Increases in temperature, altered precipitation patterns, and the occurrence and severity of extreme climatic events have been important characteristics of the climate change observed to date. This has had many and diverse impacts upon the living world, with one recent observation being a global reduction in the net primary production of all terrestrial vegetation. Increases in temperature and the frequency of extreme events are predicted to continue throughout the 21st century, and can be expected to have far reaching effects on global terrestrial ecosystems. Increases in temperature and drought occurrence could fundamentally impact upon the growth rates, species composition and biogeography of forests in many regions of the world, with many studies indicating that this process is already underway. European beech, Fagus sylvatica, is one of Europe’s most widespread and significant broadleaved tree species, forming an important and frequently dominant component of around 17 million hectares of forest. However, the species is also considered to be drought sensitive. Thus, much research interest has focused on eliciting the details of its physiological response to increased water stress, whilst dendroecological studies have attempted to identify sites and regions where reductions in growth might be found. A significant knowledge gap exists regarding a multi-regional, range-wide view of growth trends, growth variability, climate sensitivity, and drought response for the species. Predicting the potential effects of climate change on competition and species composition in mixed species forests remains an important challenge. In order to address this knowledge gap, a multi-regional tree-ring network was constructed comprising of 46 sites in a latitudinal transect across the species’ Western European range. This consisted of 2719 tree cores taken from 1398 individual trees, which were used to construct tree-ring chronologies for each site in the network. As a first step in a multi-regional assessment for F. sylvatica, a combination of the tree-ring chronologies and environmental data derived from a large scale gridded climate dataset were used in a multivariate analysis. Sites in the latitudinal transect were partitioned into geographically meaningful regions for further analysis. The resulting regions were then studied using climate-growth analysis, pointer year analysis of drought years, analysis of growth trends and growth variability, in order to examine regional variation in the response of the species to climate. Furthermore, a combination of long-term monitoring data from one specific site was combined with tree-ring sampling of multiple cohorts of F. sylvatica and one co-dominant competitor, Quercus petraea, to study the effects of an extreme drought event in 1976 on mortality and subsequent recovery. Key results of the multi-regional analysis are that large scale growth reductions are not evident in even the most southerly and driest portions of the species’ range. Radial growth is increasing, both in the north and in the core of the species’ range, with southern range edge forests maintaining stable growth. However, the variability of growth from year to year is increasing for all regions, indicative of growing stress. Crucially, the southern range edge, which previous studies had identified as an ‘at risk’ region, was shown to be more robust than expected. Climate sensitivity and drought impacts were low for this region. Instead, forests in the core of the species range, both in continental Europe and in the south of the UK, were identified as having the highest climate sensitivity, highest drought impacts, and experiencing periodic reductions in growth as a result. Northern range edge forests showed little sign of being affected by drought, instead having low climate sensitivity and strongly increasing growth trends. Extreme drought was found to affect species differently: the dominant species (F. sylvatica) failed to recover pre-drought levels of growth, whilst a transient effect of competitive release occurred for the co-dominant species (Q. petraea). There was also a long term effect on the relative abundance of the two species within the woodland, due to differences in the levels of drought induced mortality experienced by the species. This shows that in the case of extreme climatic events where thresholds in the ability of species to tolerate water stress are breached, the effects of drought can be rapid and long lasting. Drought impacts can cascade beyond that experienced by the most drought sensitive species, due to changes in competitive interactions between species in mixed species forests. The implications of this work suggest opportunities, risks and strengths for F. sylvatica. In the northern portion of the species’ range, predicted increases in productivity are confirmed by recent growth trends, indicating a good outlook for the species. At the southern range edge, F. sylvatica forests exist either in locations where precipitation is high or locations where local environmental conditions buffer them from an inhospitable regional climate. These factors result in southern range edge forests which are highly resilient to the effects of increasing climate stress. It is instead in the core of the species’ range where the most sensitive forests are found. The effects of extreme drought on a range core forest demonstrated here provide a cautionary note: where drought stress tolerance thresholds are breached, rapid and long lasting effects on growth and mortality can occur, even in regions where drought has not previously been considered to pose a strong risk to the species.

Extensive drought - and elephant-related dieback of Colophospermum mopane and Acacia tortitis, respectively, offered an opportunity for increasing understanding of the causes of drought-related patch dieback, the factors influencing elephant utilization of woody plants, and the response of woody plants to both aforementioned determinants of savanna structure and function. The dendrochronological analysis of both species was undertaken to estimate potential rates of replacement, following extensive mortality. Areas of discrete dieback were compared with adjacent paired areas of 'healthy' vegetation, which revealed, on average, 87% and 13% loss of basal area by mortality, respectively. 'Live' and 'dead' plots did not differ in soil type, topography or mean slope, but differed in vegetation structure, soil surface condition, and soil chemistry. Although there was evidence of self-thinning, neither inter - nor intra-specific competition explained dieback. 'Dead', by comparison with 'live' plots, had changed from functioning as sinks of sediment and water to sources, were less likely to retain water because of a poor soil surface condition, and were predisposed to drought effects because of a greater proportion of fines, and Na concentration. Dieback resulted from insufficient soil water for survival during a drought owing to the development of a dysfunctional landscape during 50 years of livestock ranching. Spatial heterogeneity within a landscape was suggested to enhance woodland resilience to severe droughts by ensuring the survival of plants in run-on sinks or 'drought refuge' sites. Stem sections were removed from 40 multi-stemmed C. mopane trees and prepared for examination under a dissecting microscope. It was impossible to age C. mopane, owing to a hollow and/or dark heartwood. Nevertheless, the distribution of stem diameters suggested a single recruitment event. Fire scars attributed to the last recorded fire in 1948 could explain the trees' multi-stemmed growth form and indicate that most trees of VLNR were > 50 years of age. Growth rings were identified in 29 A. tortilis trees of unknown age, but were not correlated with annual rainfall records. Growth rates varied between trees; mean ring width ranged from 1.4 to 3.5 mm (overall mean 2.4 ± 0.1 mm). A technique was proposed for predicting growth rate from annual rainfall, using selected data, and several factors potentially influencing ring width in semiarid environs were identified. Permanent ground-based transects were located within riparian (n = 16) and Acacia (n = 5) woodlands to monitor elephant utilization. Elephant had not changed the population structure of the woodlands by 2000, but had reduced stem density from 215.6 stems ha -1 (1996) to 84.4 stems ha -I (2000). Acacia tortitis trees in the woodlands had branches removed, were debarked, uprooted and broken. Acacia tortitis trees in the riverine had lower levels of utilization, whilst Acacia nilotica trees were mostly debarked. The method of elephant feeding varies within and between woody species, provided it is within the mechanical constraints of a certain size or species. Elephant behaviour is concluded to depend on spatiotemporal variation of forage abundance/quality, abundance of a preferred species, and species response (coppice or mortality). Elephant can cause a change of vegetation state, and increase spatial homogeneity of a plant population. The remnant population of woodland trees should provide the potential for recolonization, in which case the system would reflect the stable limit cycle. However, if browsing inhibits seedling recruitment, the system could reflect either a multiple stable state system or an artificial equilibrium imposed on a stable limit cycle. / Thesis (M.Sc.)-University of Natal, Pietermaritzburg, 2000.