Encroaching species are stronger anisohydric “water spenders” under elevated CO2 conditions: implications for savanna seedling establishment rates
- Authors: Reynolds, Liam Macleod
- Date: 2024-10-11
- Subjects: Xylem , Scrub encroachment , Isotopes , Photosynthesis , Plants Effect of atmospheric carbon dioxide on
- Language: English
- Type: Academic theses , Master's theses , text
- Identifier: http://hdl.handle.net/10962/465091 , vital:76572
- Description: Plant water transport systems play a fundamental role in the productivity and survival of terrestrial plants due to the vascular architecture placing a physical limit on metabolic function. Savannas have high variability in rainfall, leading multiple studies to suggest that plant water-use strategies are key mechanisms affecting seedling establishment rates. Many savannas are seeing a directional shift towards an increase in the abundance of certain woody species through a process known as bush encroachment, which has been largely attributed to the fertilising effect of rising atmospheric [CO2] on C3 trees. These species are classified as encroachers. While there have been multiple studies investigating changes in the physiology of savanna species under elevated CO2 (eCO2), few have examined how climate and eCO2 affects the fundamental water-use strategies in the seedling stage, a crucial demographic bottleneck. Here, this research provides valuable insights into the mechanisms behind bush encroachment in the context of eCO2 using results from a pot experiment at the Rhodes University Elevated CO2 facility and a field experiment. All species showed water use strategies characteristic of anisohydric “water-spenders”, however, the vulnerability to embolism and rates of water-use were different between encroachers and non-encroachers. Encroachers are better at taking advantage of water pulses, particularly under eCO2 and grass competition. This comes at the cost of higher xylem vulnerability during drought, leading to reductions in conductance when exposed to heavy water stress. The response of the photosynthetic parameters mirrored this, with encroaching species had higher rates of photosynthesis and photosystem II quantum yield than non-encroachers under the well-watered treatments. Field experiments revealed that small trees are particularly vulnerable to drought stress, when compared to medium and large trees. The outcomes of this complex response will largely depend on the extent of changes to biotic and abiotic factors across spatial and temporal zones caused by climate change. This research highlights potential hydraulic mechanisms contributing to the increase in bush encroachment, as well as providing important insights into the determinant factors that make a savanna species capable of encroachment. , Thesis (MSc) -- Faculty of Science, Botany, 2024
- Full Text:
- Authors: Reynolds, Liam Macleod
- Date: 2024-10-11
- Subjects: Xylem , Scrub encroachment , Isotopes , Photosynthesis , Plants Effect of atmospheric carbon dioxide on
- Language: English
- Type: Academic theses , Master's theses , text
- Identifier: http://hdl.handle.net/10962/465091 , vital:76572
- Description: Plant water transport systems play a fundamental role in the productivity and survival of terrestrial plants due to the vascular architecture placing a physical limit on metabolic function. Savannas have high variability in rainfall, leading multiple studies to suggest that plant water-use strategies are key mechanisms affecting seedling establishment rates. Many savannas are seeing a directional shift towards an increase in the abundance of certain woody species through a process known as bush encroachment, which has been largely attributed to the fertilising effect of rising atmospheric [CO2] on C3 trees. These species are classified as encroachers. While there have been multiple studies investigating changes in the physiology of savanna species under elevated CO2 (eCO2), few have examined how climate and eCO2 affects the fundamental water-use strategies in the seedling stage, a crucial demographic bottleneck. Here, this research provides valuable insights into the mechanisms behind bush encroachment in the context of eCO2 using results from a pot experiment at the Rhodes University Elevated CO2 facility and a field experiment. All species showed water use strategies characteristic of anisohydric “water-spenders”, however, the vulnerability to embolism and rates of water-use were different between encroachers and non-encroachers. Encroachers are better at taking advantage of water pulses, particularly under eCO2 and grass competition. This comes at the cost of higher xylem vulnerability during drought, leading to reductions in conductance when exposed to heavy water stress. The response of the photosynthetic parameters mirrored this, with encroaching species had higher rates of photosynthesis and photosystem II quantum yield than non-encroachers under the well-watered treatments. Field experiments revealed that small trees are particularly vulnerable to drought stress, when compared to medium and large trees. The outcomes of this complex response will largely depend on the extent of changes to biotic and abiotic factors across spatial and temporal zones caused by climate change. This research highlights potential hydraulic mechanisms contributing to the increase in bush encroachment, as well as providing important insights into the determinant factors that make a savanna species capable of encroachment. , Thesis (MSc) -- Faculty of Science, Botany, 2024
- Full Text:
Spatial pattern analysis of thicket expansion in a semi-arid savanna
- Authors: Putzier, Rachel Rayne
- Date: 2024-10-11
- Subjects: Arid regions South Africa Eastern Cape , Spatial analysis (Statistics) , Thicket , Lidar , Cluster analysis , Trees Mortality , Scrub encroachment
- Language: English
- Type: Academic theses , Master's theses , text
- Identifier: http://hdl.handle.net/10962/464484 , vital:76515
- Description: Woody thickening has negative economic and ecological impacts in savannas globally. While the increase of savanna trees as a form of bush encroachment has been well studied, less is known about the increase of thicket species in savannas, which is an important phenomenon resulting in the formation of closed-canopy clumps which may exclude the grass layer over time. The early stage of thicket expansion is often initiated by a nucleating savanna species which facilitates the establishment of woody thicket pioneer species, and as the thicket clump formation expands, bush clumps increase in dominance, thereby increasing the frequency of competitive interactions and leading to a possible switch from facilitative to competitive interactions. Spatial point pattern analysis provides a useful tool to elucidate these underlying patterns and ecological processes. I used high resolution LiDAR data combined with spatial point pattern analysis to understand tree-tree interactions in a semi-arid savanna in the Eastern Cape Province of South Africa. I conducted a cluster analysis based on vegetation structural variables to distinguish different stages of woody plant encroachment from open savanna to closed canopy thicket. Using the canopy height model, I quantified the change in the grass height from open savanna to closed canopy thicket clumps as an indicator of a possible biome shift. Additionally, I used spatial point pattern analyses to investigate the effect of thicket clump formation on the composition of savanna and thicket species, the overall patterns of trees, and the associations of small thicket species with large Vachellia karroo trees, which serve as clump initiators. Finally, I examined the mortality of savanna trees across increasing stages of thicket expansion using second order spatial statistics, namely the Mark- and Pair-Correlation Functions. Results confirmed that three vegetation states, influenced by elevation, are present at the study site, representing open canopy savanna (early-stage thicket encroachment), encroached savanna with low thicket dominance (intermediate-stage thicket encroachment), and highly encroached with dominant thicket clumps (late-stage thicket encroachment). These stages showed increasing tree height, canopy cover and canopy height density, as well as decreased (but not completely absent grass layer) as thicket encroachment progresses. Spatial point pattern analysis showed, as predicted, that there was an overall aggregation of trees at small-scales within early thicket clump formation, from which I inferred that facilitative relationships may exist between trees. Contrary to my predictions, at later stages of thicket clump formation I found dominant independent patterns between savanna adults and juvenile thicket species, which may result from a combination of facilitative and competitive effects. Lastly, as expected, I found that the density of V. karroo mortality increased as thicket encroachment increased, with an overall random spatial pattern of dead V. karroo across encroachment stages. As predicted, tree mortality was randomly distributed in space in the open savanna state, and as thicket clump formation increases, tree competitive mortality became more evident, as well as decreased tree performance. Overall, the study highlights the interplay between facilitation and competition in semiarid savanna where thicket clumps are expanding. Intervention strategies are suggested to target areas of intermediate thicket clump formation, as these areas provide an opportunity to remove V. karroo before the nucleation process has enabled the establishment and increase of thicket species and to ensure the grass layer is kept productive. I conclude that the use of remote sensing and LiDAR technology holds a wide range of possibilities for monitoring and managing woody encroachment in savanna systems, however these methods need to be further refined for effective use within African savanna and thicket context, which displays high spatial aggregation making typical segmentation methods difficult. , Thesis (MSc) -- Faculty of Science, Botany, 2024
- Full Text:
- Authors: Putzier, Rachel Rayne
- Date: 2024-10-11
- Subjects: Arid regions South Africa Eastern Cape , Spatial analysis (Statistics) , Thicket , Lidar , Cluster analysis , Trees Mortality , Scrub encroachment
- Language: English
- Type: Academic theses , Master's theses , text
- Identifier: http://hdl.handle.net/10962/464484 , vital:76515
- Description: Woody thickening has negative economic and ecological impacts in savannas globally. While the increase of savanna trees as a form of bush encroachment has been well studied, less is known about the increase of thicket species in savannas, which is an important phenomenon resulting in the formation of closed-canopy clumps which may exclude the grass layer over time. The early stage of thicket expansion is often initiated by a nucleating savanna species which facilitates the establishment of woody thicket pioneer species, and as the thicket clump formation expands, bush clumps increase in dominance, thereby increasing the frequency of competitive interactions and leading to a possible switch from facilitative to competitive interactions. Spatial point pattern analysis provides a useful tool to elucidate these underlying patterns and ecological processes. I used high resolution LiDAR data combined with spatial point pattern analysis to understand tree-tree interactions in a semi-arid savanna in the Eastern Cape Province of South Africa. I conducted a cluster analysis based on vegetation structural variables to distinguish different stages of woody plant encroachment from open savanna to closed canopy thicket. Using the canopy height model, I quantified the change in the grass height from open savanna to closed canopy thicket clumps as an indicator of a possible biome shift. Additionally, I used spatial point pattern analyses to investigate the effect of thicket clump formation on the composition of savanna and thicket species, the overall patterns of trees, and the associations of small thicket species with large Vachellia karroo trees, which serve as clump initiators. Finally, I examined the mortality of savanna trees across increasing stages of thicket expansion using second order spatial statistics, namely the Mark- and Pair-Correlation Functions. Results confirmed that three vegetation states, influenced by elevation, are present at the study site, representing open canopy savanna (early-stage thicket encroachment), encroached savanna with low thicket dominance (intermediate-stage thicket encroachment), and highly encroached with dominant thicket clumps (late-stage thicket encroachment). These stages showed increasing tree height, canopy cover and canopy height density, as well as decreased (but not completely absent grass layer) as thicket encroachment progresses. Spatial point pattern analysis showed, as predicted, that there was an overall aggregation of trees at small-scales within early thicket clump formation, from which I inferred that facilitative relationships may exist between trees. Contrary to my predictions, at later stages of thicket clump formation I found dominant independent patterns between savanna adults and juvenile thicket species, which may result from a combination of facilitative and competitive effects. Lastly, as expected, I found that the density of V. karroo mortality increased as thicket encroachment increased, with an overall random spatial pattern of dead V. karroo across encroachment stages. As predicted, tree mortality was randomly distributed in space in the open savanna state, and as thicket clump formation increases, tree competitive mortality became more evident, as well as decreased tree performance. Overall, the study highlights the interplay between facilitation and competition in semiarid savanna where thicket clumps are expanding. Intervention strategies are suggested to target areas of intermediate thicket clump formation, as these areas provide an opportunity to remove V. karroo before the nucleation process has enabled the establishment and increase of thicket species and to ensure the grass layer is kept productive. I conclude that the use of remote sensing and LiDAR technology holds a wide range of possibilities for monitoring and managing woody encroachment in savanna systems, however these methods need to be further refined for effective use within African savanna and thicket context, which displays high spatial aggregation making typical segmentation methods difficult. , Thesis (MSc) -- Faculty of Science, Botany, 2024
- Full Text:
The process of thicket encroachment in semi-arid savanna: community patterns and biotic interactions
- Authors: Nell, Rhys
- Date: 2022-10-14
- Subjects: Scrub encroachment , Savanna ecology , Biotic interaction , Plant nutrients , Plant-water relationships
- Language: English
- Type: Academic theses , Master's theses , text
- Identifier: http://hdl.handle.net/10962/364966 , vital:65666
- Description: Bush encroachment in savannas is widespread in South Africa and is concerning, from both socio-economic and conservation viewpoints, as it affects ecosystem services, functioning and productivity. This phenomenon depends on multiple factors such as history, vegetation, management and environmental conditions, and their interplay. Encroachment into savannas has been relatively well-documented, however understanding of the different roles of tree-tree interactions between species that occur during this process is still limited. This includes the interactions causing spatial patterning, or how interactions and outcomes change over time in terms of encroachment succession from open savanna to closed-canopy thicket. The main objectives of this research are to document thicket establishment in a savanna ecosystem and consider the ecological roles of the key woody species and the abiotic properties of their micro-sites. Determining interactive effects of species co-occurrence is critical to understanding or predicting patterns and changes in biodiversity, nutrient distribution and available water resources. It is also imperative in determining correct and effective land management practices, particularly for reducing bush encroachment and its negative effect on rangelands. All data were collected on Endwell farm, located in the Smaldeel region of the Eastern Cape, South Africa. Endwell farm is a semi-arid savanna with a mean annual rainfall of 730 mm. First, I examine and describe the thicket encroachment process by exploring the associations between species and their size classes in the field. This was done by using plot-based belt transects and looking at changes in species size-class compositions from early to late successional stages. Association rules (market basket) analysis was used to identify the most common species size-class association patterns. The association between the savanna tree Vachellia karroo and the thicket pioneer Scutia myrtina was the most prevalent at all stages, with V. karroo being central to all associations in the first stage of encroachment; during later stages of encroachment, associations shift to incorporating other thicket pioneer species. The demography and clump formation of S. myrtina was strongly linked to associations with V. karroo to initiate bush clump formation. Results suggest that mature V. karroo facilitate the establishment and growth of S. myrtina. These two species were the focus of more detailed investigations to explore the nature and magnitude of their interspecific interactions. I then examined the effects of pairwise tree interactions between V. karroo and S. myrtina on soil and leaf nutrient content. I measureddifferences betweeninter-canopy and sub-canopy soil nutrient content, and the effect of associations on plant leaf nutrients, between pair-size combinations and individual controls. Results confirmed that pair-size tree interactions affected both soil nutrient and leaf nutrient content. All individuals increased soil K, N and organic C in the sub-canopy, while association with V. karrooincreased S. myrtinafoliar N, Pand K. In contrast, association with S. myrtinaloweredV. karroofoliar N, P and K. Small S. myrtina individuals werefound to benefit most from establishing and growing next to a large V. karroo individual, through mechanisms affecting soil and foliar nutrients. Scutia myrtina individuals establishing in association with smaller size classes of V. karroo showed no significant effects. I tested for positive and negative effects of pairwise tree interactions between Vachellia karroo and Scutia myrtina on available soil water and plant water potential (Ψ). This was done by looking at differences betweeninter-canopy and sub-canopy soil moisture and bulk density and associations on plant water stress (pre-dawn and mid-day leaf Ψ), between pair-size combinations and individual controls. I also selectively removed large V. karroo individuals from pairs to confirm the effects of competition andfacilitation. Similar to other studies, results confirmed positive and negative effects of pairwise tree interactions. Small S. myrtina individuals weremost facilitated by establishing and growing up next to a large V. karroo individual, through mechanisms affecting soil water content, bulk density and leaf Ψ. Scutia myrtina establishing in association with other size classes of V. karroo were much less facilitated, showing no significant effects. In contrast, large S. myrtina showed competitive interactions with V. karroo. , Thesis (MSc) -- Faculty of Science, Botany, 2022
- Full Text:
The process of thicket encroachment in semi-arid savanna: community patterns and biotic interactions
- Authors: Nell, Rhys
- Date: 2022-10-14
- Subjects: Scrub encroachment , Savanna ecology , Biotic interaction , Plant nutrients , Plant-water relationships
- Language: English
- Type: Academic theses , Master's theses , text
- Identifier: http://hdl.handle.net/10962/364966 , vital:65666
- Description: Bush encroachment in savannas is widespread in South Africa and is concerning, from both socio-economic and conservation viewpoints, as it affects ecosystem services, functioning and productivity. This phenomenon depends on multiple factors such as history, vegetation, management and environmental conditions, and their interplay. Encroachment into savannas has been relatively well-documented, however understanding of the different roles of tree-tree interactions between species that occur during this process is still limited. This includes the interactions causing spatial patterning, or how interactions and outcomes change over time in terms of encroachment succession from open savanna to closed-canopy thicket. The main objectives of this research are to document thicket establishment in a savanna ecosystem and consider the ecological roles of the key woody species and the abiotic properties of their micro-sites. Determining interactive effects of species co-occurrence is critical to understanding or predicting patterns and changes in biodiversity, nutrient distribution and available water resources. It is also imperative in determining correct and effective land management practices, particularly for reducing bush encroachment and its negative effect on rangelands. All data were collected on Endwell farm, located in the Smaldeel region of the Eastern Cape, South Africa. Endwell farm is a semi-arid savanna with a mean annual rainfall of 730 mm. First, I examine and describe the thicket encroachment process by exploring the associations between species and their size classes in the field. This was done by using plot-based belt transects and looking at changes in species size-class compositions from early to late successional stages. Association rules (market basket) analysis was used to identify the most common species size-class association patterns. The association between the savanna tree Vachellia karroo and the thicket pioneer Scutia myrtina was the most prevalent at all stages, with V. karroo being central to all associations in the first stage of encroachment; during later stages of encroachment, associations shift to incorporating other thicket pioneer species. The demography and clump formation of S. myrtina was strongly linked to associations with V. karroo to initiate bush clump formation. Results suggest that mature V. karroo facilitate the establishment and growth of S. myrtina. These two species were the focus of more detailed investigations to explore the nature and magnitude of their interspecific interactions. I then examined the effects of pairwise tree interactions between V. karroo and S. myrtina on soil and leaf nutrient content. I measureddifferences betweeninter-canopy and sub-canopy soil nutrient content, and the effect of associations on plant leaf nutrients, between pair-size combinations and individual controls. Results confirmed that pair-size tree interactions affected both soil nutrient and leaf nutrient content. All individuals increased soil K, N and organic C in the sub-canopy, while association with V. karrooincreased S. myrtinafoliar N, Pand K. In contrast, association with S. myrtinaloweredV. karroofoliar N, P and K. Small S. myrtina individuals werefound to benefit most from establishing and growing next to a large V. karroo individual, through mechanisms affecting soil and foliar nutrients. Scutia myrtina individuals establishing in association with smaller size classes of V. karroo showed no significant effects. I tested for positive and negative effects of pairwise tree interactions between Vachellia karroo and Scutia myrtina on available soil water and plant water potential (Ψ). This was done by looking at differences betweeninter-canopy and sub-canopy soil moisture and bulk density and associations on plant water stress (pre-dawn and mid-day leaf Ψ), between pair-size combinations and individual controls. I also selectively removed large V. karroo individuals from pairs to confirm the effects of competition andfacilitation. Similar to other studies, results confirmed positive and negative effects of pairwise tree interactions. Small S. myrtina individuals weremost facilitated by establishing and growing up next to a large V. karroo individual, through mechanisms affecting soil water content, bulk density and leaf Ψ. Scutia myrtina establishing in association with other size classes of V. karroo were much less facilitated, showing no significant effects. In contrast, large S. myrtina showed competitive interactions with V. karroo. , Thesis (MSc) -- Faculty of Science, Botany, 2022
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