Doctoral Degrees (Grassland and Rangeland Science)

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Factors affecting savanna tree sapling recruitment.
(2013) Vadigi, Snehalatha.; Ward, David Mercer.
Savannas are globally important ecosystems characterized by the coexistence of trees and grasses. Woody plants, which are slow-growing dominant life forms, influence the physiognomic structure and function of savanna ecosystems. Their density and distribution provides sustenance to a vast and unique savanna biodiversity, by forming a major source of food material to large mammalian herbivores, sheltering them and through their facilitation of diverse plant species. Savanna tree existence is strongly affected by factors that determine their sapling recruitment. We defined „sapling‟ as a young tree, in the first season of its growth, which does not depend on cotyledonary reserves (=seedling stage) and relies on external resources to grow further. Sapling recruitment may strictly be defined as the progression of a young plant from seedling to sapling stage. However, we believe that savanna tree saplings, present within the grass layer in the initial years of their growth, are equally vulnerable to environmental stresses. This study examines the factors affecting tree sapling establishment in a humid savanna (1250 mm mean annual precipitation). Additionally, the effects of fire were tested in a greenhouse experiment. Dominant species from humid savannas (> 1000 mm MAP), Acacia karroo, Acacia sieberiana, Schotia brachypetala and Strychnos spinosa, and mesic savannas (approx. 750 mm MAP), Acacia nigrescens, Acacia tortilis, Colophospermum mopane and Combretum apiculatum, were studied. In this thesis I examined the effects of resource availability (water, nutrients and light), disturbances (fire and herbivory) and competition (grass) on the sapling ecology of these species. Sapling recruitment and growth were assessed in terms of survival and aboveground growth responses, i.e. total biomass, stem growth rates (used as proxy measures for assessing persistence) and leaf biomass proportion (important for producing root reserves necessary to resprout). I studied the effects of fire and a nutrient gradient on survival and growth of four Acacia species in the presence of grass competition, in a controlled greenhouse experiment. Generally, Acacias invest in defenses after herbivory. I also determined their physical and chemical defense investments in this experiment. Sapling survival was not influenced by nutrients but highly varied among the species due to fire, indicating that fires may have a differential effect on species composition at a landscape scale. Intermediate levels of nutrients were found to be beneficial for sapling growth than high and low levels. This may be due to an increase in grass competition at higher levels of nutrients. Fires did not have a positive influence on sapling defence investment. To evaluate the relative importance of resource availability on sapling tree recruitment and its interactions with grass competition, I tested the effects of water (frequent irrigation vs. rainfall), shade (presence vs. absence), nutrients (addition vs. no addition) and grass competition (presence vs. absence) on sapling survival and growth under controlled field conditions in a humid South African savanna. Treatments did not have an effect on sapling survival, indicating that mortality is not defined by resource availability and grass competition in humid savannas. Shade had the greatest negative effect on sapling growth, suppressing the beneficial effects of nutrients and absence of grass competition. Nutrient limitation and grass competition had a relatively small influence on savanna sapling growth. Frequency of water availability had no effect on sapling growth, perhaps owing to high rainfall experienced over the experimental period. Therefore, canopy shade can be considered to be an important driver of tree dynamics in humid savannas with some degree of influence by nutrient availability and grass competition. The effects of clipping (i.e. simulated herbivory of grass and tree saplings) as influenced by nutrient availability and grass competition were examined on sapling survival and growth of all study species in a humid savanna. None of the treatments had an effect on sapling survival. This signifies that herbivory alone cannot significantly decrease plant density in humid savannas. However, tree saplings grew taller with a reduction in diameter and overall biomass, implying that saplings may become more susceptible to fires after herbivory. Nutrient addition and grass competition in general had a positive and negative effect, respectively, on sapling growth. This response was prominent in the stem length growth rates of defoliated saplings of one humid and two mesic species. These results imply that clipping (or herbivory) is the major factor reducing sapling vigour to establish, but is affected by both grass competition and nutrient availability. This study shows that fire has a differential effect on sapling survival of different species, particularly between humid savanna species. Light interception among all other resources limits the recruitment of saplings into adult size classes. Clipping, nutrient availability and grass competition had a relatively small direct effect, but may interact with other factors to alter sapling establishment dynamics. Wet-season droughts in humid savannas are not a hindrance to tree establishment because sapling survival was not dependent on frequency of rainfall. Thus, in humid savannas, fires can have a major impact on tree species density and composition while canopy shade has a very high potential to alter tree distribution.
Browse : quantity and nutritive value of evergreen and deciduous tree species in semi-arid Southern African savannas.
(2012) Penderis, Caryn Anne.; Kirkman, Kevin Peter.
Browse selection, intake, utilisation, palatability, quality and production are tightly linked and need to be considered together in trying to improve our understanding of browsing dynamics and the interactions between browsers and vegetation. Such an understanding is necessary in order to re-evaluate determinations of browser carrying capacities and evaluating actual and potential impacts of browsing animals on vegetation composition and diversity. Browser carrying capacity is determined by both the quantity and the nutritive value of forage. The measurement of browse quantity and nutritive value and the matching of browse supply to browser demand are central to sustainable utilisation and the monitoring of vegetation health. South African savannas are poorly studied with respect to tree canopy growth and browse production making it difficult to quantify the available browse biomass on which browsing capacity estimations are based, and consequently difficult to estimate levels of browsing that are sustainable. This study addressed these issues by investigating browse dynamics, broadly aiming to (1) explore factors affecting browse production, biomass and nutritive value; (2) develop models to assess and monitor these parameters across seasons and properties; (3) use the resultant models in improving our understanding of how to determine browser carrying capacities. More specifically, our study sought to examine the effects of plant physiognomy, forage nutritive value, canopy stratum, defoliation, temperature, rainfall and soil nutrient status on the browse production of evergreen (Carissa bispinosa, Euclea divinorum, Gymnosporia senegalensis), semi-deciduous (Spirostachys africana, Ziziphus mucronata) and deciduous (Acacia nilotica, Dichrostachys cinerea) savanna tree species from June 2003 – June 2005 in three sites along the northern Zululand coastline of KwaZulu-Natal. Available browse biomass, during the dry season, of four key savanna tree species (A. nilotica, E. divinorum, G. senegalensis, and S. africana) was estimated through the development of allometric regression equations. Non-linear regression was used to investigate the relation between the leaf dry mass (LDM) and canopy volume (CVol) of each of the four tree species. Exponential regression (y = a + brlnx) of the natural logarithm of CVol data provided the most accurate and precise description of the tree CVol – LDM relation. A study was undertaken to determine which factors may influence browse production in a southern African savanna. Regression tree models for the browse production identified that the dominant factors influencing browse production were CVol (m3), season, species and height to the lowest leaves of the tree canopy (HL) (m). The length of the growing season had a marked effect on the production potential of savanna tree species, suggesting that improved conditions for growth, i.e. greater rainfall, soil moisture content and improved soil nutrient availability result in a longer period of rapid sustained growth. Species was identified as an important contributing factor to differences in browse production rates, suggesting the need for the development of species or species group models. Mean annual browse production of evergreen trees was greater than that of deciduous and semi-deciduous trees. Mean quarterly (three monthly) browse production was highest, for all trees, during the wet season, with the greatest difference between wet and dry season production being observed in deciduous forms. Evergreen forms showed continuous growth over the whole study, with enhanced growth over the wet season. Deciduous forms, on the other hand, concentrated growth in spurts, when environmental conditions became favourable, with most production occurring during a short growing season. Browse nutritive value was found to be greatest during the wet season, when growth and photosynthesis are at their greatest. Further, browse nutritive value was greatest in deciduous species. Evergreen trees were found to have greater acid detergent fibre (ADF) concentrations than both the deciduous and semi-deciduous trees. By contrast, crude protein (CP) concentrations were greater in semi-deciduous and deciduous species than in evergreen species. The daily CP requirements for maintenance for an adult impala (45 kg) were met by all species over all three study areas and all seasons. Daily CP requirements for growth and lactation, however, were only ever met by deciduous and semi-deciduous species, though this result was not consistent over study areas and seasons. Predictive models for the production of browse on deciduous, semi-deciduous and evergreen trees in northern Zululand were developed using multivariate adaptive regression spline functions. The best predictors of growing season browse production in all three tree guilds (defined here as a group of trees having a characteristic mode of living) were primarily measurable tree dimensions, while the prevailing environmental conditions had little impact. Differences in the production, nutritive value and available browse biomass between the different tree forms and seasons have a profound effect on the determination of browser carrying capacities and need to be incorporated into any game or conservation management plan.
The roles of competition, disturbance and nutrients on species composition, light interception and biomass production in a South African semi-arid savanna.
(2012) Mopipi, Keletso.; Kirkman, Kevin Peter.; Trollope, Winston Smuts Watts.; Morris, Craig Duncan.
Plants are the major source of food or energy required to sustain life on the planet, but humans are grappling with the deteriorating conditions of natural ecosystems such as compositional change, desertification, invasive plants and soil erosion. In the face of global climate change and growing demands for agricultural productivity, future pressures on grassland ecosystems will intensify, therefore sustainable utilization of all plant resources is of vital importance to enhance food security within the limits of good conservation. The semi-arid grasslands of southern Africa represent major grassland resources for grazing. Herbage production in these areas is determined not only by water and nutrient availability, but also by controlled and uncontrolled fires. Since fire is regarded as a natural factor in savannas, it is essential to develop a deeper understanding of the role of fire in community structure and function for the development of appropriate burning regimes. A study was conducted in the Eastern Cape of South Africa where the rural communities are faced with the challenges of rangeland degradation in the form of encroachment by unacceptable bush, karroid, macchia and less desirable grass species, as well as soil erosion. The main objective of this thesis was to investigate the roles of competition and disturbance regimes (fire and simulated non-selective grazing) on species composition, habitat productivity and the performances of selected species from this semi-arid savanna. Long-term effects of burning frequency on herbaceous species composition, Leaf Area Index (LAI), Photosynthetically Active Radiation (PAR) within the herbaceous canopy, biomass production and soil chemical properties were investigated. These studies were conducted on a fire trial set up in 1980 at the University of Fort Hare research farm in the Eastern Cape, South Africa. The treatments comprise an annual, biennial, triennial, quadrennial, sexennial and no burn control, all replicated twice in a Complete Randomized Design. The data from the trial collected between 1980 and 2008 were used to determine compositional variation for herbaceous species using the Non-metric Multidimensional Scaling and Bray-Curtis Dissimilarity tests. The PAR ceptometer was used to determine LAI and intercepted PAR, while random samples were harvested from 1m² quadrats from each plot. Soil samples were taken at four depths (0-2 cm; 2-4 cm; 4-6 cm and 6-8 cm) from each plot and analyzed for pH, Ca, K, P, total C and total N. The Resin-Bag technique was used to determine nitrogen mineralization. Burning frequency caused significant variation in herbaceous species composition over time. The species were distributed along gradients of increasing burning frequency, and these responses were in three categories: Those that increased with burning frequency such as Themeda triandra; those that decreased with burning frequency such as Melica decumbens, and those that showed little response such as Panicum maximum. The three-year burn resulted in the highest compositional variation, light interception, Leaf Area Index, aboveground biomass production, while the annual, biennial and no burn treatments resulted in the lowest. The fact that infrequent burning resulted in higher species variation, improved habitat productivity due to increased leaf area for light interception shows that appropriate use of fire can maintain a more diverse and productive savanna system. Burning frequency had significant effects on the soil properties, while soil depth did not show any significance. Frequent burning increased soil pH, K, Ca, and Na, but reduced C, N, P and N mineralization. There was a negative correlation between burning frequency and N mineralization, but no correlation existed between N mineralization and total N, total C or the C:N ratio. These results imply that frequent burning can cause nutrient losses and a greater nutrient limitation to plants in the long-term, especially soil C and N loss from combustion of organic material in the soil top layer. The ability of shade-tolerant plants to persist under shade and regular defoliation such as in burnt and grazed systems may be of greater importance for long-term productivity and sustainability of forage crops. It is therefore imperative to explore the mechanisms by which some species were favoured by frequent burning which created low shade conditions, while others were favoured by high shade conditions where burning is infrequent or absent. A pot experiment was conducted to investigate the shade tolerances of seven grass species that were abundant in the long-term fire trial. The test species were Cymbopogon plurinodis, Digitaria eriantha, Eragrostis curvula, Melica decumbens, Panicum maximum, Sporobolus fimbriatus and Themeda triandra. Individual grass tillers of each species were collected from the natural vegetation, propagated in separate seedling trays and transplanted into individual pots, and were grown under five shading treatments: full sun (0 % shading), 55 %; 70 %; 85 % and 93 % shading respectively. Shading significantly reduced the dry matter production of all the species. Biomass production of all the species decreased linearly to varying degrees with an increase in shade intensity. Digitaria eriantha and Eragrostis curvula were most adversely affected by shading, hence are classified as shade intolerant, while Melica decumbens was the least affected by shading, and is hence classified as shade tolerant. Cymbopogon plurinodis, Panicum maximum, Sporobolus fimbriatus and Themeda triandra are classified as moderately shade-tolerant. From the results it was apparent that some species could perform optimally in partial shade than in full sunlight, and these results lead to a conclusion that for satisfactory natural regeneration and seedling growth of this savanna vegetation would require a gap large enough to provide at least 30 % of ambient light. Investigating patterns in competitive effects and responses of species in these communities may not only explain the abundance of each species, but may also provide insight into the nature of forces that affect the structure and function of that community. Since fire, herbivory and soil nutrients are natural drivers of savanna community structure and function, their influence on competitive interactions of selected species were investigated. Two experiments were conducted to investigate the competitive effects and responses of eight selected common species in the area. The test species (phytometers) included one woody shrub, Acacia karroo and seven grass species namely: Cymbopogon plurinodis, Digitaria eriantha, Eragrostis curvula, Melica decumbens, Panicum maximum, Sporobolus fimbriatus and Themeda triandra. In an outdoor plot experiment the responses of the phytometers to competition from neighbours (0; 2 and eight neighbours respectively), fertility (fertilized, unfertilized) and clipping (clipping, no clipping) were investigated. The second comprised a pot experiment where the competitive effects of the species were investigated. Each species was grown under 3 levels of fertility (0 %; 50 % and 100 % Hoagland‘s solution) and clipping (clipping, no clipping) in pots filled with fine river sand and 4 neighbours. Competition intensity, soil fertility and clipping had significant effects on the biomass production of the phytometer species. Acacia karroo and Melica decumbens, exhibited the weakest competitive effects and responses, and incurred the highest mortalities after clipping and with 8 neighbours. Digitaria eriantha and Panicum maximum exhibited the strongest competitive effects and responses, especially in high fertility, and experienced the lowest mortalities. T.triandra exhibited stronger competitive effect after clipping in low fertility, while A. karroo and C. plurinodis exhibited stronger competitive effects in moderate (50 %) fertility. Cymbopogon plurinodis, Eragrostis curvula and Sporobolus fimbriatus ranked between these two extreme groups in terms of competitive effects and responses. Relative Competitive Interaction increased with soil fertility and number of neighbours in the absence of clipping. These results indicate that in general, taller or broad-leaved grass species outgrow the shorter ones, and this gives them a competitive advantage over light and soil resources. One of the range management objectives in the False Thornveld of the Eastern Cape is to promote the abundance of Themeda triandra, which is of high forage value and an indicator of rangeland that is in good condition. The general situation under livestock farming conditions in this area is that if the grass sward is optimally grazed and rested then there is a great potential for Themeda triandra to dominate.The results of the competition experiments indicated that the species exhibits strong competitive interaction, and also exhibited stronger competitive effect after clipping in low fertility. These results imply that it has a low response and a high effect, an attribute that would enhance its performance if it is moderately grazed or the area is burnt. The species is also moderately shade tolerant, and this may explain why it thrives in burnt and moderately grazed areas. These studies have demonstrated the important role that competition and disturbance in the form of fire and herbivory play in the maintenance of this savanna grassland. Through natural selection species are able to occupy different niches in the same area and coexist in a heterogeneous environment and minimize their chances of extinction.
Dry woodland and savanna vegetation dynamics in the Eastern Okavango Delta, Botswana.
(2012) Tedder, Michelle Jennifer.; Kirkman, Kevin Peter.; Bonyongo, Mpaphi Casper.; Morris, Craig Duncan.; Trollope, Winston Smuts Watts.
The Okavango Delta is an extremely dynamic system with variable vegetation comprised of permanent swamps, seasonal swamps, dry islands, floodplains and dry grassland, savanna and woodland. The system is largely driven by the interaction between fire and the annual flood, which filters down from the Okavango River catchments in Angola. While extensive research has been conducted on the flood-driven vegetation little is known about the dry woodland and savanna regions bordering these flood-driven habitats. A taxonomic classification of woody species composition resulted in eleven vegetation types. These data were then reanalyzed in terms of woody species morphology allowing these eleven vegetation types to be grouped into four functional response groups in order to provide a platform for improving the understanding of how dry woodland and savannas interact with the environment. These four groups were the savanna group mixed thornveld and the three woodland groups; mixed broadleaf woodland, shrub mopane woodland and tall mopane woodland. Burning in mixed thornveld and mixed broadleaf woodland was found to decrease woody species density and grass fuel loads and could be used for grazing management to remove unpalatable growth and improve grass species composition, while burning in shrub mopane woodland and mixed mopane woodland merely decreased the woody understory and is not recommended. Utilization dominated by grazing livestock resulted in overutilization of the grass sward leading to bush encroachment in both mixed thornveld and shrub mopane woodland, while utilization by goats alone resulted in underutilization of the grass sward and a dominance of herbaceous annuals. Livestock utilization had no effect on the occurrence of Pecheul-loeschea leubnitziae, a shrubby pioneer previously thought to be an indicator of overgrazing, however extensive P. leubnitziae cover was associated with a sward dominated by shade-tolerant grasses with low forage quality. Shrub mopane woodland and tall mopane woodland appear to be more stable vegetation states than mixed broadleaf woodland and mixed thornveld being less vulnerable to colonization by pioneer species and alteration as a result of utilization or environmental factors. For this reason management and monitoring of mixed thornveld and mixed broadleaf woodland is essential to prevent vegetation degradation and to ensure optimal forage availability for both livestock and wildlife.
Towards a predictive understanding of savanna vegetation dynamics in the eastern Lowveld of South Africa : with implications for effective management.
(2005) Peel, Michael John Stephen.; Zacharias, Peter John Kenneth.; Biggs, Chara.
The purpose of this study was to develop and test a predictive understanding of the vegetation dynamics of the Lowveld of South Africa (30°35'E to 30°40'E and 24°00'S to 25°00'S). The study covered about 5000 km2 in Adjacent Private Protected Areas (APPA) adjoining the Kruger National Park (KNP). Data gathering (800 sites; 23 properties) commenced in 1989 and those recorded up to 2004 are reported here. The value, both ecological and economic, of the wildlife and tourism industry dependent on this savanna region is discussed in both historical and current perspectives. A range of land-use objectives and anthropogenic interventions were exposed. The properties ranged in size from 30 to 800 km2 and formed an effective and extensive manipulative experiment for investigating interaction of bush density, animal stocking, use of fire and landscape-scale processes. The first descriptive classification (at 1:250 000) of the area was developed using Inverse Distance Weighted interpolations. This confirms similar landscape/vegetation patterns in the KNP and Mocambique. The current mode of determining stocking density or carrying capacity was interrogated and indices suitable for complex multi-species systems developed. This was done in the context of equilibrial/disequilibrial paradigms. Application of the original indices resulted in drought-related declines in animal biomass of 4000 kg km2 over 20 years due to overestimation of carrying capacity. The model proposed here uses rainfall, animal type, biomass and vegetation parameters to determine stocking density for both coarse (regional) and ranch-specific scales. Principal driving determinants (rainfall, geology, soil type, tree density canopy cover, animal numbers, feeding classes and fire) of vegetation structure and their influence on the herbaceous layer were investigated. Groupings on ecological potential showed 'high' potential areas are less sensitive to animal impact than those classified as 'low' potential. Sustainability, embedded in a forward-looking component viz. Strategic Adaptive Management (SAM) with well-articulated endpoints viz. Thresholds of Potential Concern (TPCs) was used to study fluctuations in animal populations with Connochaetes taurinus (Blue wildebeest) as the case study. The TPC approach provides strong pointers for proactive management aimed at maintaining the system within bands defined by TPCs supporting operationally practical and periodically reviewed objectives.
Determinants of community composition and diversity in KwaZulu-Natal mesic grasslands : evidence from long-term field experiments and pot and plot competition experiments.
(2003) Fynn, Richard Warwick Sinclair.; Kirkman, Kevin Peter.
A predictive understanding of plant community response to various environmental influences (e.g. type, timing and frequency of disturbance, site productivity, fertilization, etc.) is a general goal of plant ecology. This study sought to further understanding of mesic grassland dynamics in KwaZulu-Natal using long-term field experiments (> 50 years) and short-term pot and plot competition experiments. The specific objectives were to: 1) examine the effects of long-term burning of grassland on soil organic matter content because of its potential impact on nitrogen cycling and community composition, 2) examine patterns of community composition and species richness in response to different type, timing and frequency of disturbance (burning and mowing) in a long-term grassland burning and mowing experiment and to different type and amounts of fertilizer application in a long-term grassland fertilization experiment, 3) develop hypotheses concerning the response of different species to disturbance and fertilization, and test these hypotheses using pot and plot competition experiments, and 4) provide a general synthesis of the results of the various field, pot and plot experiments that may be used to develop a predictive theoretical framework for mesic grassland dynamics. Total soil nitrogen was lowest in sites burnt annually, intermediate in sites burnt triennially and highest in sites protected from disturbance and sites mown annually in the dormant-period (spring or winter). Winter burning reduced soil organic carbon and total soil nitrogen more than spring burning. Mineralizable nitrogen was reduced by burning. The different effects of type, timing and frequency of disturbance on total soil nitrogen appeared to be an important determinant of community composition and species richness. Short-grass species (Themeda triandra, Eragrostis capensis, Heteropogon contortus, Diheteropogon amplectens and Eragrostis racemosa) were most abundant in annually burnt sites (especially winter burnt sites), whereas medium and tall-grass species (Eragrostis curvula, Cymbopogon spp., Hyparrhenia hirta and Aristida junciformis) were most abundant in triennially burnt sites, sites protected from disturbance and sites mown annually in the dormant-period, all of which had higher total soil nitrogen than annually or biennially burnt sites. Species richness and short-grass species declined with increasing levels of nitrogen fertilization in the fertilizer experiment and declined with increasing productivity and nitrogen availability in both the fertilizer and burning and mowing experiments. Thus, it was hypothesized that the type, timing and frequency of disturbance resulted in different compositional states through different effects on soil resources (especially nitrogen), which affected the competitive balance between short and tall species. The hypothesis that composition was determined by disturbance-mediated soil nitrogen availability was supported by competition experiments, which revealed that shortgrass species were most competitive in low-nutrient/low-productivity treatments and tall-grass species most competitive in high-nutrient/high-productivity treatments. The fertilizer experiment and a competition experiment revealed that tall broad-leaved species were most competitive in sites of highest productivity, fertilized with both nitrogen and phosphorus, whereas tall narrow-leaved species were most competitive in sites of intermediate productivity, fertilized with nitrogen only. It was hypothesized that summer mowing increased the abundance of short-grass species and decreased the abundance of tall-grass species in the burning and mowing experiment by increasing the competitive ability of short-grass relative to tall-grass species, rather than the tall-grass species being less tolerant of mowing. A competition experiment revealed that tall-grass species (Hyparrhenia hirta and Panicum maximum) were as tolerant of cutting as a short-grass species (Themeda triandra). Themeda triandra was shown to become extremely competitive in cutting treatments, reducing the biomass of most other species relative to their monoculture biomass, showing that its dominance of mown sites in the burning and mowing experiment was a result of its superior competitive ability rather than greater tolerance of mowing. However, many tall erect herbaceous dicots appeared to be intolerant of summer mowing, probably because their meristems are aerial and easily removed by mowing, whereas short creeping herbaceous dicots were increased by summer mowing probably because their meristems were below the mowing height. Further, these short species would be vulnerable to shading in unmown sites. Thus, species with basal meristems (hemicryptophytes) or meristems near the soil surface (geophytes) appear to be more tolerant of mowing than species with aerial meristems (phanerophytes), but the tradeoff is that a low meristem height renders them vulnerable to shading in unmown sites. Very high litter accumulation in the sites protected from disturbance appeared to have a direct influence on community composition and species richness. Species that dominated these sites (e.g. Tristachya leucothrix & Aristida junciformis) initiated tillers below-ground and had sharp erect shoots that appeared to be an adaptation for penetrating litter. Species that initiate tillers below-ground are probably less vulnerable to the effects of shading by litter because their tiller initiation is not dependent on high light availability. The fact that Aristida junciformis was shown to have very low competitive ability in two competition experiments, suggests its dominance of protected sites was through tolerance of high litter levels rather than competitive exclusion of other species. Low grass species richness in these sites was probably a result of an inability of many species to tolerate these high litter levels. This study has revealed that inherent site productivity and its interaction with the effect of disturbance on soil resources and litter levels is a major determinant of community composition and species richness. The effect of type, timing and frequency of disturbance on soil nitrogen was able to account for the principal changes in community composition. Thus, the influence of disturbance on soil nitrogen is a unifying principle in plant ecological theory that enables greater understanding of disturbance-composition relationships. However, intolerance of certain forms of disturbance (e.g. mowing) by species with aerial meristems, or intolerance of accumulating litter in the absence of disturbance by species without sharp erect shoots, may also have important influences on composition. In addition, this study has revealed that plant traits (height, leaf width, position of tiller initiation, shoot morphology and position of meristems) were well correlated with the various effects of disturbance and fertilization on community composition, indicating that a plant trait-productivity-disturbance framework has great potential for understanding and predicting species response to disturbance and multiple limiting nutrients.
Rangeland degradation in the southern Kalahari.
(2000) Van Rooyen, Andre F.; O' Connor, Timothy Gordon.
Observations by local people in the Mier area, southern Kalahari, South Africa, indicated that degraded rangeland does not recover within a time frame acceptable to landowners. Pristine vegetation in this linear dune system consists largely of a herbaceous layer dominated by perennial grasses. Woody vegetation is sparse on dunes and interdunes in good condition. The dunes and interdunes react differently to disturbance, probably because of differences in substrate stability, soil particle size distribution and consequent differences in soil nutrient distribution and moisture content. Degraded dunes are devoid of any vegetation, except for Stipagrostis amabilis, a rhizomatous grass which remain in small clumps, and the tree Acacia haematoxylon. The latter increases in numbers probably due to the high moisture content in degraded dunes. Moisture content in degraded dunes remain high even during prolonged dry periods. Interdunes are more susceptible to degradation and are invaded by the long-lived shrub Rhigozum trichotomum and the annual grass Schmidtia kalahariensis. Both these species compete with perennial grasses for moisture. Additionally, depleted seed banks and increased seed predation by ants (Messor capensis) may also affect the re-establishment of perennial grasses in the interdunes. The main conclusion from this study is that degraded southern Kalahari rangeland cannot recover spontaneously at the landscape scale because of a negative feedback mechanism that prevents establishment and growth of seedlings. The hypothesis put forward is that rangeland ceases to react to rainfall as an ecosystem driver as it becomes degraded. Once degraded, wind controls the dynamics of the system and recovery, irrespective of rainfall, is almost impossible. In contrast, detailed studies at the level of small isolated populations of S. amabilis and S. ciliata on degraded dunes indicate high rates of population growth. The scales at which these processes exist are contradictory and may not be contradicting. Rangeland managers should take cognisance of the influence of the spatial and temporal scale at which they operate, and on which they base their decisions.
A system for supporting wetland management decisions.
(1999) Kotze, Donovan Charles.; Breen, Charles Mackie.; Klug, John R.; Hughes, Jeffrey Colin.; O'Connor, Timothy Gordon.
In South Africa, the loss of wetlands and their associated benefits has been considerable. A need was identified for a system that, using available information, would assist in achieving a balance between local, mainly short-term benefits to individuals and spatially wider and longer term benefits to society. Such a system, termed WETLAND-USE, was developed with the philosophy that:(l) wetlands have been well demonstrated to supply several indirect benefits to society (e.g. water quality enhancement); (2) the impact on these benefits can be described on a qualitative basis using field indicators that characterize the wetland and the disturbance associated with a particular land-use; (3) this information can be communicated to wetland users, which will contribute to achieving a desired balance, provided there is an enabling organizational environment and due consideration is taken of the socio-economic and organizational factors affecting wetland management. The primary conceptual framework underlying WETLAND-USE was the pressure-state-perceptions-policy framework, which depicts: the mode of use (i.e. the pressure); how this affects the state of the system (including its underlying processes and the goods and services it delivers); which in turn shape the perceptions that ultimately determine the policy pertaining to further use. This cycle is repeated at a range of organizational levels from local to national and takes place within a particular socioeconomic context. WETLAND-USE, which was designed for use by fieldworkers, and built using a rule-based, expert system approach, has two main parts, dealing largely with biophysical and social aspects respectively. Part 1, which guides the collection of data relating to the state of the wetland, assists in: (1) predicting the likely impacts of disturbances associated with a proposed land-use (the pressure) on the wetland state, and (2) providing ongoing management guidelines for particular land-uses. Part 2 assists in: (1) describing the social, land tenure and policy contexts of the wetland; and (2) establishing and maintaining organizational arrangements, local policy and management objectives and goals. Several discrete investigations were required for the development and refinement of WETLAND-USE, which was done in an iterative fashion. Initial discrete investigations fed into the development of a prototype system which was refined through evaluation using a questionnaire survey and further discrete investigations. The revised system was re-evaluated using a fieldworkshop approach and, based on the performance of the system in the field, it was revised further to produce the final system. In the two initial discrete studies, protocols were developed for characterizing key physical determinants of wetland functioning, notably: (1) degree of wetness, one of the primary functional determinants, described in the field using readily identifiable soil morphological indicators (e.g. matrix chroma and mottles) and (2) landform setting, which strongly influences local flow patterns and lateral exchange of water and water-borne materials. Graminoid plant species composition and functional groups (defined in terms of photosynthetic pathway) were then described in relation to the above physical determinants, together with rainfall, temperature and soil texture, within wetlands spanning a wide altitudinal range. This revealed that degree of wetness and altitude had the strongest influence over the vegetation parameters examined. An investigation into incorporating cumulative impacts into wetland decision making revealed that consideration should be given to: wetland loss in relation to ecoregions and catchments, and the relation of change in wetland extent, spatial configuration and context respectively to wetland function. Current conservation initiatives in KwaZulu-Natal were shown to account poorly for cumulative impacts on wetlands. Rules of thumb for making such considerations, given severe data limitations, were developed with reference to the high turn-over of species along the altitudinal gradient observed in the vegetation study. The "rules" were than applied to a case-study, the upper Mgeni catchment, as part of an initiative to engage a diversity of stakeholders in wetland information gathering and use. This resulted in the selection of priority wetlands in the catchment and an examination of the extent to which integration had been achieved vertically (across hierarchical levels) and horizontally (across organizations within particular hierarchical levels). In order to broaden the range of land-uses accounted for by the WETLAND-USE prototype, it was applied to a communally used wetland, Mbongolwane, and found to account poorly for the traditional cultivation and vegetation harvesting practices encountered. WETLAND-USE was modified to include a greater diversity of land-use types as well as enhancing its capacity to allow assessments to be conducted using the system's general criteria, thereby making WETLAND-USE more robust. In enhancing the capacity of WETLAND-USE to account for the social and organizational dimension of wetland management, the involvement of local and outside organizations in influencing wetland resource use in five sites was examined in relation to predefined frameworks. The sites, Mandlazini wetland, Mbongolwane wetland, Blood River vlei, Ntabamhlope vlei and Wakkerstroom vlei were chosen to represent a diversity of social contexts and management authorities. This revealed that in communally used areas in particular, a wide range of organizations are involved to varying degrees in influencing the use of different wetland resources. The level to which the local organizational environment contributed to sustainable use varied greatly among wetlands, but in all cases had important deficiencies: (1) self-governing resource-management organizations were largely lacking and in communal areas were weakening under contemporary conditions; and (2) although a formal management system was in place in two of the five wetlands, it was largely absent in the remaining three. There has been little involvement from extension services in facilitating local policy development and in promoting alternative land-uses which have less pressure on the state of the wetland. Local wetland management policy and collaboration among land-owners in wetlands under multiple separate ownership such as Blood River vlei was identified as being particularly poor. The evaluations of WETLAND-USE revealed that, in relation to the underlying philosophy of the thesis, WETLAND-USE had been improved through field application and incorporation of the findings of the discrete investigations. Nevertheless, important limitations of the study were highlighted, including: its high level of reliance on expert opinion in the face of a paucity of empirical data relating to the functioning of local wetlands and their attendant benefits (and how these are affected by anthropogenic disturbances), and a particularly shallow representation of socio-economic factors. The identification of these limitations was useful in highlighting key areas for further research.
Impact of stocking rate, livestock type and livestock movement on sustainable utilisation of sourveld.
(1999) Kirkman, Kevin Peter.; Tainton, Neil M.; O'Connor, Timothy Gordon.
Data collected between 1992/93 and 1996/97 from two long-term grazing trials were used to investigate the interaction between grazing animals and veld grass. In the first trial, the impacts of stocking rate and time of stocking in spring on both livestock performance and veld vigour (defined as the ability of a grass plant to regrow after defoliation) and condition were investigated. In the second trial comparisons were made, firstly between the impacts of sheep and cattle grazing, and secondly between various types and frequencies of rest, on veld vigour and condition. Treatments applied in the first trial comprised four stocking rates, namely 7, 10, 13 and 16 sheep ha¯¹ for the duration of the grazing season, and two times of stocking, namely as early as possible after spring burning and three weeks later. Sheep grazed each treatment continuously throughout the growing season. Treatments were applied to alternate blocks in a two-year cycle with each block resting for a year within a grazing cycle. Animal performance (mass gains over the season) was measured to quantify livestock performance. Herbage availability was measured on a species basis at intervals throughout each season using a dry-weight-rank procedure to determine grazing patterns. Residual effects of the grazing treatments on vigour were determined by measuring herbage regrowth on a species basis during the rest season which followed a season of grazing and comparing these measures to a previously ungrazed control treatment. Effects of the grazing treatment on proportional species composition were determined using a nearest plant point technique. Stocking rate had a non-linear effect on livestock performance, with livestock performance on the lightest stocking rate being less than on the two intermediate stocking rates. The mass gains on the heaviest stocking rate were generally the smallest. Delaying the time of stocking in spring resulted in smaller mass gains during the resultant shorter season. The sheep from both the early and late time of stocking groups had similar mean masses at the end of the season. The advantage of stocking early can thus be attributed more to saving the cost of alternative feed for the interim period than to additional mass gains due to stocking early. Quantification of livestock performance in terms of selected and available feed quality, quantity and species availability throughout each season was extremely complex due to multiple thresholds in the measured variables and no simple cause and effect relations could be established that would hold for spatial or temporal extrapolation. The negative impact of grazing on veld vigour was severe. Stocking rate and time of stocking had a secondary impact with the vigour loss positively related to increasing grazing pressure. The main factor influencing vigour loss was grazing, irrespective of time of stocking or stocking rate, as opposed to no grazing. The impact of grazing on vigour was severely negative in the palatable species, variable in the species of intermediate palatability and positive in the unpalatable species that were rarely, if ever, grazed. The stocking rate and time of stocking rate had an impact on the proportional species composition, with the more palatable species declining in proportion. There was an observable relation between impact of grazing on vigour and on species composition. Treatments applied in the second trial involved applying a full growing season rest in alternate years, half a growing season rest (late season) in alternate years and no rest to veld grazed by sheep or cattle at similar stocking rates. Residual effects of the treatments on veld vigour were determined by measuring species regrowth using a dry-weight-rank technique during the season following treatment application, and comparing it to controls ungrazed for one and two seasons respectively. Changes in proportional species composition were determined using a nearest plant point technique. The vigour of veld grazed by sheep declined rapidly relative to veld grazed by cattle. The vigour of palatable species was severely impacted, vigour of intermediate species was variably impacted and vigour of unpalatable species increased dramatically on veld grazed by sheep compared to the control treatments. Similar trends occurred in veld grazed by cattle, but to a lesser degree. Resting was beneficial for vigour recovery in both sheep and cattle treatments but it seems that the grazing treatment between rests has a greater influence on the veld vigour and condition than the rest itself. The veld grazed by sheep remained at a substantially lower productivity level than veld grazed by cattle. This was particularly evident in the change in productivity balance between palatable and unpalatable species in the sheep treatments, where palatable species vigour declined and unpalatable species vigour increased relative to veld grazing by cattle. Species composition of veld grazed by sheep deteriorated over the trial period in contrast to the veld grazed by cattle, which improved in species composition. Grazing management recommendations for sourveld should include a bias towards cattle, optimising stocking rate for improved performance and resting for enhancing vigour of the palatable grasses.
Host specificity in South African mistletoes.
(2013) Okubamichael, Desale Yosief.; Ward, David Mercer.; Griffiths, Megan Elizabeth.
Mistletoes intimately connect to their host trees with a haustorium that allows them to access nutrients and water. Mistletoes in South Africa vary greatly in their degree of host specificity. Most species occur on a wide range of host families, while others are restricted to a single host family or—at the extreme—to a single host species. Mistletoes that are host generalists at a larger spatial scale may become host-specific at a local scale. One of the challenges in mistletoe biology is determining the factors that maintain local host specificity. Birds potentially reinforce the mistletoe–host interactions by direct dispersal. However, many mistletoe species coexist while parasitising different co-occurring host species. This suggests that host trees may impose more selection pressure than birds in determining host specificity. Thus, my thesis examines the role of host trees as ecological and physiological filters that influence the infection patterns and determine host specificity of mistletoes in South Africa. The second chapter of this thesis synthesises the literature on host specificity in mistletoes. I then present the results of four field and laboratory experiments that were used examine the features affecting host specificity in representatives of two families of mistletoes (Loranthaceae and Viscaceae) in South Africa. My main research objectives focus on host abundance and morphology, host compatibility, host water and nutrient content, abiotic influences on mistletoe seedling survival and growth and mistletoe–host stomatal morphology in relation to water potential that affect nutrient acquisition by mistletoes from their host trees. The geographic mosaic approach was explored as a potential explanation for the mistletoe–host interactions that direct host specificity in mistletoes. I synthesised the available literature on the mechanisms and factors that direct mistletoe host specificity. This was supported by data analysed from South African herbarium collections, books describing the South African flora and field observations in South Africa. I suggest that host abundance (host availability through time and space) and host compatibility (as determined by genetic, morphological, physiological and chemical factors) play a primary role in determining host specificity in South African mistletoes, while differential bird dispersal strengthens or weakens mistletoe–host interactions. Analysis of the network structure of mistletoe–host interactions at different levels (e.g., at the level of population, species and genus) followed by genetic and reciprocal germination experiments may reveal the patterns and mechanisms of host specificity in mistletoes. I quantified the mistletoe–host composition, height of potential host trees and nutrient and water content of mistletoes and their hosts at Pniel Estates. Surveys of the study site revealed a single mistletoe species, Viscum rotundifolium, parasitising only Ziziphus mucronata and Ehretia rigida. Both parasitised host species were not the most abundant trees, were not the tallest trees and did not have the highest water or nutrient content of trees in the area, although these factors have been found to be good predictors for mistletoe parasitism in other studies. Subsequently, I tested mistletoe–host compatibility by conducting a germination experiment in the greenhouse by inculcating seeds of V. rotundifolium on freshly cut branches of nine available potential host trees. I found that mistletoe seeds had a greater chance of attachment and subsequent survival on branches of E. rigida and Z. mucronata as compared with seeds on co-occurring Acacia and other potential host species. This suggests that host compatibility plays a role in directing the host specificity of V. rotundifolium at Pniel Estates. I found that individuals of V. rotundifolium had more negative water potentials than their host trees and, by doing so, they passively maintain the flow of nutrients. In addition, I found evidence that the mistletoe uses active uptake to access nutrients from host phloem because the leaf tissue of a mistletoe had a nitrogen-to-calcium ratio (N:Ca) >1. Conventionally, a high N:Ca ratio (>1) in the leaf tissue of a mistletoe is taken as evidence of active uptake from host phloem because N is highly phloem-mobile while Ca is a large molecule and is phloem-immobile. This method has shortcomings discussed at greater length in the chapter but my findings suggest that the mistletoe V. rotundifolium uses a combination of passive and active nutrient uptake. I quantified the mistletoe–host community composition and host physical features (height and diameter at breast height) in two sites in KwaZulu-Natal, South Africa—Highover and Mtontwane. The mistletoe Agelanthus natalitius (Loranthaceae) is common at both sites, parasitising the most abundant host species—Acacia karroo—and the second most abundant host tree—Acacia caffra. Prevalence of mistletoe infection (percentage of trees parasitised) was positively correlated with tree size (height and diameter at breast height). The two host species did not differ significantly in height. At Highover the host species A. caffra and A. karroo had a similar prevalence of mistletoe infection but at Mtontwane a significantly higher percentage of A. caffra trees was parasitised in comparison with A. karroo. However, the intensity of mistletoe infection (mean number of mistletoes per tree) was lower for A. caffra (Highover: 0.66 ± 0.01, Mtontwane: 0.89 ± 0.04) than for A. karroo (Highover: 0.73 ± 0.04, Mtontwane: 1.03 ± 0.64). There were two highly infected big trees in Highover and one in Mtontwane where many mistletoe-dispersing birds were nesting which inflated the numbers for intensity of mistletoe infection in A. caffra, however. I tested mistletoe–host compatibility by conducting a reciprocal transplant experiment in the two study sites. I applied a paired design, using one local and one non-local mistletoe seed in each pair, with seed pairs placed on the two main host species at the different sites. Except in Highover where an unidentified pathogen retarded growth and survival, mistletoe seeds placed on the same substrate and in the same site as their source host grew a longer hypocotyl and had greater survival. Regardless of source, mistletoes placed on A. karroo had longer hypocotyls and greater survival than mistletoes on A. caffra. These results suggest that there may be adaptation of the mistletoe Agelanthus natalitius to the most frequently encountered host species, Acacia karroo. To simulate the conditions encountered by mistletoes during the dry and cold South African winter, mistletoe seedlings were monitored at different levels of microclimate (light, temperature and moisture) in a growth chamber. I found that higher light availability (20% and 40% shade versus 80% shade), cool temperatures (15°C and 20°C versus 25°C) and continuous moisture availability improved seedling development and subsequent survival of two mistletoe species (Viscum rotundifolium and Agelanthus natalitius). I studied the leaf stomata of two host–mistletoe pairs (Acacia karroo–Agelanthus natalitius and Vitex obovata–Erianthemum dregei) using a scanning electron microscope to investigate some of the underlying mechanisms that enable mistletoes to maintain more negative water potentials than their host trees and at the same time control water loss. In addition, I examined the response of mistletoes to the application of abscisic acid (ABA), a plant growth regulator that controls stomatal closure. I found that the mistletoes had a higher density of stomata and had larger stomata than their host trees. In addition, both mistletoe and host leaves closed their stomata during midday and in response to exogenous ABA. The ability of mistletoes to control water loss in this way may be one reason why mistletoes rarely kill their host trees, which would be maladaptive. The mistletoes used in my studies are known to be host generalists at a larger spatial scale but I found that they were host specific at a local scale. The results of my research suggest that host abundance and compatibility play a role in directing host specificity, while host nutrient and water status have little effect on host specificity at this local scale. The interactions between the generalist mistletoes used in my studies and their hosts are likely to vary over the geographic ranges of the mistletoe and alternate among different hosts. This may create multiple locally host–specific mistletoe populations and produce a complex geographic mosaic of mistletoe–host combinations across space and time. I suggest that mistletoe populations in South Africa may comprise numerous lineages incapable of parasitising the full range of host species, which could potentially lead to the formation of distinct host races over time. In the future, it would be interesting to document the infection patterns of these generalist mistletoe species across their entire geographic ranges in southern Africa, with particular focus on the patterns of mistletoe infection in places where the host abundance changes among sites. Host preferences may vary with changes in host frequency and host community composition. This could be paired with reciprocal transplant germination experiments in several sites to ascertain whether the mistletoe species have higher fitness on the most locally abundant hosts.
Short-term effects of mixed grazing by cattle and sheep in highland sourveld.
(1994) Hardy, Mark Benedict.; Tainton, Neil M.
Data derived from a long-term grazing trial were used to determine short-term effects of mixed grazing by cattle and sheep in Highland Sourveld. Five cattle to sheep ratio treatments (viz. 1 :0, 3:1, 1 :1, 1 :3, 0:1) were applied, each at three stocking rates (viz. 0.5 (low), 0.71 (medium) and 1.0 (high) animal unit equivalents (AUE) ha¯¹). Ratios were expressed in terms of AUE cattle: AUE sheep. Stocking rate and ratio treatments were balanced at the start of each grazing season. Fourteen-month old Hereford steers and 'two-tooth' Merino wethers were used as experimental animals. The trial comprised two components, viz. an animal production component and a simulated component. The animal production component was conducted only in the medium stocking rate treatment where the effect cattle to sheep ratio on the performance of cattle and sheep was determined for each of four grazing seasons (viz. 1989/90, four-paddock rotational grazing The low and high stocking rate 1990/91, 1991/92, 1992/93). A management system was applied. treatments were implemented by simulating a four-paddock rotational grazing system and using a single paddock for each stocking rate/ratio combination. Grazing of the simulated treatments coincided with the grazing of a specific 'test' paddock in each ratio of the medium stocking rate treatment. These 'test' paddocks and the simulated treatment paddocks were monitored to determine the impact of mixed grazing by cattle and sheep on individual grass plants and the sward. The whole of the experimental area was rested during the growing season prior to the start of the trial and all paddocks were burnt in the dormant season just before the start of the first grazing season. The trial therefore commenced with a with a uniform sward of immature herbage in all paddocks allocated to each group of animals. ln the second and subsequent growing seasons only those paddocks which had been rested in the previous season were burned prior to the onset of growth in early spring. Rainfall for the first three grazing seasons was similar to the long-term mean of 790mm whilst the last season was considered 'dry' with 554mm recorded during 1992/93. In all grazing seasons, as the proportion of cattle in the species mix increased, sheep performance increased. A decline in sheep performance was recorded in each ratio treatment from the first to the third season. This decline was attributed to the increased maturity and thus lower quality of herbage on offer to the sheep, and the fact that only one paddock available to the animals in the second and third grazing seasons had been burned prior to the start of the season. Sheep performed best during the 'dry' season where herbage quality was maintained for longer into the grazing season than in previous seasons. In contrast, cattle performance was affected by the stocking rate (animals ha¯¹) of cattle rather than the presence of sheep. As the quantity of herbage on offer per steer declined steer performance declined. Animal performance data were also used to predict the effects of adding cattle to a sheep production enterprise and vice versa. The general trends were that the introduction of cattle into a low stocking rate, sheep-only production enterprise would allow for an increase in the stocking rate of sheep while maintaining the performance of the sheep. In this way the carrying capacity of a farm may be improved. Stocki ng rate and ratio treatments varied from those established at the start of each grazing season due to the differential performance of the cattle and sheep in each treatment. As the proportion of cattle in the species mix increased, stocking rate increased and the ratio widened in favour of cattle. Stocking rate (AUE ha¯¹), calculated a posteriori for each season, was the major influence on the severity of grazing on individual plants and within patches. As stocking rate of cattle and sheep at the various ratios increased, the extent and severity of grazing increased. At stocking rates in excess of O.8AUE ha¯¹ however, sheep-only grazing resulted in a greater proportion of plants, per species and per area, being grazed more severely than was the case for an equivalent stocking rate (AUE ha¯¹ ) of cattle. Furthermore, cattle and sheep had similar effects on patch size distribution when stocked at the same number of AUE ha¯¹. There were no measurable effects of stocking rate and ratio on proportional species composition and basal cover over a two year monitoring period. A technique for estimating basal cover in tufted grasslands was developed and is presented as an appendix to the thesis. Data were also used to evaluate the use of AUE as an integral part of the grazing capaci ty concept. Results indicated that cattle and sheep cannot be equated in terms of AUE when referring to the grazing impact. It is suggested that the definition of grazing capacity should include the species of livestock and assume a grazing management system appropriate to the grazing habit of the animals concerned. Resul ts of the trial provide strong indications that, in the long-term, the current recommendations of grazing cattle together with sheep in order to prevent the degradation or loss of veld condition which occurs in sheep-only grazing systems, will not succeed. A four-paddock rotation grazing system does not appear to be an appropriate veld management system for sustainable sheep production in sour grassveld. An alternative approach to veld management is suggested in which the sheep are confined to only those areas of the farm which were burnt at the start of the grazing season. Ideally, sheep should not allocated to the same paddock for two consecutive grazing seasons.
The nitrogen economy of three irrigated temperate grass pastures with and without clover in Natal.
(1994) Eckard, Richard John.; Tainton, Neil M.
Abstract can be viewed in PDF document.
Effects of herbivores, fire and harvesting on the population dynamics of Acacia drepanolobium sjoestedt in Laikipia, Kenya.
(2007) Okello, Bell Dedan.; O'Connor, Timothy Gordon.
Effects of herbivory, fire, and tree harvesting on Acacia drepanolobium were studied using plant population dynamics as the philosophical basis of research. Specifically, growth rates, chrono-sequence of re-growth, biomass and charcoal yield, herbivory, flowering, seed production, germination, mortality and the ants of Acacia drepanolobium were studied in the black cotton ecosystem of Mpala Research Centre, Laikipia, Kenya, between September 1995 and December 2000. Acacia drepanolobium was the most abundant tree or shrub with densities ranging from 80% to 98% of all the overstorey species, but it was the least browsed of all the trees and shrubs in the black cotton ecosystem, ranging from a mean of 7.2% to 9% of the individuals browsed. The tree is inhabited by four Acacia ant species, Crematogaster mimosae, Crematogaster sjoestedti, Crematogaster nigriceps, and Tetraponera penzigi, which are believed to be obligate, and which probably play a role in the low browsing rates observed. Six herbivore treatments replicated three times (no herbivores - O; only cattle - C, all herbivores allowed - MWC {control}, mega-herbivores {elephants and giraffe} and wildlife {W} – MW only, wildlife – W - only {all wildlife except mega-herbivores} and wildlife and cattle only - WC) was the main experimental design used in understanding the dynamics of the tree species under influence of different herbivores. Mean annual height growth of A. drepanolobium trees was 24.9 cm yr-1, while the mean Relative Growth Rates ranged from 14.6 x 10 –3 cm cm-1 yr-1 to 18.7 x 10 –3 cm cm-1 yr-1. Growth rates were different among the herbivore treatments and between seasons. Shoots of the tree grew by a mean range of 6.8 cm to 9.1 cm, were similar among the treatments but differed among the seasons. Canopy volume increased over time although it fluctuated with seasons, suggesting an increase in bushiness of A. drepanolobium in the ecosystem. Trees occupied by different ant species showed differences in shoot density (number of new shoots per twig), being greater in Crematogaster nigriceps occupied trees compared with the other ant species. Swollen thorn (gall) density per unit of twig length was greatest in treatments with megaherbivores; these galls were significantly larger on trees occupied by the ant Crematogaster nigriceps. Treatments with herbivores were more spinescent than the total exclusion treatment. Spine lengths ranged from 0.8 to 2.4 cm, and recorded a progressive reduction of up to 36.36.7% in treatments without browsers suggesting a relaxation of induced defence in A. drepanolobium. Flowering in A. drepanolobium was low and staggered over the study period ranging from 0.8% to 2.0% of the trees with no differences among the treatments suggesting that the level of herbivory was not sufficient to influence reproduction of the tree in the experimental site. Consequently, seedling recruitment was very low within the experimental site. However, a nearby site recorded flowering of between 22.7% and 93.5%. Mean pod production, mean number of seeds per tree and mean weight of pods and seeds had a positive linear relationship with tree density (R2=0.77, 0.81 and 0.81 respectively). Trees occupied by Crematogaster mimosae were the most likely to flower (68%) compared with C. nigriceps (5.8%), again suggesting that ants had an effect on the tree’s reproduction. Mortality of A. drepanolobium trees averaged 0.9% to 4.2% over the study period, being significantly greater in treatments with mega-herbivores. Seedling survival ranged from 42% to 75%, being greatest in the cattle only treatment. Between 30% and 100% (mean 67.2%), of A. drepanolobium seeds were attacked by a bruchid beetle (Bruchus sp.). Seeds attacked by bruchid beetles had significantly lower germination rates. Similarly, seeds passed through a fire also recoded significantly low germination rates compared with normal seeds. Fire (3.4%) and bruchid beetles (20.7%) germination compared with (control) undamaged seeds (84%) play an important role in the population dynamics of A. drepanolobium. Fewer A. drepanolobium seeds (33%) were recovered from the surface compared with buried (72%) seeds after a fire, indicating seed loss from the effect of fire and predation. In the burn experiment, fire top-killed 16% of A. drepanolobium trees but no tree or seedling was killed. On the other hand, fire significantly reduced the density of non-A. drepanolobium trees by between 50% and 100%, with none of them showing signs of coppicing after the fire unlike top-killed A. drepanolobium trees. Woody biomass from A. drepanolobium was strongly related to stem diameter (Y = 3.77x + 1.17, R2 = 0.96, P < 0.001). Mean charcoal production from earthen kilns was 2.83 Mg ha- 1. Height and stem diameter in coppicing stands increased at a mean rate of 28.6 cm yr-1 and 0.7 cm yr-1 respectively. Biomass in coppicing stands accumulated at a mean rate of 1.3 Mg ha-1 yr-1 in a 14-year period, yielding dry biomass of 18.26 Mg ha-1 useable wood that can produce a minimum of 3.0 Mg ha-1 of charcoal. This study shows that Acacia drepanolobium populations are affected by several factors including herbivory, fire and ants. The population dynamics of this tree shows that it can be harvested for sustainable charcoal yield over a 14-year cycle.
Recovery, resilience and stability of piospere systems in the Kruger National Park.
(2010) Matchett, Katherine Jean.; Kirkman, Kevin Peter.; Ward, David Mercer.; Peel, Michael John Stephen.; Morris, Craig Duncan.
Water provision is an important tool in the management of savanna ecosystems. Artificial water sources are a potential focus for degradation (biodiversity and loss of ecosystem resilience at a range of spatio-temporal scales), because they alter plant-animal interactions and soil function and stability, through the creation of piospheres. This study was undertaken as part of a drive by the Kruger National Park (KNP) to enable managers to integrate artificial waterhole management (e.g. waterhole closure or rotation) when setting goals for heterogeneity and biodiversity conservation in the park. The over-arching goal was to quantify the relationship between water provision and different attributes of heterogeneity, as part of a broader initiative to place water provision and piospheres within an ecosystem threshold framework. Herbivore utilisation gradients (piospheres) around artificial waterholes in the KNP, described in 1990, were resurveyed in 2006-2007, against a backdrop of waterhole closure in the KNP, to contribute to an understanding of the factors governing recovery and resilience in grazing systems. The responses of the plant community and soil parameters to a relaxation of herbivore utilisation pressure around closed waterholes (recovering piospheres) were examined, as were changes in the same parameters at sites that have remained open (active piospheres). These ecosystem properties were considered in relation to structural and functional ecosystem thresholds, and the piospheres surveyed incorporated a range of rainfall and edaphic gradients in the KNP. Herbaceous basal cover and soil infiltration capacity both increased significantly between 1990 and 2006/7, regardless of waterhole status. This was linked to higher rainfall in 2006/7, compared to 1990. The only vegetation variables to respond consistently to distance from water were the remote-sensed Normalized Difference Vegetation Index (NDVI) and herbaceous species composition. NDVI increased with distance from water, and annual grasses and forbs were most abundant close to water. Perennial, disturbance-sensitive climax species increased in abundance further from water. Soil analyses (N, P, pH, organic matter, and texture) and field measurements (infiltration, compaction) revealed no systematic piosphere patterns. Waterhole closure did not result in soil or vegetation recovery, but piosphere intermittency and the increases of basal cover and infiltration rate indicated that ecosystem resilience has not been compromised vii by long-term artificial water provision in the KNP. This study has shown that the traditional piosphere model is of limited use in sub-humid savanna ecosystems like the KNP during above-average rainfall periods.
An evaluation of irrigated ryegrass/clover pastures for fat lamb production in the Highland Sourveld of Natal.
(1986) Du Plessis, Thomas Marius.; Bransby, David Ian.
The primary objective of this study was to increase net farm income in the Highland Sourveld of Natal by developing pasture based fat lamb production systems. Secondary objectives included (a) determination of the most suitable pasture species or species mixture; (b) determination of the optimum level of N fertilization; (c) establishment of biological and economic optimum stocking rates; (d) projection of expected profit and (e) verification of proposals on a farm scale. Midmar Italian ryegrass was compared to selected legumes alone, and in mixtures with ryegrass in a small plot trial conducted over two years at a 3- and 6-week harvesting frequency under irrigation. Ryegrass in combination with red and white clover provided the highest yields for both harvesting intervals in both years. When this pasture was subsequently compared to ryegrass alone under grazing, no significant difference could be detected in lamb gains at 20 and 28 ewe-lamb units/ha, but at 36 ewe-lamb units/ha the lambs on ryegrass started losing weight at 30 kg, while those on ryegrass + clover continued to gain until a market weight of 40 kg was achieved. In a small plot trial little response was detected to added N above 175 kg N/ha on ryegrass + clover, and under grazing, no significant difference was observed between 275; 375 and 4,75 kg N/ha over two single year periods. Relationships between average daily gain (ADG) and stocking rate, ADG and disc meter height, and stocking rate and disc meter height were mostly well described by linear functions. Expected pre-weaning AOG was 150 g higher (p ≤ 0,01) than post-weaning ADG (100 g) in the first year, and a similar difference was observed in the second year. For the period common to all stocking rates an average maximum gain/ha of 824 kg was achieved at a stocking rate of 33,5 lambs/ha and a disc meter height of 4,5 cm. By lambing in April, lambs could be marketed in September at 40 kg and more weaned lambs could be grazed on the pasture and marketed in December, leading to an expected gain/ha of 1400 to 2200 kg, and expected profit/ha of R 1254 to R 2771, depending on stocking rate. When evaluated under farm conditions, this system produced 2060 kg live weight/ha and a profit of R 3206/ha. It is concluded from the present study that the fat lamb production system based on irrigated ryegrass + clover pastures can be recommended with confidence to farmers in the Highland Sourveld of Natal.
Beef production from kikuyu and Italian ryegrass.
(1985) Bartholomew, Peter Edward.; Tainton, Neil M.
Four grazing trials to characterise cultivated pastures, in terms of beef production, were conducted in Bioclimate 3 of Natal. Dual purpose and British beef type cows were run on kikuyu at stocking rates from 2,81 to 7,30 cows plus calves per ha. During the eight seasons of the trial the seasonal rainfall varied from 580 to 933 mm. There was a positive linear relationship between rainfall and pasture yield with maximum yield of kikuyu being recorded during February - March. Stocking rate affected pasture yields only during favourable rainfall seasons. Crude protein (CP) and crude fibre (CF) of kikuyu fluctuated markedly within and between seasons. However, CP increased and CF decreased as stocking rate increased. There were significant relationships between stocking rate and (a) calf performance, (b) calf livemass gain, (c) period required to attain maximum mass, (d) period on pasture for the cows, and (e) cow mass change: Weaners were run on irrigated Italian ryegrass at 5, 7 and 9 weaners per ha for four seasons. Stocking rate had little effect on the growth pattern of the pasture but affected dry matter yields. Reducing the stocking rate resulted in increased pasture yields and CF content but reduced CP levels of material on offer. Steers exhibited higher gains than heifers but lower carcass grades and stocking rates for maximum gain per ha (SRmax). Livemass gains of 1315 and 1224 kg per ha can be expected at SRmax of 6,85 and 9,54 for steers and heifers respectively. Yearling heifers run at four stocking rates on kikuyu for one season showed a negative linear relationship between stocking rate and gain and a positive linear relationship between pasture height and gain. A SRmax of 8,85 allows for a livemass gain of 1 040 kg per ha. The effect of feeding concentrates on foggaged kikuyu was evaluated. Foggaged kikuyu can be used as a source of roughage for fattening steers. However, as the steers became adapted to the concentrate the intake of kikuyu declined from 39 to 19% of their daily intake. Regressions derived from the characterisation trials allow for developing beef systems for different situations.
Development and testing of a remote controlled oesophageal fistula valve for goats.
(1993) Raats, Jan Gabriël.; Tainton, Neil M.; Trollope, Winston Smuts Watts.
A remote control sampling technique was developed for the collection of oesophageal fistula samples from goats. Number and size of samples can be varied and collected throughout the day without disturbing the animal's normal feeding behaviour. The equipment developed and tested in this study consists of an oesophageal fistula valve which allows the fistula to be opened and closed, a rechargeable battery pack and motor to operate the valve, a portable radio and receiver to activate the valve motor, and a harness to attach the equipment to the body of the animal In addition, a closing device to effectively seal large oesophageal fistulae (> 1 050 mm²), which in turn is required to accommodate the valve, was developed. During field tests with the fistula valve, 10 % of I 027 sampling attempts failed due to blockage of the valve, and an average of 1.3 incidences of equipment failure were recorded per animal per sampling day, from an average of 9.9 extrusa collections per day. Observed feeding behaviour (grazing/browsing) as well as grass / bush ratio of fistula valve and standard fistula bag samples of four goats, formed the basis for the evaluation of this technique. In addition, extrusa recovery rates, measured under controlled conditions, were used in the evaluation of this sampling method. Differences in extrusa composition between the fistula valve and fistula bag techniques varied substantially during the browsing period within a camp and also between camps. Furthermore, during high frequencies of observed grazing, there were large differences between the fistula valve and fistula bag methods. During this study, the fistula valve technique provided a more realistic estimate (R²=.91) of the observed feeding behaviour of goats than the fistula bag technique (R²=.63). Under controlled conditions, the large oesophageal fistula, with or without the valve, enables high and consistent extrusa recovery rates (87 % recovery; SD 7.5).
The bioeconomic implications of various stocking strategies in the semi- arid savanna of Natal.
(1994) Hatch, Grant Peter.; Tainton, Neil M.; Ortmann, Gerald Friedel.
Climatic and market uncertainty present major challenges to livestock producers in arid and semi-arid environments. Range managers require detailed information on biological and economic components of the system in order to formulate stocking strategies which maximise short-term financial risk and minimise long-term ecological risk. Computer-based simulation models may provide useful tools to assist in this decision process. This thesis outlines the development of a bioeconomic stocking model for the semi-arid savanna of Natal. Grazing trials were established at two sites (Llanwarne and Dordrecht) on Llanwarne Estates in the Magudu area of the semi-arid savanna or Lowveld of Natal. The Lowveld comprises a herbaceous layer dominated by Themeda triandra, Panicum maximum and P. coloratum and a woody layer characterised by Acacia species. The sites differed initially in range composition. Llanwarne was dominated by Themeda triandra, Panicum maximum and P. coloratum, while Dordrecht with a history of heavy stocking was dominated by Urochloa mosambicensis, Sporobolus nitens and S. iocladus. Three treatments were stocked with Brahman-cross cattle at each site to initially represent 'light'(0.17 LSU ha-'), 'intermediate' (0.23 LSU ha-') and 'heavy' (0.30 LSU ha-') stocking. Data collected at three-week intervals over seven seasons (November 1986 to June 1993 or 120 measuring periods) provided the basis for the development of a stocking model LOWBEEF - OWveid BioEconomic Efficiency Forecasting) which comprised two biological sub-models (GRASS and BEEF), based on step-wise multiple linear regression models, and an integrated economic component (ECON). The GRASS model predicted the amount of residual herbage at the end of summer (kg ha-') and the forage deficit period (days) over which forage supplementation would be required to maintain animal mass. Residual herbage mass at the end of summer (kg ha-') was significantly related (P < 0.01) to cumulative summer grazing days (LSU gd ha-'), rainfall (mm) (measured 1 July to 30 June) and range condition (indexed as the sum of the proportions of T. triandra, P. maximum and P. coloratum). The forage deficit period (days) over which herbage mass declined below a grazing cut-off of 1695 kg ha-' was significantly related (P < 0.01) to residual herbage mass at the end of summer. The BEEF model predicted the livemass gain over summer (kg ha¯¹) which was significantly related (P< 0.01) to rainfall (mm) stocking rate (LSU ha¯¹) but interestingly not to condition. The economic component (ECON) reflected the difference between gross income (R ha¯¹) and total costs, which were based on fixed and variable cost structures (using 1993 Rands), including demand-related winter costs, to reflect net returns to land management (R ha¯¹). A conceptual model of range dynamics based on three discrete states, was to developed to summarise the effects of rainfall and stocking rate in semi-arid savanna. State 1, characterised by iocladus and S. nitens, was associated with heavy stocking. Movement towards State 2, characterised by T. triandra and P. maximum, was associated with periods of above-average rainfall. Drought conditions, which comprised a major system disturbance led to stability at State 3, dominated by U. mosambicensis. Post-drought recovery was influenced by predrought composition and stocking levels where tuft numbers, basal cover and seedbank were significantly reduced by increased stocking within a sward dominated by species of low stature such as Aristida congesta subsp Urochloa mosambicensis, Sporobolus nitens, Sporobolus iocladus and Tragus racemosa. It was suggested that extensive soil loss may lead to stabilisation across an irreversible threshold at a forth state characterised by shallow species such as Tragus racemosa Aristida congesta subsp. congesta. Sensitivity of optimum economic stocking rate net return to price and interest rate fluctuations, and wage and feed cost increases were examined for various rainfall and range condition scenarios. Net return and optimum economic stocking rate increased as rainfall and range condition increased through the effect of increased residual herbage mass at the end summer, decreased forage deficit periods and reduced supplementary feed costs. Net return was highly responsive to changes beef price where an increase in beef price led to an increase in optimum economic stocking rate and net return. The effect of reduced prices may be compounded by dry where supply-driven decreases in price may occur. This suggested that for dry seasons the optimum stocking rate was the lightest within the range of economic stocking rates. Although an increase in interest rates would increase variable costs and lead to reduced returns, the influence of interest rates on enterprises will vary in relation to farm debt loads. Increased labour costs would result in a corresponding decline in net return although optimum economic stocking rate would remain unaffected. Increased supplementary feed cost had little influence on net return relative to the effect of demand-driven increases in feed costs as rainfall decreased. The distribution of net returns for stocking strategies of 0.20, 0.30 and 0.40 LSU ha¯¹ and climate-dependent stocking (where stocking levels were varied in relation to rainfall and hence forage availability) and range condition scores of 10, 50, 80 and a dynamic range model were examined for a 60 year rainfall sequence (1931-1991). While a range score of 10 would see residual herbage mass decline to below a grazing cut-off of 1695 kg ha¯¹ before the end of summer, a range score of 80 suggested that, irrespective of stocking strategy within the range investigated, herbage would not become limiting. This suggested that irrespective of stocking strategy a range score of 10, established across an irreversible soil loss threshold, would reflect accumulated losses over the 60 year period. In contrast, a range score of 80 would lead to positive accumulated returns. A dynamic range model (where range composition was related to previous seasons rainfall) and a climate-dependent stocking strategy, suggested that herbage would not become limiting by the end of summer and forage deficit periods would be restricted to an average of 88 days per year. Such an approach would yield a higher accumulated cash surplus than fixed stocking strategies. Incorporation of stochastic rainfall effects allowed the development of cumulative probability distributions based on 800-year simulations to evaluate the risk associated with various stocking strategies. Range condition played a major role in determining the risk of financial loss where decreased range condition was associated with enhanced risk. An increase in stocking rate resulted in increased variability in returns. Although the risk of forage deficits and financial losses may be reduced with lighter stocking, this may be at the cost of reduced returns during wetter seasons. Increased stocking may increase the probability of higher returns during wetter seasons although this may at the cost of increased risk of forage deficits and highly negative returns during dry seasons. Importantly, ecological risk may increase as stocking is increased. A flexible or climate-dependent strategy, where stock numbers are adjusted according to previous seasons rainfall, combine financial benefits of each approach and reduce financial risk. Although errors may carry high ecological costs where, for example, the effect of an above-average rainfall season would be to increase stock numbers into a subsequent dry season, the probability of incurring such error was low. Current livestock production systems in the semi-arid savanna of Natal based on breeding stock may not be appropriate in a highly variable environment where low rainfall may require extended periods of upplementary feeding or force the sale of breeding stock. A change in emphasis from current systems to a mixed breeding system, where the level of breeding stock would be set at the optimum economic stocking rate for drier seasons, may decrease both financial and ecological risk. Growing stock may either be retained or purchased during wetter seasons to reach the optimum economic stocking rate for such seasons. although growing stock may display a greater tolerance to restricted intake (during dry seasons) than would breeding stock, additional growing may be rapidly sold in response to declining rainfall with no influence on the breeding system. Integration of wildlife into current cattle systems may be an important means of reducing financial risk associated with variable rainfall and profitability and ecological risk associated with woody plant encroachment.
Towards formalized adaptive management in succulent valley bushveld.
(1993) Stuart-Hill, Gregory Colin.; Tainton, Neil M.
This study was designed to provide the means for implementing formal scientific vegetation management 1n the succulent valley bushveld of the eastern Cape, South Africa. Nowhere in the world has a detailed, effective and practical veld management system being developed entirely from research, and even the most successful management systems rely heavily on the intuition of people. A process, formally called 'adaptive management', combines this intuition with scientific testing and the overall objective of this study was to provide a framework for formalized adaptive management in succulent valley bushveld. On analyzing the process of adaptive management, the following knowledge 'tools' were identified: (i) a management system for immediate implementation; (ii) a technique for vegetation assessment; (iii) a technique for monitoring vegetation change; (iv) a technique for monitoring forage use and recovery; (v) a list of key forage species; (vi) a model to set initial stocking rates; (vii) a method of recording essential information; and (viii) a database of ecological principles. Providing these 'tools' became the goals of this study. These topics covered almost all facets of rangeland science, and the approach was to address these in a 'top down' manner, rather than sub-optimize by specializing on anyone component. Most of the 'tools' were achieved to a greater or lesser extent and are presented as a series of publications. However, a central tool, that for monitoring vegetation change, remains outstanding despite comprehensive testing of a range of traditional botanical methods. Indeed, critical review revealed that this 'missing tool' is a problem which is common in all vegetation communities in South Africa - despite the impression created by vegetation researchers that adequate techniques are indeed available. This is serious because land managers are not able to evaluate the impact of their efforts and the government is unable to monitor the effectiveness of their research and extension services, costing millions of public monies annually. The implication also, is that vegetation cannot be managed scientifically (management implies monitoring). Either formal adaptive management is not practicable, or researchers are operating from an inappropriate paradigm; specifically that of providing techniques for their research projects and claiming that these (or derivatives of these) are adequate for farm or regional scale monitoring. More generally, research has often become an end in it's self, with research quality being judged by criteria which are of little significance to the real world and which damage efficiency. Perhaps, the real value of vegetation research lies in the experiential learning which the researcher gains not the inevitably parochial results.
The influence of fertiliser nitrogen on soil nitrogen and on the herbage of a grazed kikuyu pasture in Natal.
(1994) Hefer, Graham Daniel.; Tainton, Neil M.; Miles, Neil.
The work reported in this thesis was designed to develop a better understanding of the fate of fertiliser nitrogen applied to a tropical pasture under field conditions, with the eventual objective of improving the economy of livestock production off such pastures. This involved an examination of the concentrations of soil total nitrogen, ammonium nitrogen and nitrate nitrogen at different depths within the soil profile following the application of different levels of fertiliser nitrogen to a grazed kikuyu (Pennisetum clandestinum) pasture, as well as the influence of such applications on pasture yield and some elements of pasture quality. The trial was conducted over a two year period at Broadacres in the Natal Mistbelt. A labelled [15]NH[4]N0[3] fertiliser experiment was also conducted to ascertain how the labelled ammonium ion moved through the soil, roots and herbage after being applied in spring onto a kikuyu pasture. In the absence of fertiliser N, a total of 15.45 t/ha of soil N was recorded at an average concentration of 0.15%. More than 30% of the soil total N was, however, situated within the top 10cm of soil. organic matter (OM) content in the top 0-10cm of the profile was high (4.75%), reflecting an accumulation of organic matter in this zone. However, as organic C (and thus c: N ratios) declined with depth, so too did soil total N concentration. Not surprisingly, fertiliser measurably increase soil total N, N applications did not but indirectly may have affected soil N dynamics by increasing net mineralisation (due to its "priming" effect) thereby stimulating plant growth and thus increasing the size of the organic N pool through greater plant decay. Total soil N concentration did not change significantly from the first to the second season. This could be attributed to the fact that N gains and losses on the pastures, being over 15 years old, were probably in equilibrium. Generally similar trends in soil total N down the profile over both seasons was further confirmation of this. Before the application of any fertiliser, 331.9 kg NH[4]-N was measured in the soil to a depth of 1m, on average, over both seasons. This amount represented only 2.1% of the soil total N in the profile. The concentration of NH[4]-N followed a quadratic trend down the soil profile, irrespective of the amount of fertiliser N applied, with the largest concentrations accumulating, on average, in the 0-10cm and 75-100cm depth classes and lowest concentrations in the 50-75cm depth class. Laboratory wetting/drying experiments on soil samples collected from a depth of 75-100cm showed that NH[4]-N concentrations declined only marginally from their original concentrations. A high organic C content of 1.44% at this depth was also probable evidence of nitrification inhibition. Analysis of a similar Inanda soil form under a maize crop did not exhibit the properties eluded to above, suggesting that annual turn-over of the soil was causing mineralisation-immobilisation reactions to proceed normally. Addition of fertiliser N to the pasture significantly increased the amount of NH[4]-N over that of the control camps. Furthermore, the higher the application rate, the greater the increase in NH[4]-N accumulation within the soil profile. As N application rates increased, so the NH[4]-N:N0[3]-N ratio narrowed in the soil complex. This was probably due to NH[4]-N being applied ln excess of plant requirements at the high N application rates. On average, 66.7 kg more NH[4]-N was present in the soil in the first season than in the second after fertilisation. Although this amount did not differ significantly from spring through to autumn, during early spring and late summer/autumn concentrations were higher than in mid-summer. Observed soil NH4-N trends were also very similar to the soil total N trends within both seasons, suggesting that soil total N concentrations might well play an important role in determining soil NH4-N concentrations. Before fertilisation, only 45.6 kg N0[3]-N, representing 0.29% of the soil total N, was on average, found in the profile to a depth of 1m. The highest concentration of N0[3]-N was lodged in the top 10cm of the soil. Nitrate-N declined, on average, with depth down the profile. However, during the second season, even though the concentration of N03-N declined down the profile, it increased with depth during relative to that of the first season, suggesting the movement of N0[3]-N down the profile during this period. Fertilisation significantly increased the concentration of N0[3]-N above that of the control camps. Concentrations increased as fertiliser application rates increased, as did N0[3]-N concentrations with depth. This has important implications regarding potential leaching of N03-N into the groundwater, suggesting that once applications reach levels of 300 kg N/ha/season or more, applications should become smaller and more frequent over the season in order to remove this pollution potential. On average, 94.3 kg N0[3]-N/ha was present down to a depth of 1m over both seasons. However, significantly more N0[3]-N was present in the second season than in the first. This result is in contrast to that of the NH[4]-N, wherein lower concentrations were found in the second season than in the first. No specific trends in N0[3]-N concentration were observed within each season. Rather, N0[3]-N concentrations tended to vary inconsistently at each sampling period. Nitrate N and ammonium N concentrations within each month followed a near mirror image. A DM yield of 12.7 t/ha, averaged over all treatments, was measured over the two seasons. A progressive increase in DM yield was obtained with successive increments of N fertiliser. The response of the kikuyu to the N applied did, however, decline as N applications increased. A higher yield of 1.8 t DM/ha in the first season over that of the second was difficult to explain since rainfall amount and distribution was similar over both seasons. On average, 2.84% protein N was measured in the herbage over both seasons. In general, protein N concentrations increased as N application rates increased. On average, higher concentrations of protein-N were measured within the upper (>5cm) than in the lower <5cm) herbage stratum, irrespective of the amount of N applied. Similar bi-modal trends over time in protein-N concentration were measured for all N treatments and within both herbage strata over both seasons, with concentrations tending to be highest during early summer (Dec), and in early autumn (Feb), and lowest during spring (Oct), mid-summer (Jan) and autumn (March). spring and autumn peaks seemed to correspond with periods of slower growth, whilst low mid-summer concentrations coincided with periods of high DM yields and TNC concentrations. The range of N0[3]-N observed in the DM on the Broadacres trial was 0.12% to 0.43%. As applications of fertiliser N to the pasture increased, N0[3]-N concentrations within the herbage increased in a near-linear fashion. On average, higher concentrations of N0[3]-N, irrespective of the amount of fertiliser N applied, were measured wi thin the upper (>5cm) than the lower <5cm) herbage stratum. A similar bi-modal trend to that measured with protein-N concentrations was observed in both seasons for N0[3]-N in the herbage. High concentrations of N0[3]-N were measured during spring (Nov) and autumn (Feb), and lower concentrations in midsummer (Dec & Jan), very early spring (Oct) and early autumn (March). During summer, declining N0[3]-N concentrations were associated with a corresponding increase in herbage DM yields. A lack of any distinctive trend emerged on these trials in the response of TNC to increased fertilisation with N suggests that, in kikuyu, applied N alone would not materially alter TNC concentrations. Higher concentrations of TNC were determined in the lower <5cm) height stratum, on average, than in the corresponding upper (>5cm) stratum. This may be ascribed to the fact that TNCs tend to be found in higher concentrations where plant protein-N and N0[3]-N concentrations are low. A P concentration of 0.248% before N fertilisation, is such that it should preclude any necessity for P supplementation, at least to beef animals. Herbage P concentrations did, however, drop as N fertiliser application rates were increased on the pasture, but were still high enough to preclude supplementation. Even though no significant difference in P concentration was measured between the two herbage strata, a higher P content prevailed within the lower <5cm) herbage stratum. On average, 2.96% K was present within the herbage material in this trial. The norm for pastures ranges between 0.7 and 4.0%. On these trials, applications of fertiliser N to the camps did not significantly affect K concentrations within the herbage. The lower <5cm) herbage stratum, comprising most of the older herbage fraction, was found to contain the highest K concentration in the pasture. The presence of significantly (although probably biologically non-significantly) less K within the herbage in the second season than in the first may be linked to depletion of reserves of · this element in the soil by the plant and/ or elemental interactions between K and other macro-nutrients. An average Ca content of 0.35% within the herbage falls within the range of 0.14 to 1.5% specified by the NRC (1976) as being adequate for all except high-producing dairy animals. Increasing N application rates to the pasture increased the Ca content within the herbage . No significant differences in Ca concentration were found between the upper (>5cm) and lower <5cm) herbage strata over both seasons, even though the lower stratum had a slightly higher Ca concentration, on average, than the upper stratum. Calcium concentrations did not vary between seasons, probably because concentrations tend rather to vary according to stage of plant maturity, season or soil condition. However, higher concentrations of the element were measured in the second season than in the first. The reason for this is unknown. On average, 0.377% Mg was present within the herbage over both seasons. This compares favourably with published data wherein Mg concentrations varied from 0 . 04 to 0.9% in the DM, with a mean of 0.36%. All camps with N applied to them contained significantly more Mg in their herbage than did the material of the control camps. On these trials, the Ca :Mg ratio is 0.92: 1, which 1S considered to be near the optimum for livestock and thus the potential for tetany to arise is minimal. Magnesium concentrations remained essentially similar within both herbage strata, regardless of the rate of fertiliser N applied. As in the case of Ca, Mg concentrations within the herbage were significantly higher in the second season than in the first. Calcium:phosphate ratios increased, on average in the herbage, as N application rates increased. This ratio was high in spring, dropped off in summer and increased again into autumn, suggesting that the two ions were following the growth pattern of the kikuyu over the season. The K/Mg+Ca ratios were nearly double that of the norm, suggesting that the pasture was experiencing luxury K uptake which may be conducive to tetany in animals grazing the pasture. This ratio narrowed as N application rates were increased, probably as a result of ion dilution as the herbage yields increased in response to these N applications. The ratio was low in spring (October), but increased to a peak in December, before declining again to a low in March.

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