Thesis (PhD)--Stellenbosch University, 2015. / ENGLISH ABSTRACT: Surface fires are known to affect trees of different species differently, depending on the capacity of the bark to shield the cambium from heat. Tree bark characteristics differ among species and thus potentially influence the protective ability against cambium damage. The objectives of this study were to compare the protective role of bark against fire for selected indigenous and exotic species in the Western Cape, South Africa, and to investigate post-fire growth impacts following surface fire damage on Pinus radiata. In the first part of the study, trees were felled and billets of 25 cm height harvested from different heights along the trunk. Bark thickness, compass direction, stem diameter at breast height, bark moisture content and relative height of the sample in the stem were tested for their effect on heat insulation capacity of bark. Heating experiments were conducted at 400°C on the fresh billets with intact bark. Time to heat the cambium to lethal 60°C was determined. The second part of the study investigated the role of bark surface topology, bark density and bark chemical composition for its fire resistance. The same size billets were harvested from the lower trunk section of selected species. Surface topology was characterised by means of x-ray based computer tomography, density by moisture saturation method and bark chemical composition by thermo-gravimetrical analyses (TGA). The third part of the study investigated the impact of high intensity surface fires on growth of an 18 year old Pinus radiata plantation which was exposed to a ground fire 5 years prior to the analysis. Tree ring measurements were done on cores obtained by non-destructing coring method and various growth indices, based on yearly basal area increment (iBA) used to quantify growth response to the fire damage. Statistical analysis based on correlation, multi-model inference and multiple regression revealed no significant influence of compass direction and diameter at breast height. Heat resistance was mainly determined by bark thickness and to a lesser degree by moisture content. In several species relative height at the stem modulated the bark thickness effect. Higher up the stem bark of the same bark thickness offered less protection against heat. The results also suggest that in particular bark topology plays a role, while the correlations with bark density and chemical composition could not be secured statistically. A main finding was that fissures in the bark play a significant role. A regression model showed a significant influence of fissure width, fissure frequency and the minimum bark thickness to the cambium, which is a function of fissure depth. The results show that structural bark parameters are a necessary addition to explain heat resistance of bark. Statistical analysis employing one-way Anova and incorporating Tamhane’s T2 Post Hoc test revealed significant growth reductions following high intensity surface fire damage on Pinus radiata in the fire year with the impact being passed on to the following year. The recovery phase extended a two year period. During this time the trees showed increased diameter growth probably due to increased water availability. / AFRIKAANSE OPSOMMING: Dit is bekend dat oppervlakvure in bos-ekostelsels verskillende boomspesies verskillend affekteer, afhangende van die vermoë van bas om die kambium van hitte te beskerm. Baseienskappe verskil tussen spesies en het dus 'n potensiële invloed op die beskermende vermoë teen kambiumskade. Die doelwit van hierdie studie was om die beskermende rol van bas teen vuur te vergelyk tussen inheemse en uitheemse spesies in Wes-Kaapland, Suid Afrika asook om die na-vuur impak op groei te ondersoek met brandskade aan Pinus radiata. In die eerste deel van die studie is bome geoes en stompe van 25 cm lengte van verskillende hoogtes verwyder. Basdikte, kompasrigting, stompdiameter by borshoogte, basvoggehalte en die relatiewe hoogte van die stomp in die stam is getoets vir hul invloed op hitte-isolasiekapasiteit van bas. Verhittingseksperimente is gedoen teen 400 °C op die vars stompe wat steeds bas opgehad het. Die tyd om die bas tot by 'n skadelike 60 °C te verhit is bepaal. Die tweede deel van die studie het die rol van basoppervlaktopologie, basdigtheid, en die bas chemiese samestelling ondersoek in vuurweerstand. Dieselfde grootte stompe is geoes van die laer dele van die stam van uitgesoekte spesies. Oppervlaktopologie is bepaal deur middel van X-straal rekenaartomografie, digtheid deur die versadigingsvoggehaltemetode, en chemiese samestelling deur termo-gravimetriese analise (TGA). Die derde deel van die studie het die impak van hoë intensiteit oppervlakvure op groei van 18-jaar oue Pinus radiata ondersoek. Jaarringmetings is gedoen op inkrementboorsels wat nie-destruktief bekom is en verskeie groei-indekse, gebaseer op jaarlikse basale oppervlak aanwas, is gebruik om die groeireaksie op brandskade te kwantifiseer. Statistiese analise gebaseer op korrelasie, multi-model inferensie, en veelvuldige regressie het gewys dat kompasrigting en deursnee op borshoogte nie 'n beduidende invloed gehad het nie. Hitteweerstand was hoofsaaklik bepaal deur basdikte, en in 'n mindere mate basvoggehalte. By verskeie spesies het die relatiewe hoogte die basdikte-effek gemoduleer. Hoër in die stam het dieselfde dikte bas minder beskerming gebied as bas van laer in die stam. Die resultate impliseer dat basoppervlaktopologie ook 'n rol speel in hitteweerstand terwyl basdigtheid en chemiese samestelling nie 'n statisties beduidende rol gespeel het nie. 'n Belangrike bevinding was dat gleuwe of openinge in die bas 'n beduidende rol speel. 'n Regressiemodel wys dat 'n beduidende invloed deur gleufwydte, gleuffrekwensie en die minimum basdikte na die kambium. Die resultate wys dat strukturele basparameters 'n belangrike bykomende rol speel om hitteweerstand van bas te verduidelik. Die statistiese analise waar eenrigting ANOVA met Tamhane se T2 Post Hoc toets gebruik is toon dat ‘n beduidende groeivermindering teweeg gebring is in die Pinus radiata as gevolg van skade veroorsaak deur hoë intensiteit oppervlakvure waarvan die impak eers in die jaar na die vuur sigbaar was. Die herstelfase het oor twee jaar gestrek. Gedurende hierdie tyd het die bome 'n toename in deursneegroei getoon, waarskynklik as gevolg van verhoogde waterbeskikbaarheid.

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. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2013.