Resprouting Response among Savanna Tree Species in Relation to Stem Size, Woody Removal Intensity and Herbicide Application.

Mechanical and chemical methods are widely used to control woody plant encroachment in many African countries. However, very little is known about the effectiveness of these control methods among woody species of different ages. We conducted a field experiment to determine the effects of different tree removal treatments (10%, 20%, 50%, 75% and 100%) and herbicide application (Picloram; 6 mL L-1) on the resprouting ability and vigour of 12 woody plant species. We examined 20 plots (30 m × 30 m) that were each subjected to tree removal, followed by herbicide application on half of the stems for each plot. All the tree species in this study resprouted after cutting. The applied concentration of herbicide significantly reduced the shoot production for Ehretia rigida, Vachellia robusta and Ziziphus mucronata, with a marginal effect for Dichrostachys cinerea. The diameter of stems was an important factor in determining resprouting ability, with shoot production decreasing with increasing stem diameter. However, stem diameter did not affect shoot length and diameter for all species. We found that woody plants are more likely to resprout and survive as juveniles than as adults after cutting and that herbicide only affected four of the twelve species at a concentration of 6 mL L-1. Thus, testing the amount of Picloram needed to kill certain woody species may be of importance for land users in southern African savannas.

The utilisation of domestic goats in rural and peri-urban areas of KwaZulu-Natal, South Africa.

Goats are a common ruminant in livestock husbandry systems in communal areas of southern Africa, but less so in peri-urban areas. While dynamics of goat farming in the former are relatively well-understood, little is known about it in peri-urban environments. We investigated the contribution of small-scale farming of goats to household livelihoods in rural and peri-urban areas in KwaZulu-Natal Province of South Africa. We used a semi-structured questionnaire survey to elicit views of 115 respondents on the contribution of goats to household income at two rural (Kokstad, Msinga) and two peri-urban (Howick, Pietermaritzburg) locations. Goats contributed to household income as a source of cash and meat in various sociocultural contexts such as at weddings, funerals and festive period gatherings (e.g. Easter and Christmas), including payment for household needs and expenses including food, school fees and medico-cultural consultations. These findings were more pronounced in rural areas, where more goats were kept than in peri-urban areas, which also had smaller herds per household. Goats were a source of cash in numerous ways including sales of their skins after slaughter and use of skins through value addition to make household craft items such as stools that could be sold for cash. None of the farmers milked their goats. Goat famers also kept cattle (52%), sheep (23%) and chickens (67%). Goat ownership appeared to be more profitable in rural areas and contributed to a lesser extent as a source of income in peri-urban areas where goats were largely kept for sale. There is potential for increased value addition of goat products to increase returns from small-scale goat farming in rural and peri-urban settings. Artefacts and cultural symbols derived from goat products are pervasive amongst Zulu people, and represent an additional avenue of research into 'hidden' valuation of goats.

Utilisation of Woody Plants by the Cape Porcupine in Mesic Savannas in South Africa.

Herbivory plays a fundamental role in determining the structure of savannas. The impacts of small and medium-sized mammalian herbivores on trees in savannas remain poorly understood because most research attention focuses on large herbivores such as elephants whose destructive effects on trees can be pervasive at landscape scales. On the other hand, feeding activities of generalist herbivores such as Cape porcupines on woody plants can lead to tree mortality. This study investigated the utilisation of woody plants by the Cape porcupine in three mesic savanna sites in South Africa. We determined the woody plant diet of the porcupine for the early and late dry seasons at Roodeplaat Farm in Gauteng Province, and at Goss Game Farm and Bisley Valley Nature Reserve in KwaZulu-Natal Province. Thirty and twenty randomly located quadrats (30 m × 30 m) were laid at Roodeplaat and Goss, respectively, while 10 smaller quadrats (10 m × 10 m) were laid at Bisley. We measured stem diameter and the length and width of bark scars made by porcupines on stems of woody plants. We collected ten dung samples from each study site in the wet and dry seasons for quantification of woody material in porcupine diet. Porcupine foraging behaviour impacted different tree species at each site: Vachellia robusta at Roodeplaat, Spirostachys africana at Goss and Vachellia nilotica at Bisley. Each of these trees was dominant at each site. More scarring and tree mortality were recorded at Bisley with almost 70% tree sapling mortality occurring on trees that porcupine fed on. The size of bark scars was greater at Goss (P < 0.01) than at Roodeplaat and Bisley, which were similar. The area of bark damage on S. africana trees differed significantly by stem diameter size class (P = 0.007) and was greater for small stems (size class < 7.1 cm) than the larger stems (size classes 7.1-14 cm and 14.1-21). For all the study sites, dung samples revealed that woody material contributed over 80% of the porcupine diet during the dry season, but was lower at 35% during the wet season for Roodeplaat, although it was consistently high for Bisley at 79%. Porcupine foraging activities substantially contributed to tree mortality at each site. We posit that porcupine induced mortality on dominant tree species at each site may contribute to structural heterogeneity in woody plant vegetation in mesic savannas.

Soil microbes and associated extracellular enzymes largely impact nutrient bioavailability in acidic and nutrient poor grassland ecosystem soils.

Understanding the role of soil microbes and their associated extracellular enzymes in long-term grassland experiments presents an opportunity for testing relevant ecological questions on grassland nutrient dynamics and functioning. Veld fertilizer trials initiated in 1951 in South Africa were used to assess soil functional microbial diversity and their metabolic activities in the nutrient-poor grassland soils. Phosphorus and liming trials used for this specific study comprised of superphosphate (336 kg ha-1) and dolomitic lime (2250 kg ha-1) (P + L), superphosphate (336 kg ha-1) (+ P) and control trials. These soils were analyzed for their nutrient concentrations, pH, total cations and exchange acidity, microflora and extracellular enzyme activities. The analysed soil characteristics showed significant differences except nitrogen (N) and organic carbon (C) concentrations showing no significant differences. P-solubilizing, N-cycling and N-fixing microbial diversity varied among the different soil treatments. ß-glucosaminidase enzyme activity was high in control soils compared to P-fertilized and limed soils. Alkaline phosphatase showed increased activity in P-fertilized soils, whereas acid phosphatase showed increased activity in control soils. Therefore, the application of superphosphate and liming influences the relative abundance of bacterial communities with nutrient cycling and fixing functions which account for nutrient bioavailability in acidic and nutrient stressed grassland ecosystem soils.

Alien invasive Leucaena leucocephala successfully acquires nutrients by investing in below-ground biomass compared to native Vachellia nilotica in nutrient-amended soils in South Africa.

Soils in grasslands and savannas of southern Africa are acidic and nutrient-poor. Legume plants, such as Vachellia nilotica and alien invasive Leucaena leucocephala, are a major component of the vegetation there. Vachellia nilotica can establish in drought-prone environments, and is invasive in high rainfall areas. Leucaena leucocephala is an emerging invasive in South Africa and is ranked among the world's 100 most invasive alien species. Alien plants can invade native habitats through their adaptability to low-resource soils, and thus can out-compete and displace native vegetation. We investigated the effects of phosphorus (P) deficiency and soil acidity on legume-microbe symbiosis, nitrogen (N) nutrition and carbon (C) growth costs of these two legumes in grassland soils. We used as inoculum and growth substrate soils collected from a long-term (>65 years) nutrient and lime-addition trial, the Veld Fertilizer Trial (VFT), located at Ukulinga Research Farm near Pietermaritzburg in South Africa. We used soils from three VFT treatments: soils fertilized with superphosphate (336 kg ha-1) applied once per year (+P), soils fertilized with superphosphate (336 kg ha-1) applied once per year with dolomitic lime (2250 kg ha-1) applied once every 5 years (P+L) and soils with no superphosphate and no dolomitic lime applications (Control). Seeds of V. nilotica and L. leucocephala were germinated and grown independently in these soils in green house conditions and harvested after 125 days for measurement of growth, legume-microbe symbiosis, N nutrition and C growth costs. Results showed that the two legumes had different growth adaptations. Vachellia nilotica grown in control soils and +P soils nodulated with various Burkholderia spp., while L. leucocephala did not nodulate in all soil treatments. Both legumes utilized for growth both atmospheric- and soil-derived N across all treatments thereby decreasing C growth costs. Vachellia nilotica grown in +P soils accumulated the most biomass and N nutrition. Leucaena leucocephala maximized specific N assimilation rates by investing in below-ground biomass accumulation in control soils. This shows that L. leucocephala possesses traits that are successful in acquiring nutrients by investing in below-ground biomass and relying on utilization of N from both the soil and the atmosphere.

Nitrogen Source Preference and Growth Carbon Costs of Leucaena leucocephala (Lam.) de Wit Saplings in South African Grassland Soils.

Leucaena leucocephala (Fabaceae) is native to Central America and has invaded many climatic regions of the tropics. In South Africa, the species is categorized as an emerging or incipient weed used as fodder, timber, firewood and in erosion control on degraded habitats. The species is common along the eastern subtropical regions of KwaZulu-Natal (KZN) Province, where it invades grasslands, savannas and edges of forests. Soils of these ecosystems are characterized as nutrient deficient and acidic. Using a pot trial, we determined the effects of the nutrient addition treatments on microbial symbiosis, N nutrition and biomass accumulation of L. leucocephala under greenhouse conditions. After 180 days of growth, plants were harvested, and their utilization of N derived from the atmosphere and from the soil was quantified through determination of δ15N values. L. leucocephala maintained growth and N nutrition by relying on both atmospheric- and soil-derived N across all soil treatments. The NDFA was significantly higher in high P (N1 + P, N2 + P and N3 + P) soils. L. leucocephala was able to nodulate with intermediate and fast-growing strains from the Mesorhizobium and Rhizobium genus in N2 + P grown plants. This shows that L. leucocephala possesses traits that are successful in acquiring nutrients, especially in nutrient limited conditions, by establishing plant symbiosis with multiple bacteria and relying on extracting N from the soil and from the atmosphere through the symbiosis.

Altering Nitrogen Sources Affects Growth Carbon Costs in Vachellia nilotica Growing in Nutrient-Deficient Grassland Soils.

Vachellianilotica (L.) Willd. Ex Del. is a multipurpose leguminous tree that is common in grassland and savanna ecosystems in southern and eastern Africa. These ecosystem soils are reported to be acidic and nutrient-limited, specifically with regards to nitrogen (N) and phosphorus (P). The presence of this plant in these terrestrial ecosystems improves soil fertility benefiting the surrounding vegetation due to its ability to fix atmospheric N. This study seeks to understand the N-fixing bacteria symbiosis and physiological adaptations of V. nilotica in these acidic and nutrient-deficient KwaZulu-Natal soils. The soils used for this study were collected from the Ukulinga Grassland Nutrient Experiment located at the Ukulinga research farm of the University of KwaZulu-Natal, Pietermaritzburg, South Africa. Due to long-term soil nutrient addition treatments, these soils offered a diverse nutrient variation for better understanding the effects of acidity and nutrient variation on microbial symbiosis, plant nutrition, and biomass accumulation of V. nilotica. V. nilotica was able to maintain growth by relying on both atmospheric and soil-derived N across all treatments decreasing carbon (C) growth costs. There was an increased reliance on atmospheric-derived N of un-nodulated high N-treated plants. The plants grown in high N + P soils were able to nodulate with various species from the Mesorhizobium genus, which resulted in increased biomass compared to other plants. The results of this study show that V. nilotica can alter N sources to reduce C growth costs. In addition, both nodulating and free-living soil N2 fixing bacteria such as Caulobacter rhizosphaerae, Sphingomonas sp. and Burkholderia contaminans identified in the experimental soils may play an important role under P-deficient conditions.

No safety net in the face of climate change: The case of pastoralists in Kunene Region, Namibia.

Over the past decade, pastoralists in Kunene Region, Namibia, have endured recurrent drought and flood events that have culminated in the loss of their primary form of livelihood-pastoralism. Most pastoralists are finding it difficult to sustain their livelihoods, and their communities have fallen into extreme poverty. Ecosystem-based Adaptation (EbA) approaches are increasingly acknowledged as having the potential to enhance the adaptive capacity of vulnerable communities. The first step is to develop an understanding of how affected communities live, their perceptions of and how they respond to climate change and the biophysical impacts of climate change in their communities. This study aims to collect this information in order to explore the use of EbA to help pastoralists adapt to climate change. We examined an isolated pastoral Himba community, to understand their perceptions, experiences and understanding of climate change and its related impacts on their livelihoods. A nested mixed-methods approach using structured interviews was employed to address the study objectives. Interview results revealed that pastoralists lack scientific knowledge of climate change, and they have no access to climate change information. Though pastoralists have coping and adaptation approaches at the community level (such as making gardens, fishing, etc.), these have become ineffective as climatic uncertainty and change persist. Furthermore, pastoralists no longer get benefits from the environment, such as food and fodder. Despite this, there are currently no biodiversity interventions at the community level to address the impacts of climate change. Pastoralists have indicated their adaptation needs, particularly the provision of water supply to grow food. This is an open avenue to explore EbA approaches, specifically ecological restoration, while still addressing the need of the pastoralists. There is an urgent need to develop new practical adaptation strategies, including restoration options that will strengthen their adaptive capacity.

Differential patterns of nitrogen nutrition and growth cost of the indigenous Vachellia sieberiana and the introduced Chromolaena odorata in the savannah environment.

Vachellia sieberiana fixes atmospheric nitrogen (N) and distributes it back into ecosystems. We hypothesize that biological nitrogen fixation in this plant species is limited by competition from the invasive shrub, Chromolaena odorata. Competition would therefore result in the legume plant switching its limited nitrogen (N) sources in phosphorus-poor soils in savannah ecosystems when resources have to be shared. This study investigated the different patterns of N use and growth costs by a native and an introduced leguminous shrubby species. We propose that the two species sharing the same environment might result in competition. The competitive effect would induce in the indigenous legume to better utilize atmospheric-derived N modifying plant growth kinetics and plant mineral concentrations. Seedlings of V. sieberiana were cultivated in natural soil inoculum with low levels of phosphorus (mg L-1 ± SE) of 3.67 ± 0.88. The experiments were divided into two treatments where (i) seedlings of V. sieberiana were subjected to competition by cultivating them together with seedlings of C. odorata, and (ii) seedlings of V. sieberiana were cultivated independently. Although V. sieberiana was subjected to competition, the N2-fixing bacteria that occupied the nodules was Mesorhizobium species, similar to plants not subjected to competition. Total plant biomass was similar between treatments although V. sieberiana plants subjected to competition accumulated more below-ground biomass and showed higher carbon construction costs than plants growing individually. Total plant phosphorus and nitrogen decreased in seedlings of V. sieberiana under competition, whereas no differences were observed in percent N derived from the atmosphere (%NDFA) between treatments. The specific nitrogen utilization rate (SNUR) was higher in V. sieberiana plants subjected to competition while specific nitrogen absorption rate (SNAR) showed the opposite response. Vachellia sieberiana is highly adapted to nutrient-poor savannah ecosystems and can withstand competition from invasive shrubs by utilizing both atmospheric and soil nitrogen sources.

An African grassland responds similarly to long-term fertilization to the Park Grass experiment.

We compared the results of a long-term (65 years) experiment in a South African grassland with the world's longest-running ecological experiment, the Park Grass study at Rothamsted, U.K. The climate is warm and humid in South Africa and cool and temperate in England. The African grassland has been fertilized with two forms of nitrogen applied at four levels, phosphorus and lime in a crossed design in 96 plots. In 1951, about 84% of plant cover consisted of Themeda triandra, Tristachya leucothrix and Setaria nigrirostris. Currently, the dominant species are Panicum maximum, Setaria sphacelata and Eragrostis curvula, making up 71% of total biomass. As in the Park Grass experiment, we found a significant (additive) interaction effect on ANPP of nitrogen and phosphorus, and a (marginally significant) negative correlation between ANPP and species richness. Unlike the Park Grass experiment, there was no correlation between ANPP and species richness when pH was included as a covariate. There was also a significant negative effect of nitrogen amount and nitrogen form and a positive effect of lime on species richness and species diversity. Soil pH had an important effect on species richness. Liming was insufficient to balance the negative effects on species richness of nitrogen fertilization. There was a significant effect of pH on biomass of three abundant species. There were also significant effects of light on the biomass of four species, with only Panicum maximum having a negative response to light. In all of the abundant species, adding total species richness and ANPP to the model increased the amount of variance explained. The biomass of Eragrostis curvula and P. maximum were negatively correlated with species richness while three other abundant species increased with species richness, suggesting that competition and facilitation were active. Consistent with the results from the Park Grass and other long-term fertilization experiments of grasslands, we found a positive effect of soil pH and a negative effect of nitrogen amount on species richness, a more acutely negative effect on species richness of acidic ammonium sulphate fertilizer than limestone ammonium nitrate, a negative relationship between species richness and biomass, and a positive effect on species richness of lime interacting with nitrogen.

Predator satiation and recruitment in a mast fruiting monocarpic forest herb.

BACKGROUND AND AIMS: Cross-pollination and satiation of seed predators are often invoked to explain synchronous mast reproduction in long-lived plants. However, explanations for the synchronous death of parent plants are elusive. The roles of synchronous seeding and post-reproductive mortality were investigated in a perennial monocarpic herb (Isoglossa woodii) in coastal dune forest in South Africa. METHODS: Pre-dispersal seed predation and seed production were assessed by measuring fruit and seed set of inflorescences sprayed with insecticide or water and with no spray treatments. Seed predation was measured at different densities of I. woodii plants by monitoring removal rates of seed from the forest floor. The influence of adult plants on establishment of I. woodii seedlings was assessed by monitoring growth and survivorship of seedlings in caged and uncaged 1 × 1 m plots in understorey gaps and thickets. KEY RESULTS: Fruit and seed set were similar between spray treatments. An I. woodii stem produced 767·8 ± 160·8 seeds (mean ± s.e.) on dune crests and 1359·0 ± 234·4 seeds on the foredune. Seed rain was greater on the foredune than in other topographic locations. Seed predation rates were 32 and 54 % on dune crests and in dune slacks, respectively, and decreased with seed abundance, number of inflorescences per stem and plant height. Seedling recruitment was greater beneath synchronously dying adult plants than in natural understorey gaps (no I. woodii). However, seedling growth rate beneath I. woodii mid-way through its life-cycle was less than in gaps, although survivorship was similar. CONCLUSIONS: The selective advantage of masting in I. woodii derives from satiation of both pre- and post-dispersal seed predators. In addition, post-seeding mortality of adult plants facilitates seedling establishment. Satiation of seed predators and the benefits of seedling establishment are strong drivers of the evolution of synchronous monocarpy in I. woodii.

Ecological filtering by a dominant herb selects for shade tolerance in the tree seedling community of coastal dune forest.

The regeneration niche is commonly partitioned along a gradient from shade-tolerant to shade-intolerant species to explain plant community assembly in forests. We examined the shade tolerance of tree seedlings in a subtropical coastal forest to determine whether the ecological filtering effect of a dominant, synchronously monocarpic herb (Isoglossa woodii) selects for species at either end of the light response continuum during the herb's vegetative and reproductive phases. Photosynthetic characteristics of seedlings of 20 common tree species and the herb were measured. Seedlings were grown in the greenhouse at 12-14% irradiance, and their light compensation points measured using an open-flow gas exchange system. The light compensation points for the tree species were low, falling within a narrow range from 2.1 ± 0.8 µmol m(-2) s(-1) in Celtis africana to 6.4 ± 0.7 µmol m(-2) s(-1) in Allophylus natalensis, indicating general shade tolerance, consistent with a high and narrow range of apparent quantum yield among species (0.078 ± 0.002 mol CO(2) mol(-1) photon). Rates of dark respiration were significantly lower in a generalist pioneer species (Acacia karroo) than in a forest pioneer (C. africana), or in late successional phase forest species. We argue that the general shade tolerance, and phenotypic clustering of shade tolerance, in many tree species from several families in this system, is a result of ecological filtering by the prevailing low light levels beneath the I. woodii understorey, which excludes most light-demanding species from the seedling community.