Magister Scientiae (Biodiversity and Conservation Biology) - MSc (Biodiv and Cons Biol) / The Succulent Karoo Biome (SKB) in South Africa is widely reputed to house Earth’s greatest diversity of succulent plants. It is also famous for spectacular displays of annual flowers after good rains. The area experiences winter rainfall which infrequently exceeds 100 mm per annum but certain parts of the SKB can get 250 mm. Irrigated agriculture on a large scale was therefore not a viable option when European farmers began colonizing the land. The land was conquered from the indigenous Khoekhoe herders and San hunter-gatherers, South Africa’s first peoples. The biome underwent extreme transformation in the last 200 years following colonisation which resulted in homogenization of the landscape and extinction of many succulents thus reducing biodiversity.

A considerable part of the global pool of terrestrial carbon is stored in high latitude soils. In these soils, repeated cycles of freezing and thawing creates soil motion (cryoturbation) that in combination with other cryogenic disturbance processes may play a profound role in controlling the carbon balance of the arctic soil. Conditions for cryogenic soil processes are predicted to dramatically change in response to the ongoing climate warming, but little is known how these changes may affect the ability of arctic soils to accumulate carbon. In this thesis, I utilize a patterned ground system, referred to as non-sorted circles, as experimental units and quantify how cryogenic soil processes affect plant communities and carbon fluxes in arctic soils. I show that the cryoturbation has been an important mechanism for transporting carbon downwards in the studied soil over the last millennia. Interestingly, burial of organic material by cryoturbation appears to have mainly occurred during bioclimatic events occurring around A.D. 900-1250 and A.D. 1650-1950 as indicated by inferred 14C ages. Using a novel photogrammetric approach, I estimate that about 0.2-0.8 % of the carbon pool is annually subjected to a net downward transport induced by the physical motion of soil. Even though this flux seems small, it suggests that cryoturbation is an important transporter of carbon over centennial and millennial timescales and contributes to translocate organic matter to deeper soil layers where respiration proceeds at slow rates. Cryogenic processes not only affect the trajectories of the soil carbon, but also generate plant community changes in both species composition and abundance, as indicated by a conducted plant survey on non-sorted circles subjected to variable differential frost heave during the winter. Here, disturbance-tolerant plant species, such as Carex capillaris and Tofieldia pusilla, seem to be favoured by disturbance generated by the differential heave. Comparison with findings from a previous plant survey on the site conducted in the 1980s suggest that the warmer temperatures during the last decades have resulted in decreased differential heave in the studied non-sorted circles. I argue that this change in cryogenic activity has increased abundance of plants present in the 1980s. The fact that the activity and function of the non-sorted circles in Abisko are undergoing changes is further supported by their contemporary carbon dioxide (CO2) fluxes. Here, my measurements of CO2 fluxes suggest that all studied non-sorted circles act as net CO2 sources and thus that the carbon balance of the soils are in a transition state. My results highlight the complex but important relationship between cryogenic soil processes and the carbon balance of arctic soils.

Peatland ecosystems only cover 2-3 % of the Earth‟s surface but they represent significant carbon stores, holding approximately one third of the global soil carbon (C). The major peat forming genera Sphagnum appears to be highly sensitive to increased N availability. Many studies have shown decreased productivity of Sphagnum which could lead to a decrease in the amount of C stored, especially as many studies also show an increase in the decomposition rate with higher N deposition. However, the overall effects of N on CO2 fluxes of Sphagnum remain unclear. The present study aimed to look at the effects of increased N on Sphagnum productivity, decomposition and CO2 fluxes after long-term N additions (> 5 years) using a field experiment at Whim Moss in southern Scotland where N deposition has been manipulated employing a very realistic application coupled to rainfall since 2002. The experiment also has treatments with PK addition to test the effects of removing P and/or K-limitation. Measurements of plant tissue nutrient concentrations, visual assessments of Sphagnum viability, and pore water analysis were also carried out. Nitrogen additions increased tissue N, and decreased Sphagnum shoot extension and productivity. Simultaneous P and K additions alleviated the effects of N on tissue N concentrations and growth, although this was only significant for shoot extension. Visual assessments correlated well with tissue chemistry and productivity; the decline in health was associated with high %N and reduced productivity. Interestingly, in the present study increased N decreased the mass loss and again when PK was added with N decomposition rates were more similar to the control. With respect to the carbon balance of the site and the sustainability of peatlands the results suggest that the negative effect of N on C assimilation may be partially offset by the reduced decomposition rates. The CO2 measurements showed a large loss of C as CO2 from all the Sphagnum plots which was exacerbated by adding N especially when the air temperature increased. The positive temperature response of ecosystem respiration with N additions suggests that in high N deposition areas climate change and subsequent temperature rises will increase C losses from bogs.

Planting trees on agricultural land either as farm woodlands or agroforestry (trees integrated with farming) is one option for reducing the level of atmospheric carbon dioxide. Trees store carbon as biomass, and may increase carbon storage in the ground. A review of the literature outlined uncertainty relating to changes in carbon storage after planting trees on agricultural land. The aim of this thesis is to deter¬mine the impact of tree planting on arable and pasture land in terms of above and belowground carbon storage and thereby address these uncertainties, and assess the implications for the Woodland Carbon Code: a voluntary standard for carbon storage in UK woodlands. Measurements of soil organic carbon to a depth of 1.5 m were taken at two field sites in Bedfordshire in the UK: a 19 year old silvoarable trial, and a 14 year old silvopasture and farm woodland. On average 60% and 40% of the soil carbon (rel¬ative to 1.5 m) was found beneath 0.2 and 0.4 m in depth respectively. Whilst tree planting in the arable system showed gains in soil organic carbon (12.4 t C ha−1 at 0–40 cm), tree planting in the pasture was associated with losses of soil organic carbon (6.1–13.4 t C ha−1 at 0–10 cm). Evidence from a nearby mature grazed woodland indicate that these losses may be recovered. No differences associated with tree planting were found to the full 1.5 m, though this may be due to a lack of statistical power. Measurements of above and belowground biomass, and the root distribution of 19 year old poplar (Populus spp.) trees (at the silvoarable trial) and ash (Fraxinus excelsior) trees ranging from 7 to 21 years (at several field sites across Bedfordshire) were made, involving the destructive harvest of 48 trees. These measurements suggest that Forestry Commission yield tables overestimate yield for poplar trees grown in a silvoarable system. An allometric relationship for determining ash tree biomass from diameter measurements was established. The biophysical model Yield-SAFE was updated to take into account root growth, and was parameterised using field measurements. It was successfully used to describe existing tree growth at two sites, and was then used to predict future biomass carbon storage at the silvoarable trial. Measurements indicate that losses in soil carbon at relatively shallow depths can offset a large proportion of the carbon stored in tree biomass, but assessing changes on a site by site basis may be prohibitively expensive for schemes such as the Woodland Carbon Code.

La rapide augmentation des gaz à effet de serre (GES) dans l’atmosphère - dont le CO2 – due aux activités humaines est considérée comme responsable des changements climatiques en cours et futurs. Les écosystèmes terrestres sont potentiellement des "puits" importants de C et pourraient contribuer à l'atténuation des GES. Les prairies permanentes (steppes, savanes, prairies de montagne, ...) couvrent 40% de la surface terrestre (hors calotte glaciaire) et leurs sols représentent potentiellement un énorme "puits" permettant de stocker du C naturellement (GIEC 2001). Cependant, les processus impliqués et leur régulation restent à préciser. L’objectif de la thèse était d’analyser l’effet des pratiques de pâturage sur le stockage de C dans le sol. Cette analyse a été réalisée à partir de données acquises sur deux dispositifs ‘long terme’ en prairie permanente (SOERE ACBB) et en s’intéressant aux flux de C entre les différents compartiments de l’agro-écosystème sous différentes intensités de pâturage afin i) d’étudier notre capacité à estimer le stockage de C dans le sol après 10 ans d’application de traitement, en comparant deux méthodes (méthode utilisant des tours à flux et mesure du stock de C du sol) ii) d’apporter des connaissances sur les mécanismes et régulations agissant sur les dynamiques de stockage du C. Les résultats de la comparaison des deux méthodes de mesures testées ont indiqué une séquestration nette de C dans le sol, avec un taux de séquestration moyen mesuré avec les deux méthodes de 2.21 t C ha-1 an-1 et de 2.29 t C ha-1 an-1, sans différence significative entre traitements, mais avec une tendance à une séquestration plus élevée avec la gestion plus intensive. Chaque méthode permet d’accéder à des informations différentes. L'approche avec les tours à flux permet d'identifier des interactions entre le climat et les pratiques de gestion sur les flux de C dans les prairies. Les inventaires de sol ont permis de montrer que le carbone se stocke également dans les couches plus profondes de sol. Alors que les communautés végétales ont évolué sous l’effet des traitements différenciés de pâturage, les mesures ne montrent pas d’évolution des stocks de C totaux ni des matières organiques particulaires. L’analyse des flux de C entre les différents compartiments de l'écosystème, après 7 ans d’application des traitements, montre que les traitements avec une intensité faible ou nulle ont conduit à une réduction des flux de carbone entre les compartiments du continuum de dégradation du C, tandis que les stocks de carbone des racines et des POM ne sont pas affectés par les traitements. Une étude complémentaire conduite pour estimer les productions racinaires indique que la réponse des racines (stocks et production) et des stocks de matières organiques particulaires pourrait être en partie découplée de la réponse du compartiment aérien de la végétation. A l’issue de cette étude, il nous apparaît qu’une approche plus intégrative du fonctionnement de l’écosystème est nécessaire pour accroître notre capacité de prédiction de l’impact des pratiques sur le stockage du C en prairie. / The fast increase of greenhouse gases in the atmosphere, such as carbon dioxide, due to human activities is consider as the main cause of actual climate change. Terrestrial ecosystem are considered as a huge "sink" of C and may contribute to decrease greenhouse gases. Permanent grasslands cover 40% of land and their soil may contribute to sequester C (GIEC 2001). However, the processes involved and their regulations remain to be specified. The aim of the thesis was to analyze the effect of grazing management on soil C storage. This analysis was made from data acquired on two long term permanent grassland sites (SOERE ACBB) and by studying C fluxes between the different agroecosystem compartments under different grazing intensities for i) estimating our capacity to measure soil C storage after 10 years of grazing treatments by comparing two methods (soil inventories vs net carbon storage measurements), ii) to provide knowledge on the mechanisms and regulations affecting the dynamics of soil C sequestration. Comparing results of both methods, measurements indicated a net C sequestration in soil, with an average sequestration rate of 2.21 t C ha-1 yr-1 and 2.29 t C ha-1 year-1 and no significant difference between treatment but a tendency to a higher sequestration with more intensive management. Each method provides access to different information. The approach with flux towers allows a better understanding of the role and interactions between climate and practices on C fluxes in grasslands. Soil inventories showed carbon is store in deeper soil layers. While plant communities have evolved as a result of differentiated grazing treatments, measurements show no changes in total C stocks and particulate organic matter. Analysis of C fluxes after 7 years of differentiate grazing treatments, showed that treatments with low or zero grazing intensity led to a reduction of carbon fluxes between the compartments of the continuum of degradation, while carbon stocks in roots and POM were not affected by treatments. A complementary study conducted to estimate root production indicates that the response of roots (stocks and production) and stocks of particulate organic matter may be partly decoupled from the response of the aerial vegetation compartment. This study indicates that a more integrative approach on ecosystem functioning is necessary to increase our ability to predict the impact of management practices on C storage in grassland.

Tropical savannas and woodlands are the dominant vegetation cover in Southern Africa covering 4 million km2. Their large spatial extent means they are potentially a globally important store of biomass carbon with implications for global climate, and an area of high biodiversity value. They provide natural resources such as food, fuel and timber that help sustain the livelihoods of over 100 million people. The ability of these savanna woodlands to maintain these important ecological functions is under question due to increases in land use and land cover change. This thesis addresses a set of science questions aimed at (i) improving our knowledge of the amount of carbon and biodiversity stored in these ecosystems and how they co-vary, (ii) how these variables are spatially distributed at landscape scales and the factors which underlie these patterns, and (iii) how they respond over time to human disturbance. In Chapter 2 I examine how patterns in aboveground woody carbon storage (AGC) are linked to differences in forest structure, tree species diversity and floristic composition across a recently established network of 25 permanent sample plots in south-east Tanzania. Large stems were a significant contributor to plot-level AGC stocks with the top 3% of individuals (>40cm) in terms of size containing 35% of the total measured C. This data can potentially be used to simplify future measurements of biomass in these systems. Tree species diversity was positively related to AGC indicating the potential to align forest conservation efforts. The linear relationship suggests a functional relationship between the variables and is consistent with ecological theory on niche complementarity and selection effects, however based on the available data the mechanisms underlying this relationship can only be theorised. Changes in tree species composition were also noted across plots with differences in vegetation structure between plots explaining 16% of the variation in composition, with environmental differences related to climate and soils explaining only 3%. In Chapter 3, the focus shifts to understanding larger-scale spatial patterns in AGC. Field plots are spatially limited in this regard, therefore radar remote sensing data was used to generate a map of AGC in order to improve our knowledge on what principally controls its spatial variability at landscape scales. Results showed that factors related topography, climate and soils explained very little of the variation in C stocks across the landscape (r2 = 15 – 20%). Differences in slope angle and topographic position were important in discriminating between low biomass savannas and moderate biomass woodlands, while differences in annual precipitation were more important in separating woodlands and denser forests. A large proportion of the variation in C stocks (~80%) was unexplained highlighting the role of unmeasured variables. It is suggested that fire may play a key role in shaping patterns in tree species composition and C stocks across these landscapes. This data has important implications for a local REDD+ project which is aiming to generate carbon credits through improved fire management. In the second part of the thesis the attention shifts to understanding the long-term ecological impacts of shifting cultivation and the sensitivity and resilience of these woodlands to anthropogenic change. In Chapter 4 I examined how carbon stored in trees and soils recover across a 40-year chronosequence of abandoned agricultural land, and how this patchy disturbance impacts spatial pattern in tree species composition and diversity. I show that re-growing woodlands can act as carbon sinks through the accumulation of woody biomass (0.83 tC ha-1 yr-1), with soil texture having no clear impact on accumulation rates. Re-growing woodlands were also found to contain considerable biodiversity value by promoting novel species assemblages. Bulk soil carbon stocks appeared to be largely unaffected by the full cycle of shifting cultivation. However in Chapter 5 I show evidence of a previously unquantified legacy effect of land clearance on soil CO2 production with more recently abandoned fields (c. 6 years) exhibiting significantly higher efflux rates than the older abandonments (15 -25 years) and mature woodlands. Total soil nitrogen was the most important predictor of soil respiration across the plots (r2 = 0.3) followed by fine root density (r2 = 0.12). Soils in the younger sites were found to be more nitrogen rich which was used to explain the greater CO2 fluxes in these areas, however, it is still unclear why this pattern exists. The thesis concludes by discussing the wider implications of the results, as well as outlining further work needed to solidify some of the conclusions drawn in this thesis.

Carbon Capture and Storage (CCS) is a suite of technologies available to directly reduce carbon dioxide (CO2) emissions to the atmosphere from fossil fuelled power plants and large industrial point sources. For a safe deployment of CCS it is important that CO2 injected into deep geological formations does not migrate out of the storage site. Characterising and understanding possible migration mechanisms and pathways along which migration may occur is therefore crucial to ensure secure engineered storage of anthropogenic CO2. In this thesis naturally occurring CO2 accumulations in the subsurface are studied as analogue sites for engineered storage sites with respect to CO2 migration pathways and mechanisms that ensure the retention of CO2 in the subsurface. Geological data of natural CO2 reservoirs world-wide has been compiled from published literature and analysed. Results show that faults are the main pathways for migration of CO2 from subsurface reservoirs to the surface and that the state and density of CO2, pressure of the reservoir, and thickness of the caprock influence the successful retention of CO2. Gaseous, low density CO2, overpressured reservoirs, and thin caprocks are characteristics of insecure storage sites. Two natural CO2 reservoirs have been studied in detail with respect to their fault seal properties. This includes the first study of how fault rock seals behave in CO2 reservoirs. It has been shown that the bounding fault of the Fizzy Field reservoir in the southern North Sea can with hold the amount of CO2 trapped in the reservoir at current time. A initially higher gas column would have led to across fault migration of CO2 as the fault rock seals would not have been able to withhold higher pressures. Depending on the present day stress regime the fault could be close to failure. At the natural CO2 reservoir of St. Johns Dome, Arizona, migration of CO2 to the surface has been occurring for at least the last 500 ka. Fault seal analysis shows that this migration is related to the fault rock composition and the orientation of the bounding fault in the present day stress field. Using the U-Th disequilibrium method the ages of travertine deposits of the St. Johns Dome area have been determined. The results illustrate that along one fault CO2 migration took place for at least 480 ka and that individual travertine mounds have had long lifespans of up to ~350 ka. Age and uranium isotope trends along the fault have been interpreted as signs of a shrinking CO2 reservoir. The amount of CO2 calculated to have migrated out of the St. Johns Dome is up to 113 Gt. Calculated rates span from 5 t/yr to 30,000 t/yr and indicate that at the worst case large amounts of CO2 can migrate rapidly from the subsurface reservoir along faults to the surface. This thesis highlights the importance of faults as fluid pathways for vertical migration of CO2. It has been also shown that they can act as baffles for CO2 migration and that whether a fault acts as pathway or baffle for CO2 can be predicted using fault seal analysis. However, further work is needed in order to minimise the uncertainties of fault seal analysis for CO2 reservoirs.

abstract: Rangelands are an extensive land cover type that cover about 40% of earth’s ice-free surface, expanding into many biomes. Moreover, managing rangelands is crucial for long-term sustainability of the vital ecosystem services they provide including carbon (C) storage via soil organic carbon (SOC) and animal agriculture. Arid rangelands are particularly susceptible to dramatic shifts in vegetation cover, physical and chemical soil properties, and erosion due to grazing pressure. Many studies have documented these effects, but studies focusing on grazing impacts on soil properties, namely SOC, are less common. Furthermore, studies testing effects of different levels of grazing intensities on SOC pools and distribution yield mixed results with little alignment. The primary objective of this thesis was to have a better understanding of the role of grazing intensity on arid rangeland soil C storage. I conducted research in long established pastures in Jornada Experimental Range (JER). I established a 1500m transect in three pastures originating at water points and analyzed vegetation cover and SOC on points along these transects to see the effect of grazing on C storage on a grazing gradient. I used the line-point intercept method to measure and categorize vegetation into grass, bare, and shrub. Since soil adjacent to each of these three cover types will likely contain differing SOC content, I then used this vegetation cover data to calculate the contribution of each cover type to SOC. I found shrub cover and total vegetation cover to decrease, while grass and bare cover increased with decreasing proximity to the water source. I found areal (g/m2) and percent (go SOC to be highest in the first 200m of the transects when accounting for the contribution of the three vegetation cover types. I concluded that SOC is being redistributed toward the water source via foraging and defecation and foraging, due to a negative trend of both total vegetation cover and percent SOC (g/g). With the decreasing trends of vegetation cover and SOC further from pasture water sources, my thesis research contributes to the understanding of storage and distribution of SOC stocks in arid rangelands. / Dissertation/Thesis / Masters Thesis Biology 2018

Rising atmospheric CO2 concentrations have motivated efforts to better quantify reservoirs and fluxes of Earth’s carbon. Of these fluxes from the atmosphere, one that has received relatively little attention is the atmospheric carbon sink associated with carbonate mineral dissolution. Osterhoudt (2014) and Salley (2016) explored new normalization techniques to improve and standardize a process for measuring this flux over large river basins. The present research extends this work to the 490,600 km2 Ohio River drainage basin and 11 subbasins. The study estimated the DIC flux leaving these basins between October 1, 2013, and September 30, 2014, based on secondary hydrogeochemical, geologic, and climatic data. The total annual DIC flux for the Ohio River basin was estimated to be 7.54 x 1012 g carbon (C). The time-volume normalized value of DIC flux for the Ohio basin was 3.36 x 108 g C/km3 day, where the km3 refers to the amount of water available during the year. This was within 71.4% agreement with the Barren River data (Salley, 2016) and within 63.9% agreement with the Green River data (Osterhoudt, 2014). In general, normalized DIC flux values of sub-basins containing at least modest amounts (more than 8%) of exposed carbonates (Tennessee, Cumberland, Green, Kentucky, Licking, Monongahela, and Allegheny) were in strong agreement with the normalized DIC flux of the Ohio River basin, whereas inclusion of basins with little or no near surface carbonates (Wabash, Great Miami, Scioto and Kanawha) yielded poor agreement. Regression analysis yielded strong agreement between DIC flux and the normalization parameters for the carbonate-bearing sub-basins (R2 = 0.97, p =

The boreal forest is one of the worlds largest forest biomes and comprises a major portion of the terrestrial carbon (C) sink. Quantifying the net C change in forest ecosystems is an important step in understanding and modeling the global C cycle. The goals of this project were: to estimate and compare the total change in ecosystem C over a 10-year period in two boreal forest stands using biometric and eddy-covariance approaches, and to evaluate the year-to-year changes in C uptake. This study utilized 10 years of eddy-covariance data and ecosys model data from the Old Aspen (OA) and Old Jack Pine (OJP) sites in central Saskatchewan, part of the Boreal Ecosystem Research and Monitoring Sites (BERMS). According to the eddy-covariance and C stock approaches, between 1994 and 2004 the net change in C storage at OA was 15.6 ± 4.0 and 18.2 ± 8.0 Mg C ha-1, respectively. At OJP, the 10-year net change in C storage from eddy-covariance was 5.8 ± 2.0 Mg C ha-1 in comparison to 6.9 ± 1.6 Mg C ha-1 from the carbon stock approach. While both sites were sinks of C between 1994 and 2004, the greatest increase in C occurred in different components - the forest floor at OA (14.6 Mg C ha-1) and in the living vegetation at OJP (8.0 Mg C ha-1). In 2004, total ecosystem C content was greater at OA (180.6 Mg C ha-1) than OJP (78.9 Mg C ha-1), with 50% (OA) and 39% (OJP) of the C in the detritus and mineral soil pools. During the 10-year period of eddy-covariance measurements, there was a positive correlation between both annual and growing season gross ecosystem photosynthesis (GEP) and live stem C biomass increment at OA, whereas no significant relationships were found at OJP. Stem C increment accounted for 30% of total net primary productivity (NPP) at both sites, and NPP/GEP ratios were 0.36 and 0.32 at OA and OJP, respectively. Overall, this study found good agreement between eddy-covariance and biometric estimates of ecosystem C change at OA and OJP between 1994 and 2004. Over that period at OA, eddy-covariance estimates of photosynthesis captured the inter-annual variability in C uptake based on the growth of tree rings.