O objetivo geral desta tese foi examinar diferentes 1 questões associadas ao 2 processo de invasão de plantas para investigar os mecanismos, impactos e 3 medidas de recuperação da comunidade vegetal, abordando diferentes estratégias 4 metodológicas que incluem estudos observacionais, experimentos de remoção e 5 uma revisão metodológica. Os resultados do Capítulo 1 mostraram como as 6 relações entre condições climáticas e estrutura da paisagem podem determinar o 7 grau de invasão de plantas na escala regional. Observou-se que os padrões de 8 invasão dos campos sulinos estão principalmente relacionados com maior 9 densidade de estradas, menor cobertura de campo nativo e com o aumento do 10 déficit hídrico. Além disso, constatou-se que a gramínea Eragrostis plana é a 11 planta invasora mais abundante dos campos sulinos. No entanto, os resultados do 12 experimento do Capítulo 2 mostraram que sua invasão não pode ser explicada 13 pela riqueza de espécies ou composição dos grupos funcionais de plantas da 14 comunidade residente. A invasão de E. plana foi principalmente associada ao 15 distúrbio causado pela remoção de biomassa na comunidade. Neste sentido, as 16 comunidades campestres poderiam ser resistentes à invasão de E. plana até que 17 algum distúrbio aumente sua vulnerabilidade. Por outro lado, através da 18 comparação entre comunidades invadidas, removidas e não-invadidas, os 19 resultados do experimento do Capítulo 3 contribuíram no entendimento do 20 impacto real da invasão de E. plana, em termos de redução da riqueza e cobertura 21 de plantas nativas na comunidade. No entanto, embora os métodos de remoção 22 utilizados reduziram a cobertura da invasora, não foram suficientes para conseguir 23 sua erradicação local. Além disso, após três anos de remoção de E. plana, as comunidades se tornaram distintas às invadidas mas não 1 foram semelhantes às 2 comunidades não invadias, o qual poderia indicar que outras medidas de 3 restauração são ainda necessárias. Embora os experimentos de remoção de 4 espécies sejam úteis para investigar questões associadas ao processo de 5 invasão, existem limitações importantes a considerar, como foi evidenciado nos 6 Capítulos 2 e 3. Neste sentido, oferecemos a revisão bibliográfica sistemática do 7 Capítulo 4, onde se discute o potencial dos métodos de remoção utilizados para 8 estudar a resistência e a recuperação da comunidade à invasão, apontando 9 algumas limitações. Como resultado, o Capítulo 4 mostrou que a maioria dos 10 trabalhos não propõem controles adequados nos experimentos, o que pode dar 11 lugar a confundimento de efeitos. Desta forma, foram desenvolvidas algumas 12 sugestões para serem consideradas nos experimentos de remoção de espécies, 13 com o objetivo de continuar avançando nesta temática. As informações geradas 14 nesta tese podem contribuir para o entendimento do processo de invasão de 15 plantas nos ecossistemas campestres, com vistas ao manejo, à conservação e à 16 restauração das comunidades invadidas, adquirindo um senso crítico no 17 planejamento de desenhos experimentas. / The general aim of this thesis was to investigate different 1 issues associated 2 with plant invasion process to understand the mechanisms, impacts and 3 community recovery, by employing different methodological strategies such as 4 observational studies, removal experiments and a literature review. The results 5 from Chapter 1 showed how the interactions between climate and landscape 6 structure can determine the level of invasion of South Brazilian grasslands, 7 highlighting that invasion is mainly related to high road density, less native 8 grassland cover and increased aridity. Furthermore, it was confirmed that 9 Eragrostis plana is the most important invasive species in the South Brazilian 10 grasslands. However, the results from the experiment of Chapter 2 showed that its 11 invasion could not be explained by the species richness or functional group 12 composition in the community. Eragrostis plana invasion was associated with the 13 disturbance effect caused by the amount of removed biomass. Thus, grassland 14 communities may be resistant to E. plana invasion until some disturbance 15 increases their vulnerability. Moreover, by comparing invaded, removed and non16 invaded communities, the experimental results of Chapter 3 highlighted the 17 ecological impact of E. plana invasion, in terms of richness reduction and native 18 species cover. However, although removals methods reduced the cover of the 19 invasive species, they were not enough to locally extinct it. Moreover, after three 20 years of the invasive removal, communities became different from invaded ones 21 but not resembling non-invaded references, which suggest that community 22 recovery may require restoration strategies. Although removal experiments have 23 been useful to investigate certain issues associated with invasion process, there are important limitations to consider, as was shown in Chapter 1 2 and 3. For this 2 purpose, our systematic review presented in Chapter 4 discussed the potential of 3 removal methods for assessing community resistance and recovery from invasion. 4 There, we showed that most of the studies did not use adequate controls in 5 removal experiments, which can lead to confounding effects. Thus, we developed 6 suggestions to be considered in experimental designs to advance the 7 methodological technique of removals. The information generated in this thesis can 8 contribute to the understanding of plant invasion process in South Brazilian 9 grasslands and, consequently, aid to management, conservation and restoration of 10 invaded communities by acquiring a critical sense in experimental designs.

O objetivo geral desta tese foi examinar diferentes 1 questões associadas ao 2 processo de invasão de plantas para investigar os mecanismos, impactos e 3 medidas de recuperação da comunidade vegetal, abordando diferentes estratégias 4 metodológicas que incluem estudos observacionais, experimentos de remoção e 5 uma revisão metodológica. Os resultados do Capítulo 1 mostraram como as 6 relações entre condições climáticas e estrutura da paisagem podem determinar o 7 grau de invasão de plantas na escala regional. Observou-se que os padrões de 8 invasão dos campos sulinos estão principalmente relacionados com maior 9 densidade de estradas, menor cobertura de campo nativo e com o aumento do 10 déficit hídrico. Além disso, constatou-se que a gramínea Eragrostis plana é a 11 planta invasora mais abundante dos campos sulinos. No entanto, os resultados do 12 experimento do Capítulo 2 mostraram que sua invasão não pode ser explicada 13 pela riqueza de espécies ou composição dos grupos funcionais de plantas da 14 comunidade residente. A invasão de E. plana foi principalmente associada ao 15 distúrbio causado pela remoção de biomassa na comunidade. Neste sentido, as 16 comunidades campestres poderiam ser resistentes à invasão de E. plana até que 17 algum distúrbio aumente sua vulnerabilidade. Por outro lado, através da 18 comparação entre comunidades invadidas, removidas e não-invadidas, os 19 resultados do experimento do Capítulo 3 contribuíram no entendimento do 20 impacto real da invasão de E. plana, em termos de redução da riqueza e cobertura 21 de plantas nativas na comunidade. No entanto, embora os métodos de remoção 22 utilizados reduziram a cobertura da invasora, não foram suficientes para conseguir 23 sua erradicação local. Além disso, após três anos de remoção de E. plana, as comunidades se tornaram distintas às invadidas mas não 1 foram semelhantes às 2 comunidades não invadias, o qual poderia indicar que outras medidas de 3 restauração são ainda necessárias. Embora os experimentos de remoção de 4 espécies sejam úteis para investigar questões associadas ao processo de 5 invasão, existem limitações importantes a considerar, como foi evidenciado nos 6 Capítulos 2 e 3. Neste sentido, oferecemos a revisão bibliográfica sistemática do 7 Capítulo 4, onde se discute o potencial dos métodos de remoção utilizados para 8 estudar a resistência e a recuperação da comunidade à invasão, apontando 9 algumas limitações. Como resultado, o Capítulo 4 mostrou que a maioria dos 10 trabalhos não propõem controles adequados nos experimentos, o que pode dar 11 lugar a confundimento de efeitos. Desta forma, foram desenvolvidas algumas 12 sugestões para serem consideradas nos experimentos de remoção de espécies, 13 com o objetivo de continuar avançando nesta temática. As informações geradas 14 nesta tese podem contribuir para o entendimento do processo de invasão de 15 plantas nos ecossistemas campestres, com vistas ao manejo, à conservação e à 16 restauração das comunidades invadidas, adquirindo um senso crítico no 17 planejamento de desenhos experimentas. / The general aim of this thesis was to investigate different 1 issues associated 2 with plant invasion process to understand the mechanisms, impacts and 3 community recovery, by employing different methodological strategies such as 4 observational studies, removal experiments and a literature review. The results 5 from Chapter 1 showed how the interactions between climate and landscape 6 structure can determine the level of invasion of South Brazilian grasslands, 7 highlighting that invasion is mainly related to high road density, less native 8 grassland cover and increased aridity. Furthermore, it was confirmed that 9 Eragrostis plana is the most important invasive species in the South Brazilian 10 grasslands. However, the results from the experiment of Chapter 2 showed that its 11 invasion could not be explained by the species richness or functional group 12 composition in the community. Eragrostis plana invasion was associated with the 13 disturbance effect caused by the amount of removed biomass. Thus, grassland 14 communities may be resistant to E. plana invasion until some disturbance 15 increases their vulnerability. Moreover, by comparing invaded, removed and non16 invaded communities, the experimental results of Chapter 3 highlighted the 17 ecological impact of E. plana invasion, in terms of richness reduction and native 18 species cover. However, although removals methods reduced the cover of the 19 invasive species, they were not enough to locally extinct it. Moreover, after three 20 years of the invasive removal, communities became different from invaded ones 21 but not resembling non-invaded references, which suggest that community 22 recovery may require restoration strategies. Although removal experiments have 23 been useful to investigate certain issues associated with invasion process, there are important limitations to consider, as was shown in Chapter 1 2 and 3. For this 2 purpose, our systematic review presented in Chapter 4 discussed the potential of 3 removal methods for assessing community resistance and recovery from invasion. 4 There, we showed that most of the studies did not use adequate controls in 5 removal experiments, which can lead to confounding effects. Thus, we developed 6 suggestions to be considered in experimental designs to advance the 7 methodological technique of removals. The information generated in this thesis can 8 contribute to the understanding of plant invasion process in South Brazilian 9 grasslands and, consequently, aid to management, conservation and restoration of 10 invaded communities by acquiring a critical sense in experimental designs.

O objetivo geral desta tese foi examinar diferentes 1 questões associadas ao 2 processo de invasão de plantas para investigar os mecanismos, impactos e 3 medidas de recuperação da comunidade vegetal, abordando diferentes estratégias 4 metodológicas que incluem estudos observacionais, experimentos de remoção e 5 uma revisão metodológica. Os resultados do Capítulo 1 mostraram como as 6 relações entre condições climáticas e estrutura da paisagem podem determinar o 7 grau de invasão de plantas na escala regional. Observou-se que os padrões de 8 invasão dos campos sulinos estão principalmente relacionados com maior 9 densidade de estradas, menor cobertura de campo nativo e com o aumento do 10 déficit hídrico. Além disso, constatou-se que a gramínea Eragrostis plana é a 11 planta invasora mais abundante dos campos sulinos. No entanto, os resultados do 12 experimento do Capítulo 2 mostraram que sua invasão não pode ser explicada 13 pela riqueza de espécies ou composição dos grupos funcionais de plantas da 14 comunidade residente. A invasão de E. plana foi principalmente associada ao 15 distúrbio causado pela remoção de biomassa na comunidade. Neste sentido, as 16 comunidades campestres poderiam ser resistentes à invasão de E. plana até que 17 algum distúrbio aumente sua vulnerabilidade. Por outro lado, através da 18 comparação entre comunidades invadidas, removidas e não-invadidas, os 19 resultados do experimento do Capítulo 3 contribuíram no entendimento do 20 impacto real da invasão de E. plana, em termos de redução da riqueza e cobertura 21 de plantas nativas na comunidade. No entanto, embora os métodos de remoção 22 utilizados reduziram a cobertura da invasora, não foram suficientes para conseguir 23 sua erradicação local. Além disso, após três anos de remoção de E. plana, as comunidades se tornaram distintas às invadidas mas não 1 foram semelhantes às 2 comunidades não invadias, o qual poderia indicar que outras medidas de 3 restauração são ainda necessárias. Embora os experimentos de remoção de 4 espécies sejam úteis para investigar questões associadas ao processo de 5 invasão, existem limitações importantes a considerar, como foi evidenciado nos 6 Capítulos 2 e 3. Neste sentido, oferecemos a revisão bibliográfica sistemática do 7 Capítulo 4, onde se discute o potencial dos métodos de remoção utilizados para 8 estudar a resistência e a recuperação da comunidade à invasão, apontando 9 algumas limitações. Como resultado, o Capítulo 4 mostrou que a maioria dos 10 trabalhos não propõem controles adequados nos experimentos, o que pode dar 11 lugar a confundimento de efeitos. Desta forma, foram desenvolvidas algumas 12 sugestões para serem consideradas nos experimentos de remoção de espécies, 13 com o objetivo de continuar avançando nesta temática. As informações geradas 14 nesta tese podem contribuir para o entendimento do processo de invasão de 15 plantas nos ecossistemas campestres, com vistas ao manejo, à conservação e à 16 restauração das comunidades invadidas, adquirindo um senso crítico no 17 planejamento de desenhos experimentas. / The general aim of this thesis was to investigate different 1 issues associated 2 with plant invasion process to understand the mechanisms, impacts and 3 community recovery, by employing different methodological strategies such as 4 observational studies, removal experiments and a literature review. The results 5 from Chapter 1 showed how the interactions between climate and landscape 6 structure can determine the level of invasion of South Brazilian grasslands, 7 highlighting that invasion is mainly related to high road density, less native 8 grassland cover and increased aridity. Furthermore, it was confirmed that 9 Eragrostis plana is the most important invasive species in the South Brazilian 10 grasslands. However, the results from the experiment of Chapter 2 showed that its 11 invasion could not be explained by the species richness or functional group 12 composition in the community. Eragrostis plana invasion was associated with the 13 disturbance effect caused by the amount of removed biomass. Thus, grassland 14 communities may be resistant to E. plana invasion until some disturbance 15 increases their vulnerability. Moreover, by comparing invaded, removed and non16 invaded communities, the experimental results of Chapter 3 highlighted the 17 ecological impact of E. plana invasion, in terms of richness reduction and native 18 species cover. However, although removals methods reduced the cover of the 19 invasive species, they were not enough to locally extinct it. Moreover, after three 20 years of the invasive removal, communities became different from invaded ones 21 but not resembling non-invaded references, which suggest that community 22 recovery may require restoration strategies. Although removal experiments have 23 been useful to investigate certain issues associated with invasion process, there are important limitations to consider, as was shown in Chapter 1 2 and 3. For this 2 purpose, our systematic review presented in Chapter 4 discussed the potential of 3 removal methods for assessing community resistance and recovery from invasion. 4 There, we showed that most of the studies did not use adequate controls in 5 removal experiments, which can lead to confounding effects. Thus, we developed 6 suggestions to be considered in experimental designs to advance the 7 methodological technique of removals. The information generated in this thesis can 8 contribute to the understanding of plant invasion process in South Brazilian 9 grasslands and, consequently, aid to management, conservation and restoration of 10 invaded communities by acquiring a critical sense in experimental designs.

In depth studies of the vegetation of large parts of southern Africa, including detailed maps and descriptions of vegetation units are mainly limited to small areas. Meanwhile, various agricultural practises have led to destruction or deterioration of the quality of natural grassland ecosystems. The agricultural sector in developed and rapidly developing areas of southern Africa is confronted with problems like veld deterioration and the loss of natural areas that effectively contribute to the depopulation of rural areas. The necessity to identify, classify and describe the vegetation types and communities within the Grassland Biome was stressed by Mentis and Huntley (1982). 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 programmes. The phytosociological classification of northwestern KwaZulu-Natal forms part of this project. Known previous vegetation studies of this area were conducted on a large scale and a considerable time ago, which underlines the necessity for a more comprehensive and phytosociologically refined investigation of this area. The study area lies in the northern part of KwaZulu-Natal and comprises the Drakensberg mountains and slopes in the west and undulating plains and bushveld valleys in the east and south. Relevés were compiled in 526 stratified random sample plots over an area of 9300 km2, comprising the northwestern part of KwaZulu-Natal. The vegetation was classified by means of TWINSP AN and Braun-Blanquet procedures. A phytosociological investigation of this vegetation revealed great variation in floristic composition. Further refinement of the data disclosed five major vegetation types containing nine plant communities. The topography and geology of the study area contributes greatly to the diversity of the vegetation, but poor agricultural practises have caused deterioration of the vegetation. / Dissertation (MSc)--University of Pretoria, 1998. / Plant Science / unrestricted

This study investigates the impact of arrival order and seed traits on plant community assembly and biodiversity in a grassland ecosystem in northern Sweden. A field experiment was conducted using 20 different grassland species, with germination pre-tests in the greenhouse and sowing experiments in the field. Seed mass and germination speed were selected seed traits to provide insights into the competitive ability and early life history strategies of the sown species. In the field study, four arrival order treatments with both slow- and fast-germinating species were performed on 70 m2 plots. After three years, the aboveground biomass of the established species was harvested and analyzed, revealing that species arriving second exhibited significantly lower biomass compared to those arriving first. Our results thereby suggest that arrival order has a significant impact on the productivity of species in grassland ecosystems. Priority effect strength (PES) was assessed using two different equations, with results indicating no significant relationship between PES and seed mass or germination speed. Biodiversity analysis revealed that plots with simultaneous late arrival had significantly lower Shannon diversity indices compared to arrival earlier in the season. These findings suggest that arrival order significantly influences plant productivity, while the effect on biodiversity in grassland ecosystems is less clear. Further, seed traits such as mass and germination speed may not play a significant role in determining the strength of priority effects during community assembly.

Research in grassland ecosystems worldwide has highlighted the crucial role many burrowing herbivore mammals provide as so-called “ecosystem engineers”. Many of these examples come from North America and have mostly been focused on Prairie Dogs (Cynomys sp.), which, due to their burrowing and grazing activities, are considered as a species of high ecological importance. Multiple studies have shown that their loss reduces grassland ecosystem functions and that their activities increase plant, arthropod, bird and mammal biodiversity, grassland heterogeneity, provide benefits to cattle, among many other key ecological roles. However, prairie dog species are distributed throughout multiple landscapes in North America, and most of the studies have been on mixed prairie grasslands. It is therefore not surprising that, when comparing the effects of different prairie dog species across ecoregions, the role of prairie dogs is not as clear and results vary, depending on spatial context, type of grasslands, climate and environmental conditions. Yet, grasslands are considered as some of the most highly threatened ecosystems, having high rates of conversion, desertification and biodiversity loss. Therefore, it is imperative to understand the effects prairie dog species have on the ecosystem. In this thesis, I focus on the Mexican prairie dog (C. mexicanus), a species in drastic decline due to habitat loss, caused by agriculture, overgrazing, human settlement, fragmentation, diseases (e.g. plague) and past eradication programs. C. mexicanus is considered endangered by the IUCN red list of threatened species and The Convention of International Trade in Endangered Species of Wild Fauna and Flora. It is also endemic to the Grassland Priority Conservation Area (GPCA) of El Tokio, located in Northeast Mexico within the Chihuahuan desert ecoregion and designated as a GPCA by the Commission for Environmental Cooperation (a cooperation between Canada, U.S. and Mexico) in 2009. Grasslands within GPCA El Tokio are highly fragmented as a result of many years of intense agricultural practices and other anthropogenic activities and around 90% of the original grasslands have been lost. However, it is considered an area of ecological importance due to its unique vegetation and edaphic associations, as well as being a key habitat for migratory birds. Up until now, very few studies have explored the role C. mexicanus has on plant taxonomical diversity and soil properties. Moreover, no studies focusing on prairie dogs have looked into the effect the species has on plant functional diversity or soil properties. Furthermore, GPCA El Tokio is a data-scarce area. Therefore, at this point, basic information and system understanding are urgently needed in order to identify which conservation efforts are most promising and which specific sites these efforts should focus on. Understanding the influence of prairie dog disturbance on vegetation parameters (taxonomic and functional), soil properties and their interaction with environmental conditions through time, will increase knowledge on the risks and vulnerability of grasslands, promoting solutions that can support grassland management. Furthermore, there is a clear gap in the literature about the varying effects of prairie dogs depending on different grassland types and environmental conditions. To study such conditions, I first classified and controlled for factors that could influence prairie dog ecosystem engineering effects addressing the following two general questions: (1) Which grassland types are present in GPCA El Tokio? (2) Do the different grassland types present in GPCA El Tokio influence prairie dog colony sizes and burrow density? When influencing factors were confirmed, I then assess the effects of prairie dog disturbance on vegetation and soil to answer the following questions: (3) Are there clear distinctions across GPCA El Tokio grassland environmental variables? (4) Do the different grassland types present in GPCA El Tokio influence prairie dog colony sizes and burrow density? To do so I selected a representative sample of sites covering the varying environmental conditions present in GPCA El Tokio grasslands, a data-driven clustering approach was used. Once clusters were defined, average burrow density was estimated from remote sensing imagery. To study vegetation measures I first, sampled plant species (92 species and 28 families and 6 traits) to obtain taxonomical and functional diversity measures, and obtained variables related to nutrient availability, carbon and climate regulation, and water regulation and purification. To study the effects on soil a total of 11 soil properties were obtained (382 samples were analyzed). Field work was done during the wet season and repeated during the dry season to obtain season variations of all measures. Soil measures were further obtained at depths of 0-10cm and 10 to 20cm. Results from the study classify GPCA El Tokio into 8 different grassland types with 4 clusters being the most dominant, named Agriculture, Arid, Calcareous, Mountain. I found agricultural grasslands had significantly lower colony sizes compared to calcareous grasslands when outliers were considered, but no differences when outliers were removed and burrow density varied depending on the grassland type, with mountain grasslands having a significantly lower burrow density compared to agricultural and calcareous grasslands. Regarding the effects of prairie dogs on vegetation and soil my findings suggest that functional metrics and community weighted mean (CWM) analyses responded to interactions between prairie dog disturbance, grassland type and season, whilst species diversity and cover measures were less sensitive to the role of prairie dog disturbance. I found weak evidence that prairie dog disturbance has a negative effect on vegetation structure, except for minimal effects on C4 and graminoid cover, but which depended mainly on season. Grassland type and season explained most of the effects on plant functional and taxonomic diversity as well as CWM traits. In the case of soil properties WOP grasslands tended to have higher nutrient availability than WP grasslands. However, mounds played a key role within WP grasslands. Mounds reduce compaction and increase nutrient levels of soil organic carbon, nitrogen, potassium, magnesium, and phosphorus. Such an effect was particularly present in calcareous and arid grasslands, and in many cases the effect was enhanced under wet conditions. In general, the effect of disturbance on soil property changes was dependent on environmental conditions. Overall, my study shows evidence that grassland type and season have a stronger effect than prairie dog disturbance on the vegetation of this short-grass, water-restricted grassland ecosystem.:3 TABLE OF CONTENTS Declaration of conformity 2 1 Acknowledgments 3 2 Extended Summary 5 4 List of Figures 11 5 List of Tables 13 1 Introduction 1 1.1 Importance of grasslands 1 1.2 Grassland conservation and ecosystem engineers 2 1.3 Prairie dogs: ecosystem engineers of grasslands in North America 3 1.4 Distribution of prairie dogs across North American grasslands 6 1.5 Current threats and conservation of the Mexican prairie dog C. mexicanus 7 1.6 Research questions, aims and objectives 9 2 Scientific background 13 2.1 Differences in grassland definitions and classifications 13 2.2 Dryland biases in grassland research 13 2.3 Research gaps on C. mexicanus 15 2.4 Functional diversity and disturbance 16 2.5 Soil complexity and limited research 17 3 Study Area: Grassland Priority Conservation Area of El Tokio 19 3.1 General overview 19 3.2 Bioregions, geology, topography and soils 20 3.3 Climate 23 3.4 Ecoregions, Land cover and Biodiversity 24 3.5 Anthropogenic activities and environmental issues 28 4 Assessing and classifying factors that could influence prairie dog ecosystem engineering effects 31 4.1 Data-driven identification of grassland types 33 4.1.1 Introduction 33 4.1.2 Methods 33 4.1.3 Results 36 4.1.4 Discussion and conclusion 39 4.2 C. mexicanus colony trends and differences across grassland types 41 4.2.1 Introduction 41 4.2.2 Methods 42 4.2.3 Results 48 4.2.4 Discussion and conclusion 52 5 Prairie dog disturbance effects on vegetation and soil across different grassland types and seasons 55 5.1 General methodology 57 5.1.1 Selection of grassland locations 57 5.1.2 Defining study sites and plots 58 5.1.3 Vegetation and soil sampling design 59 5.2 The effect of C. mexicanus on taxonomical and functional diversity across different seasons and grassland types 61 5.2.1 Introduction 61 5.2.2 Methods 63 5.2.3 Results 65 5.2.4 Discussion and conclusion 72 5.3 Soil property responses to the presence of C. mexicanus and its mounds across different grassland types, seasons and depths 77 5.3.1 Introduction 77 5.3.2 Methods 78 5.3.3 Results 81 5.3.4 Discussion and conclusion 88 6 Synthesis 95 6.1 Key findings 95 6.2 Management and Conservation opportunities 97 6.2.1 Designing management plans to suit existing environmental conditions 97 6.2.2 Functional diversity: an opportunity to identify patterns of vegetation across environmental conditions and scales. 98 6.3 Future research 99 6.3.1 Exploration of spatial and temporal effects 99 6.3.2 Mechanistic understanding of disturbance dynamics 100 6.3.3 A need for plant and soil ecological data 101 7 References 103 8 Appendix 131 8.1 Appendix 1. Prairie dog research reference list 132 8.2 Appendix 2. List of individual colonies with areas and other relevant information 140 8.3 Appendix 3. Historical and present colony data. 142 8.4 Appendix 4. Correlation results 144 8.5 Appendix 5. Top 3 models 146 8.6 Appendix 6. Species list per grassland type. 151 8.7 Appendix 7. Correspondence analysis (CA) for each grassland 154 8.8 Appendix 8. Post-hoc results 155 8.9 Appendix 9. Top 3 dominant species. 157 8.10 Appendix 10. HPD credible interval and ROPE. 159

No description available.

abstract: The plateau pika (Ochotona curzoniae), a small burrowing lagomorph that occupies the high alpine grassland ecosystems of the Qinghai-Tibetan Plateau in western China, remains a controversial subject among policymakers and researchers. One line of evidence points to pikas being a pest, which has led to massive attempts to eradicate pika populations. Another point of view is that pikas are a keystone species and an ecosystem engineer in the grassland ecosystem of the QTP. The pika eradication program raises a difficult ethical and religious dilemma for local pastoralists, and is criticized for not being supported by scientific evidence. Complex interactions between pikas, livestock, and habitat condition are poorly understood. My dissertation research examines underpinning justifications of the pika poisoning program leading to these controversies. I investigated responses of pikas to habitat conditions with field experimental manipulations, and mechanisms of pika population recovery following pika removal. I present policy recommendations based on an environmental ethics framework and findings from the field experiments. After five years of a livestock grazing exclusion experiment and four years of pika monitoring, I found that grazing exclusion resulted in a decline of pika habitat use, which suggests that habitat conditions determine pika population density. I also found that pikas recolonized vacant burrow systems following removal of residents, but that distances travelled by dispersing pikas were extremely short (~50 m). Thus, current pika eradication programs, if allowed to continue, could potentially compromise local populations as well as biodiversity conservation on the QTP. Lethal management of pikas is a narrowly anthropocentric-based form of ecosystem management that has excluded value-pluralism, such as consideration of the intrinsic value of species and the important ecological role played by pikas. These conflicting approaches have led to controversies and policy gridlock. In response, I suggest that the on-going large-scale pika eradication program needs reconsideration. Moderation of stocking rates is required in degraded pika habitats, and Integrated Pest Management may be required when high stocking rate and high pika density coexist. A moderate level of livestock and pika density can be consistent with maintaining the integrity and sustainability of the QTP alpine steppe ecosystem. / Dissertation/Thesis / Doctoral Dissertation Biology 2016

Enquanto em regiões temperadas o conhecimento sobre a relação solo-vegetação é consolidado, nos trópicos e subtrópicos é preliminar. É urgente a necessidade de se determinar os fatores abióticos que controlam padrões vegetacionais visando dar suporte a estudos de recuperação e conservação. O presente estudo analisa a relação entre fatores abióticos e vegetação campestre na forma de três artigos científicos (capítulos I, II e III) e um artigo de revisão (capítulo IV). Nos primeiros três artigos, analisou-se a variância da vegetação em diferentes escalas ecológicas; e no artigo de revisão, discutiu-se de forma aplicada a variância de fatores bióticos e abióticos em resposta à degradação. Dessa forma, a presente tese objetivou responder as seguintes questões: (I) Que diferenças podem ser observadas dentro de uma mesma espécie quanto à funcionalidade e suas estratégias de alocação sob diferentes graus de disponibilidade de recursos?; (II) Quão variáveis são as propriedades físicas e químicas do solo em diferentes escalas espaciais; e existem propriedades dos solos que podem explicar com maior precisão a distribuição das espécies em diferentes escalas espaciais? (III) Que porcentagem de variância da vegetação pode ser explicada por propriedades pedológicas e climáticas; e quais características de solo e clima melhor explicam esses padrões de vegetação? No capítulo IV é apresentado um modelo conceitual sobre degradação dos campos e sua aplicação aos campos do Rio Grande do Sul (RS). Para responder as questões acima usei dados ao nível de espécie de campos calcáreos da Alemanha (capítulo I); dados ao nível de comunidade em seis áreas campestres do Rio Grande do Sul, sul do Brasil (capítulos II e III); e através da revisão de literatura relacionada à degradação, quanto à capacidade de recuperação dos campos do RS (capítulo IV). Os resultados evidenciaram que: (I) dentro de espécies ficaram evidentes duas estratégias frente à limitação de recursos, enquanto a resposta dos atributos aos diferentes tratamentos se mostrou constante; (II) a variação dos parâmetros do solo relaciona-se à escala espacial aplicada e a variância da vegetação geralmente responde a diferentes parâmetros de solo em diferentes escalas; (III) 45% da variância da vegetação entre biomas nos campos do RS foi explicada por características pedológicas e climáticas, sendo em grande parte governada pela precipitação anual e a porcentagem de saturação por alumínio do solo; e (IV) o modelo conceitual apresenta variações ao longo de dois eixos (biótico e abiótico) e poderá servir de suporte a estudos de conservação e recuperação de campos tropicais e suptropicais, bem como facilitar a tomada de decisões quanto ao manejo e conservação. Como conclusão geral, verificou-se que a vegetação campestre responde a variações ambientais em diferentes escalas espaciais e pode adotar diferentes estratégias para sobrepor filtros ambientais e processos de degradação. O entendimento da relação entre a vegetação e o meio abiótico é de grande importância para tomada de decisões quanto ao emprego de formas alternativas de manejo e conservação. / Whereas in temperate regions the abiotic-biotic relationship is well-known, in the tropics and subtropics our understanding is still preliminary. There is an urgent need to determine abiotic factors that control vegetation patterns in order to give support to restoration and conservation approaches. The present thesis analyses the relationship between abiotic factors and grassland vegetation in three original research papers (chapters I, II and III) and a review paper (chapter IV). In the first three papers, vegetation variance in response to abiotic factors was analyzed at different ecological scales; and in the fourth, the variance in biotic and abiotic factors in response to degradation process was discussed with a more applied view. Thus in this thesis the aim is to answer the following questions: (I) Which differences can be found in functional plant traits and allocation strategies within species at different levels of water and nutrient availability?; (II) How variable are physical and chemical parameters in different spatial scales; and are there soil parameters that can more accurately explain plant distribution in different spatial scales? (III) How much of RS grassland vegetation variance can be explained by soil and climatic properties; and which climatic and soil properties better explain these vegetation patterns? In chapter IV a conceptual model of grassland degradation is presented and applied to Rio Grande do Sul (RS) grasslands. To address these questions I used species-level data in a calcareous grassland in Germany (chapter I); community-level data in six sites in RS, South Brazilian grasslands (chapter II and III); and a review of literature studies concerning RS grassland degradation and restorability (chapter IV). The results showed that: (I) at a intraspecific level, the study species showed two allocation strategies in relation to resource stress, while the responses of individual traits to the soil treatments were consistent across species; (II) soil parameters variation are related to the measurement scale applied and the vegetation variance often responds to different soil parameters at different scales; (III) climatic and soil properties explained 45% of vegetation variance between biomes in RS grasslands and the main factors controlling its variance are annual precipitation and percent aluminum saturation; and (IV) the conceptual model is displayed as biotic and abiotic changes along the axes and can serve as a general framework to study degradation and restorability of tropical and subtropical grasslands, and further it may facilitate decisions on alternative management and conservation. As a general conclusion, the grassland vegetation responds to changes in the environment in different scales and may use different strategies to overcome environmental selective forces and degradation process. The understanding of this relationship is of high importance to facilitate decisions on alternative management and conservation.

Enquanto em regiões temperadas o conhecimento sobre a relação solo-vegetação é consolidado, nos trópicos e subtrópicos é preliminar. É urgente a necessidade de se determinar os fatores abióticos que controlam padrões vegetacionais visando dar suporte a estudos de recuperação e conservação. O presente estudo analisa a relação entre fatores abióticos e vegetação campestre na forma de três artigos científicos (capítulos I, II e III) e um artigo de revisão (capítulo IV). Nos primeiros três artigos, analisou-se a variância da vegetação em diferentes escalas ecológicas; e no artigo de revisão, discutiu-se de forma aplicada a variância de fatores bióticos e abióticos em resposta à degradação. Dessa forma, a presente tese objetivou responder as seguintes questões: (I) Que diferenças podem ser observadas dentro de uma mesma espécie quanto à funcionalidade e suas estratégias de alocação sob diferentes graus de disponibilidade de recursos?; (II) Quão variáveis são as propriedades físicas e químicas do solo em diferentes escalas espaciais; e existem propriedades dos solos que podem explicar com maior precisão a distribuição das espécies em diferentes escalas espaciais? (III) Que porcentagem de variância da vegetação pode ser explicada por propriedades pedológicas e climáticas; e quais características de solo e clima melhor explicam esses padrões de vegetação? No capítulo IV é apresentado um modelo conceitual sobre degradação dos campos e sua aplicação aos campos do Rio Grande do Sul (RS). Para responder as questões acima usei dados ao nível de espécie de campos calcáreos da Alemanha (capítulo I); dados ao nível de comunidade em seis áreas campestres do Rio Grande do Sul, sul do Brasil (capítulos II e III); e através da revisão de literatura relacionada à degradação, quanto à capacidade de recuperação dos campos do RS (capítulo IV). Os resultados evidenciaram que: (I) dentro de espécies ficaram evidentes duas estratégias frente à limitação de recursos, enquanto a resposta dos atributos aos diferentes tratamentos se mostrou constante; (II) a variação dos parâmetros do solo relaciona-se à escala espacial aplicada e a variância da vegetação geralmente responde a diferentes parâmetros de solo em diferentes escalas; (III) 45% da variância da vegetação entre biomas nos campos do RS foi explicada por características pedológicas e climáticas, sendo em grande parte governada pela precipitação anual e a porcentagem de saturação por alumínio do solo; e (IV) o modelo conceitual apresenta variações ao longo de dois eixos (biótico e abiótico) e poderá servir de suporte a estudos de conservação e recuperação de campos tropicais e suptropicais, bem como facilitar a tomada de decisões quanto ao manejo e conservação. Como conclusão geral, verificou-se que a vegetação campestre responde a variações ambientais em diferentes escalas espaciais e pode adotar diferentes estratégias para sobrepor filtros ambientais e processos de degradação. O entendimento da relação entre a vegetação e o meio abiótico é de grande importância para tomada de decisões quanto ao emprego de formas alternativas de manejo e conservação. / Whereas in temperate regions the abiotic-biotic relationship is well-known, in the tropics and subtropics our understanding is still preliminary. There is an urgent need to determine abiotic factors that control vegetation patterns in order to give support to restoration and conservation approaches. The present thesis analyses the relationship between abiotic factors and grassland vegetation in three original research papers (chapters I, II and III) and a review paper (chapter IV). In the first three papers, vegetation variance in response to abiotic factors was analyzed at different ecological scales; and in the fourth, the variance in biotic and abiotic factors in response to degradation process was discussed with a more applied view. Thus in this thesis the aim is to answer the following questions: (I) Which differences can be found in functional plant traits and allocation strategies within species at different levels of water and nutrient availability?; (II) How variable are physical and chemical parameters in different spatial scales; and are there soil parameters that can more accurately explain plant distribution in different spatial scales? (III) How much of RS grassland vegetation variance can be explained by soil and climatic properties; and which climatic and soil properties better explain these vegetation patterns? In chapter IV a conceptual model of grassland degradation is presented and applied to Rio Grande do Sul (RS) grasslands. To address these questions I used species-level data in a calcareous grassland in Germany (chapter I); community-level data in six sites in RS, South Brazilian grasslands (chapter II and III); and a review of literature studies concerning RS grassland degradation and restorability (chapter IV). The results showed that: (I) at a intraspecific level, the study species showed two allocation strategies in relation to resource stress, while the responses of individual traits to the soil treatments were consistent across species; (II) soil parameters variation are related to the measurement scale applied and the vegetation variance often responds to different soil parameters at different scales; (III) climatic and soil properties explained 45% of vegetation variance between biomes in RS grasslands and the main factors controlling its variance are annual precipitation and percent aluminum saturation; and (IV) the conceptual model is displayed as biotic and abiotic changes along the axes and can serve as a general framework to study degradation and restorability of tropical and subtropical grasslands, and further it may facilitate decisions on alternative management and conservation. As a general conclusion, the grassland vegetation responds to changes in the environment in different scales and may use different strategies to overcome environmental selective forces and degradation process. The understanding of this relationship is of high importance to facilitate decisions on alternative management and conservation.