Tree size and herbivory determine below-canopy grass quality and species composition in savannahs

Large single-standing trees are rapidly declining in savannahs, ecosystems supporting a high diversity of large herbivorous mammals. Savannah trees are important as they support both a unique flora and fauna. The herbaceous layer in particular responds to the structural and functional properties of a tree. As shrubland expands stem thickening occurs and large trees are replaced by smaller trees. Here we examine whether small trees are as effective in providing advantages for grasses growing beneath their crowns as large trees are. The role of herbivory in this positive tree-grass interaction is also investigated. We assessed soil and grass nutrient content, structural properties, and herbaceous species composition beneath trees of three size classes and under two grazing regimes in a South African savannah. We found that grass leaf content (N and P) beneath the crowns of particularly large (ca. 3. 5 m) and very large trees (ca. 9 m) was as much as 40% greater than the same grass species not growing under a tree canopy, whereas nutrient contents of grasses did not differ beneath small trees

Frequent burning promotes invasions of alien plants into a mesic African savanna

Fire is both inevitable and necessary for maintaining the structure and functioning of mesic savannas. Without disturbances such as fire and herbivory, tree cover can increase at the expense of grass cover and over time dominate mesic savannas. Consequently, repeated burning is widely used to suppress tree recruitment and control bush encroachment. However, the effect of regular burning on invasion by alien plant species is little understood. Here, vegetation data from a long-term fire experiment, which began in 1953 in a mesic Zimbabwean savanna, were used to test whether the frequency of burning promoted alien plant invasion. The fire treatments consisted of late season fires, lit at 1-, 2-, 3-, and 4-year intervals, and these regularly burnt plots were compared with unburnt plots. Results show that over half a century of frequent burning promoted the invasion by alien plants relative to areas where fire was excluded. More alien plant species became established in plots that had a higher frequency of burning. The proportion of alien species in the species assemblage was highest in the annually burnt plots followed by plots burnt biennially. Alien plant invasion was lowest in plots protected from fire but did not differ significantly between plots burnt triennially and quadrennially. Further, the abundance of five alien forbs increased significantly as the interval (in years) between fires became shorter. On average, the density of these alien forbs in annually burnt plots was at least ten times as high as the density of unburnt plots. Plant diversity was also altered by long-term burning. Total plant species richness was significantly lower in the unburnt plots compared to regularly burnt plots. These findings suggest that frequent burning of mesic savannas enhances invasion by alien plants, with short intervals between fires favouring alien forbs. Therefore, reducing the frequency of burning may be a key to minimising the risk of alien plant spread into mesic savannas, which is important because invasive plants pose a threat to native biodiversity and may alter savanna functioning

Structure and composition of Androstachys johnsonii woodland across various strata in Gonarezhou National Park, southeast Zimbabwe

A study on the structure and composition of Androstachys johnsonii Prain (Euphorbiaceae) woodland across three strata was conducted in Gonarezhou National Park (GNP), southeast Zimbabwe. Specifically, the objectives of the study were: (i) to determine the spatial structure and composition of A. johnsonii woodland in GNP and (ii) to determine factors that influence the structure and composition of A. johnsonii woodland in GNP. This study was based on a stratified random design with three major soil groups, and 30 plots were sampled in May 2010. The three soil strata were comprised of soils derived from (i) rhyolite, (ii) malvernia and (iii) granophyre bedrocks. A total of 1258 woody plants were assessed and 41 woody species were recorded. There were significant differences in mean tree heights, tree densities, basal area and species diversity in A. johnsonii woodland across the three soil strata. In contrast, there were no significant differences in the mean number of dead plants per ha in the three study strata in the GNP. Our study findings suggest that A. johnsonii woodland in GNP is being degraded. GNP management should develop a monitoring program through establishing monitoring plots in A. johnsonii woodland, and further studies need to be carried out, particularly on recruitment of A. johnsonii in the GNP

Influence of fire frequency on Colophospermum mopane and Combretum apiculatum woodland structure and composition in northern Gonarezhou National Park, Zimbabwe

We investigated the long-term effects of fire frequency on Colophospermum mopane and Combretum apiculatum woodland structure and composition in northern Gonarezhou National Park (GNP), Zimbabwe. Fire frequency was categorised as high (every 1–2 years), medium (every 3–4 years) and low (every 5–6 years). The following variables were measured or recorded: plant height, species name, canopy depth and diameter, basal circumference, number of stems per plant, plant status (dead or alive) and number of woody plants in a plot. There was a positive correlation (r = 0.55, P = 0.0007) between annual area burnt (total from January to December) and annual rainfall (average over two rain stations per rain year, July to June) between 1972 and 2005. A total of 64 woody species were recorded from C. mopane and C. apiculatum woodlands. Mean plant height increased from 4.5 to 8.2 meters in C. mopane woodland and from 4.5 to 5.1 meters in C. apiculatum woodland in areas subjected to high and low fire frequencies. In C. mopane woodland, low fire frequency was characterised by a significantly low density of woody plants (P 0.05). Our results suggest that C. mopane and C. apiculatum woodlands are in a state of structural transformation. Fire frequency effects, however, appear to be woodland specific. Fire management strategies in GNP should take into consideration annual rainfall and the different vegetation type

Spatio-temporal dynamics of woody vegetation structure in a human-modified South African savanna

A thesis submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Doctor of Philosophy. 25 October 2016 in Johannesburg, South Africa.Ecosystem services, nature’s benefit to people, contribute to human well-being. Extensive reliance on, and unsustainable use of, natural resources is typical of the rural poor in developing countries and can lead to ecosystem degradation, decreased ecosystem service provision, and increased vulnerabilities of rural populations. Most ecosystem services are intangible or difficult to quantify, but fuelwood provisioning can be measured directly and can serve as a proxy for the status of other ecosystem services (e.g. aesthetic and spiritual services, nutrient cycling, carbon sequestration). South African rural communities have a high reliance on fuelwood despite extensive access to electricity. Within this context, live wood harvesting occurring around rural settlements in increasing amounts has been deemed unsustainable. However, the ‘fuelwood crisis’ of the 1970s, and subsequent predictions of woodland collapse through fuelwood supply-demand models, has still not occurred despite substantial population growth in developing countries. Hypothesised reasons for modelled supply-demand mismatches are based on underestimation of fuelwood supply and woodland regeneration, as well as overestimation of fuelwood demand by discounting behavioural adaptability of users. It is likely that the spatial configuration of fuelwood use allows for the co-adaptability of both humans and ecosystems. A lack of understanding of the spatial configuration of these social-ecological dynamics limits our insights into current and future adaptive responses and thus, the degree of sustainability. This thesis aimed to assess woody biomass stocks and vertical structure changes, as a proxy for provisioning ecosystem services, in a spatially and temporally explicit manner, to describe the status and impact of wood extraction in semi-arid, savanna communal lands. Using repeat, airborne light detection and ranging (LiDAR) data from 2008 and 2012, we surveyed three-dimensional woodland structure in Bushbuckridge Municipality communal lands – the grazing and harvesting areas for densely populated rural settlements in former Apartheid ‘homelands’ in South Africa. Woody biomass in 2008 ranged from 9 Mg ha-1 on gabbro geology to 27 Mg ha-1 on granitic geology. Land-use pressure was associated with compensatory regrowth of savanna tree species through post-harvest coppice in the 1-3m height class. Woody biomass increased at all sites, contrary to previous fuelwood models of the area. Change detection in the vertical canopy structure revealed that biomass increases were almost solely attributable to the 1-3m and 3-5m height classes. These changes were exacerbated by wood extraction intensity in the communal lands – the communal land with the highest wood extraction pressure experienced the greatest biomass increases, likely a strong regrowth response to high harvesting levels. Within communal lands, areas closest to roads and settlements experienced substantial biomass increases as a result of shrub level gains. This relationship was mediated by the usage gradient – the greater the wood extraction pressure on the communal land, the larger and more spatially coalesced the ‘hotspots’ of shrub-level increases were in relation to ease of access to the communal land. However, biomass increases are not necessarily indicative of woodland recovery, as shrub-level increases were coupled with losses of trees >3m in height. To explore these tall tree dynamics further, we tracked >450 000 individual tree canopies over two years over contrasting landscapes – a private reserve containing elephants, two communal lands under different wood extraction pressures, and a nature reserve fenced off from both elephants and humans. Humans are considerable drivers of treefall (defined here as a ≥75% reduction in the maximum height of each tree canopy) in communal lands. Humanmediated biennial treefall rates were 2-3.5 fold higher than the background treefall rate of 1.5% treefall ha-1 (in the control site – the reserve containing neither elephants nor humans). Elephant-mediated treefall was five fold higher than the background rate. Rate and spatial patterns of treefall were mediated by geology and surface water provision in the elephantutilised site where relative treefall was higher on nutrient-rich geology, and intense treefall hotspots occurred around permanent water points. Human-mediated rates and spatial patterns of treefall were influenced by settlement and crop-land expansion, as well as ease of access to communal lands. Frequent fires facilitated the persistence of trees >3m in height, but was associated with height loss in trees <3m. The combined loss of large trees and gain in shrubs could result in a structurally simple landscape with reduced functional capacity. Shrub-level increases in the communal lands are likely an interactive combination of newly established woody encroachers and strong coppice regrowth in harvested species. The more intensely used the communal land, the greater the bush thickening and the stronger the relationship between biomass gains and structural changes in the lowest height classes. The exacerbation of bush thickening in natural resource-dependent communities has critical implications for ecosystem service provision. There is potential for coppice regrowth to provide fuelwood to communities using ‘tree thinning’ programmes, but there is a lack of data on the quantity and quality of the regrowth, as well as the sustainability of coppice, the impacts of different harvesting methods, and the potential feedbacks with changing climate and CO2 fertilisation. Woody resource spatial distribution in communal lands is centred around settlement-level wood extraction pressure, as well as natural resource accessibility in the woodlands. In highly utilised areas, woodland regenerative capacity has been underestimated. Additionally, natural resource extraction is still highly localised, even at the communal land scale, with major structural changes occurring around the periphery or close to existing infrastructure. However, it is these underrated coupled adaptive responses in social-ecological systems that explain the failure of fuelwood supply-demand models’ predictive abilities. Nevertheless, loss of large trees in the landscape and the persistence of ‘functionally juvenile’ coppice stands will have implications for seedling production and establishment in the landscape with repercussions for the future population structure and ecosystem service provision. I discuss the implications of increased natural resource reliance in an African development context and the positive feedback between rural poverty and environmental impoverishment. Potential constraints to the data are unpacked, together with opportunities for further research in this area.LG201

Topo-edaphic controls over woody plant biomass in South African savannas

The distribution of woody biomass in savannas reflects spatial patterns fundamental to ecosystem processes, such as water flow, competition, and herbivory, and is a key contributor to savanna ecosystem services, such as fuelwood supply. While total precipitation sets an upper bound on savanna woody biomass, the extent to which substrate and terrain constrain trees and shrubs below this maximum remains poorly understood, often occluded by local-scale disturbances such as fire and trampling. Here we investigate the role of hillslope topography and soil properties in controlling woody plant aboveground biomass (AGB) in Kruger National Park, South Africa. Large-area sampling with airborne Light Detection and Ranging (LiDAR) provided a means to average across local-scale disturbances, revealing an unexpectedly linear relationship between AGB and hillslope-position on basalts, where biomass levels were lowest on crests, and linearly increased toward streams (&lt;i&gt;R&lt;/i&gt;&lt;sup&gt;2&lt;/sup&gt; = 0.91). The observed pattern was different on granite substrates, where AGB exhibited a strongly non-linear relationship with hillslope position: AGB was high on crests, decreased midslope, and then increased near stream channels (&lt;i&gt;R&lt;/i&gt;&lt;sup&gt;2&lt;/sup&gt; = 0.87). Overall, we observed 5-to-8-fold lower AGB on clayey, basalt-derived soil than on granites, and we suggest this is due to herbivore-fire interactions rather than lower hydraulic conductivity or clay shrinkage/swelling, as previously hypothesized. By mapping AGB within and outside fire and herbivore exclosures, we found that basalt-derived soils support tenfold higher AGB in the absence of fire and herbivory, suggesting high clay content alone is not a proximal limitation on AGB. Understanding how fire and herbivory contribute to AGB heterogeneity is critical to predicting future savanna carbon storage under a changing climate

Above ground woody community attributes, biomass and carbon stocks along a rainfall gradient in the savannas of the central lowveld, South Africa

Enumeration of carbon stocks at benchmark sites is a necessary activity in assessing the potential carbon sequestration and possible generation of credits through restoration of intensively impacted sites. However, there is a lack of empirical studies throughout much of the savannas of sub-Saharan Africa, including South Africa. We report an estimation of species specific and site biomass and carbon stocks, and general vegetation structural attributes from three protected areas along a rainfall gradient in the central lowveld, South Africa. Estimates of biomass and carbon stocks were effected through destructive sampling to establish locally derived allometric equations. There was a gradient of increasing woody density, height of the canopy, number of species, density of regenerative stems and a greater proportion of stems in small size classes from the arid locality to the mesic locality, with the semi-arid locality being intermediate. The proportion of spinescent species decreased with increasing rainfall. The mesic locality was significantly more woody than either the arid or semi-arid sites, having double the biomass, four times the density and 40% higher basal area. Above ground carbon pools were also higher; carbon stocks were approximately 9 t/ha for the arid and semi-arid sites and 18 t/ha for the mesic site.

Effects of repeated burning on woody vegetation structure and composition in a semi-arid southern African savanna

The objective of this study was to investigate the effects of repeated dry season annual hot fires on woody plants in a semiarid southern African savanna in Zimbabwe. Parts of the National University of Science and Technology (NUST) research fields in Bulawayo, Zimbabwe have been burnt annually in the dry season between 1994 and 2003 in order to control bush encroachment. The present study was carried out in both the burnt and unburnt sites of the NUST research fields consisting of Acacia karroo-Colophospermum mopane vegetation. The study adopted a randomised block design and woody vegetation data were collected from a total of 10 plots. Variables measured and recorded included woody plant height, density, number of stems per plant, proportion of multistemmed plants, proportion of dead stems, basal area, fire damage and number of species per plot. The study results indicate that there were significant differences (P 0.05) in density, number of species per plot and number of stems per plant in woody plants between the burnt and unburnt sites. The study results suggest that repeated dry season annual hot fires leads to thinner and shortstemmed plants in semiarid savanna ecosystems. Repeated burning also increased the proportion of multistemmed plants and proportion of dead stems in the burnt site. Despite burning sections of the study area annually, bush encroachment control has not been effectively achieved. The study findings points to the need of adaptive management strategies in the use of fires in managing vegetation in semiarid savanna ecosystem

Water and available nitrogen as co-determinants of a mesic savanna in Kruger National Park, South Africa

Includes bibliographical references (leaves 66-75).Savanna is a summer rainfall, tropical and subtropical vegetation type with a continuous grass cover and discontinuous cover of trees. Rainfall plays an important role in savanna as it triggers many ecosystem processes such as nitrogen mineralization. This rainfall is however extremely variable with cycles of wet and dry years. The impacts of rainfall variability on N mineralization rates and tree-grass dynamics are not well known. This study seeks to provide a quantitative understanding on the influence of rainfall amount on N mineralization rates, and how this interaction (of water and available N) may affect structural diversity in savannas. Rainfall manipulation treatments were set up to simulate different rainfall amounts. These treatments included a wet treatment which received normal rainfall regime plus irrigation( 150 % of rain). The control treatment had no rainfall manipulation and received the normal rainfall regime of the year (l00 % rain). The dry treatment received half of the normal rainfall (50 %). Furthermore, the treatments had different plant cover combinations of tree-grass, grass only and tree only. Volumetric soil water content and nitrogen mineralization rates were determined in all treatments from August to June for two growing seasons. N mineralization rates were determined by incubating soil cores in situ for +1- 28 days. Grass biomass, tree saplings height and basal area were measured to determine the treatment effects on plant growth. Leaf area and nutrient contents of leaves were quantified to establish physiological response of saplings to different rainfall regimes was also determined. N mineralization rate was highest in wet treatments (0.55 - 0.29 flg NI g soil! day) and lowest in dry treatments ( 0.33 - 0.11 flg NI g soil! day). Grass biomass increased in wet treatments (1901.3 - 2079.2 Kglha) and decreased in dry treatments (722.3 - 880.6 Kg/ha), while the saplings' growth decreased in wet treatments (190 cm) and increased in dry treatments (265 cm). However, saplings experienced greatest growth when grass was removed, than when grass growth was suppressed by dry conditions. These results show that increase in water and available N in wet treatments led to high grass productivity and intense competition on tree sapling which showed little growth. In dry treatments, grasses were limited by water and low N availability as shown by the reduction in grass productivity. Reduced grass competition in dry treatments, and lack of competition in grass removal treatments opened an opportunity for saplings to grow faster. This shows that grass competition plays an important role as it can limit saplings from growing beyond sizes where they are vulnerable to fire and browsing. This study shows that it is not only the impact of rainfall that is driving tree-grass coexistence and structural diversity in savannas, but the combined effects of rainfall and availabl