretamente as diferentes fases da fenologia, assim como as condições de umidade dos diferentes tipos de solo ao qual a vegetação está associada. O sensoriamento remoto é uma ferramenta que torna possível a aquisição de informações para análise espacial e temporal da vegetação. Este trabalho apresenta um estudo sobre as variações da resposta espectral da vegetação campestre natural na Área de Proteção Ambiental (APA) do Ibirapuitã, localizada no bioma Pampa, avaliando suas relações com os elementos do clima. Para isso, a metodologia compreendeu a coleta de dados climáticos de temperatura média do ar e precipitação pluvial; dados orbitais a partir de imagens de média resolução espacial do sensor MODIS para o período de 2000 a 2009, com posterior estimativa da correlação estatística entre estes dados. Além disso, utilizou-se o produto SRTM para auxiliar definição da grade amostral, em função da altimetria e solo. Os resultados mostraram os períodos de crescimento/desenvolvimento da vegetação campestre natural associados às estações do ano, com um período de crescimento nas estações quentes, diminuição da reflectância na região espectral do visível e aumento da reflectância no infravermelho próximo. Observou-se também, um período de senescência na estação fria do ano, onde o comportamento espectral da vegetação apresenta um aumento da reflectância na região espectral do visível e diminuição da reflectância no infravermelho próximo. As diferenças na reflectância da vegetação para cada tipo de solo foi observada para solos mais profundos, capazes de armazenar mais água. De maneira geral, verificou-se que a resposta espectral da vegetação campestre apresentou relação com as variações de temperatura média do ar e precipitação na região, respondendo às variáveis meteorológicas com um atraso aproximado de dezesseis (16) dias. Verificou-se assim que dados MODIS do infravermelho próximo, são adequados para monitorar o início da estação de máximo crescimento da vegetação campestre natural da APA do Ibirapuitã que se inicia na primavera. Este período está relacionado com a temperatura média do ar. / The knowledge of phenology is based on observations of periods of plant development. Inter-annual changes in temperature and rainfall may directly influence the different stages of phenology, as well as the moisture conditions of different soil types to which the vegetation is associated. Remote sensing is a tool which allows the acquisition of information for spatial and temporal analysis of vegetation. This work has aimed to study the variations of the natural grassland spectral response in the Ibirapuitã environmental protected area, which belongs to the Pampa biome, evaluating their relationship with the elements of climate. The methodology has consisted of collection of meteorological data, such as air temperature and rainfall, and orbital data from medium spatial resolution MODIS images, which covered the period from 2000 to 2009. It has also performed a statistical correlation among those data. In addition, a SRTM-defining product has employed as sampling grid, depending on elevation and soil. The results have showed periods of growth and development of natural grassland vegetation associated with the seasons, with a growth period during the warm seasons, and with a decrease of the reflectance in the visible spectrum and with an increase in the near infrared one. It was also observed a period of senescence during the cold season, where the spectral characteristics of the vegetation present a increase of the reflectance in the visible spectrum and with an decrease in the near infrared one. The differences in reflectance of vegetation in each soil type have showed most prominent in the deeper ones, which have a large water storage capacity. Overall, it has been observed a tight coupling between the spectral response of grassland vegetation and changes in air temperature and rainfall in the study area, so that the former has responded to the meteorological variables with a delay of approximately sixteen (16) days. Near-infrared MODIS data have been showed suitable to monitor the beginning of the season of maximum growth of natural grassland vegetation in the APA of the Ibirapuitã of which begins in spring. This period is mainly related to air temperature.

Grasslands in the western United States face severe environmental threats including those brought about by climate change, such as changes in precipitation regimes and altered fire cycles; land-use conversion and development; and the introduction, establishment, and spread of non-native species. Lehmann’s lovegrass (Eragrostis lehmanniana) was introduced to the southwestern United States in the early 1900s. Since its introduction, it has become the dominant grass in the mid-elevation grasslands of southern Arizona, including the Santa Rita Experimental Range (SRER), where it has displaced native grasses including Arizona cottontop, three awns, and gramas. Like all plants in terrestrial ecosystems, this grass harbors fungal symbionts that can be important for its establishment and persistence. This thesis focuses on fungal symbionts of Lehmann’s lovegrass and has two components. First, the diversity and distributions of endophytes in Lehmann’s lovegrass are evaluated in the context of biotic and abiotic factors in the SRER. Culturing from roots and shoots of Lehmann’s lovegrass at points beneath and outside the canopy of native mesquites, which are encroaching on grasslands over time, provides insight into how a single plant species can exhibit local variation in the composition of its symbionts. Second, the thesis is used as the basis for engagement of students in science, technology, engineering, and mathematics (STEM) through the development and implementation of classroom- and field activities centered on endophytes, which help high school students address core learning aims while also gaining real research experience. Engaging students in important questions relevant to their local environment can catalyze interest in science and help students cross the threshold into research. The contributions of such approaches with respect to learning not only fulfills key next-generation science standards and common core objectives, but provides students with a meaningful introduction to the excitement, importance, and accessibility of science.

Afforestation is considered to be one of the leading land-use changes affecting ecosystem function and diversity. This study investigates the impact of pine afforestation on microclimate and soil in fragments of highly vulnerable Afromontane grassland at Groenvaly. Three major challenges for afforestation research are identified as (i) the range and intricacy of the impact of afforestation, (ii) differences in measurement and monitoring periods and (iii) a lack of focus on biomes adjacent to plantations. The approach here aimed to address these three areas. Air microclimate data were collected for 24 months within a plantation site, a control grassland site and a grassland fragment using three Davis Vantage Pro2 weather stations. Soil temperature data were logged on iButtonsTM for 18 months and soil samples from four seasons were analysed for moisture content, nitrogen (N), nitrate, ammonium, phosphorous (P), pH, sodium (Na), calcium (Ca), potassium (K), magnesium (Mg) and soil organic carbon (SOC). All data were statistically analysed at within-site, between-site, seasonal and mean scales and each analysis highlighted different conclusions. Results for the chemicals properties of the individual grassland fragments did not exhibit within-site variation except for K and P and between-site variation was only evident for N, nitrate, moisture and SOC. Solar irradiance was reduced in the fragments only during winter while SOC and P in the fragments only differed from the control grassland sites in summer and autumn respectively. Mean values for P, pH, Na, Ca, K, SOC and soil moisture within the fragments’ soil were between those of the control grassland and the plantation while N and Mg values were closer to the plantation than the control grassland. Mean values for air temperature, wind speed, solar irradiance and humidity within the fragment were closer to the control grassland than the plantation. Soil temperatures at 2cm in the fragments were similar to the control grassland, while temperatures at 10cm below the surface were lower than both the control grassland and the plantation sites in winter. Maximum air temperatures in the fragments were lowered in summer and raised in winter but minimum air temperatures were raised in the fragments across all seasons. Results of this study show that there are different impacts in different seasons while overall mean data indicate that the fragments’ soil is affected, and microclimate is unaffected, by the plantation. The impact of the change in soil and microclimate in grassland fragments requires more investigation to determine if grassland fragments are a suitable conservation strategy in pine plantations. / Dissertation (MSc)--University of Pretoria, 2013. / gm2014 / Geography, Geoinformatics and Meteorology / unrestricted

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.

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 Arts / Department of Geography / J. M. Shawn Hutchinson / Grasslands of the Flint Hills are often burned as a land management practice. Remote sensing can be used to help better manage prairie landscapes by providing useful information about the long-term trends in grassland vegetation greenness and helping to quantify regional differences in vegetation development. Using MODIS 16-day NDVI composite imagery between the years 2001-10 for the entire Flint Hills ecoregion, BFAST was used to determine trend, seasonal, and noise components of the image time series. To explain the trend, 4 factors were considered including hydrologic soil group, burn frequency, and precipitation deviation from the 30 year normal. In addition, the time series data was processed using TIMESAT to extract eight different phenometrics: Growing season length, start of season, end of season, middle of season, maximum value, small integral, left derivative, and right derivative. Phenometrics were produced for each year of the study and an ANOVA was performed on the means of all eight phenometrics to assess if significant differences existed across the study area. A K-means cluster analysis was also performed by aggregating pixel-level phenometrics at the county level to identify administrative divisions exhibiting similar vegetation development. For the study period, the area of negatively and positively trending grassland were similar (41-43%). Logistic regression showed that the log odds of a pixel experiencing a negative trend were higher in sites with clay soils and higher burning frequencies and lower for pixels having higher than normal precipitation and loam soils. Significant differences existed for all phenometrics when considering the ecoregion as a whole. On a phenometric-by-phenometric basis, unexpected groupings of counties often showed statistically similar values. Similarly, when considering all phenometrics at the same time, counties clustered in surprising patterns. Results suggest that long-term trends in grassland conditions warrant further attention and may rival other sources of grassland change (e.g., conversion, transition to savannah) in importance. Analyses of phenometrics indicates that factors other than natural gradients in temperature and precipitation play a significant role in the annual cycle of grassland vegetation development. Unanticipated, and sometimes geographically disparate, groups of counties were shown to be similar in the context of specific phenology metrics and this may prove useful in future implementations of smoke management plans within the Flint Hills.

A plant ecological study was conducted in the grasslands of Transkei to develop a better understanding of the distribution, structure and composition of the plant communities found in the area. The agricultural sector in developed and rapidly developing areas of southern Africa is faced with problems like veld deterioration and the loss of natural areas that effectively contribute to the depopulation of rural areas. The Grassland Biome Project was initiated to address the problem of increased destruction of natural resources because the most suitable area for agriculture falls within the grasslands of South Africa. The aim of the Grassland Biome Project is to integrate knowledge, comprehension and expertise, which will enable scientists to forecast the results of the available options of grassland management programs. Vegetation of Transkei forms part of this project. The study area (Transkei) borders on the kingdom of Lesotho in the north and Indian Ocean in the southeast. In the northeast it borders on KwaZulu-Natal and in the northwest, and in the west on the eastern Cape Province. Relevés were compiled in 379 random sample plots and four major plant communities, namely: Bush and Grassland vegetation of drier areas, Wet and Disturbed grassland, Grassland of Undulating Areas, and Moist Grasslands on Flat Plains, were found. The vegetation was classified by means of TWINSPAN and Braun-Blanquet procedures were found. The vegetation was classified by means of TWINSPAN and Braun-Blanquet procedures. / Dissertation (MSc)--University of Pretoria, 2005. / Geography, Geoinformatics and Meteorology / Unrestricted