A dissertation submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of requirements for the degree of Master of Science. School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Johannesburg, South Africa. 3 June 2016. / Currently Eucalyptus plantations in the warm and cool temperate parts of South Africa are being exposed to damaging temperature extremes and unseasonal frost events that, in particular, have detrimental effects on juvenile plantations. To accommodate these conditions, E. grandis and E. nitens have been selected for hybridization in efforts to identify and select clones suitable for successful plantation establishment in affected areas. Biochemical and physiological responses of plants to cold shock and simulated frost conditions offers a means for this type of selection. In this study, the responses of E. grandis, E. nitens and 8 characterized E. grandis x E. nitens (GN) hybrid clones to cold shock and simulated frost conditions were evaluated. The responses elicited were used as an indication of the eucalypts low temperature and frost tolerance potential, based on levels of: reactive oxygen species (ROS), phenolic acids (PA), starch, total soluble sugars (TSS), chlorophyll fluorescence (CF) and relative electrolyte conductance (REC). Plants were subjected to standard growth conditions of 25°C day/14°C night temperature and a 12h photoperiod for 7 days and subsequently cold shocked at 5°C for 24h. Frost conditions were simulated by freezing excised leaf discs from 2°C to -6°C at a rate of -4°C/h with a one hour hold at -6°C. The results showed an upregulation of ROS in E. grandis, GN 1, GN 4 and GN 6, 30-90 minutes into the cold shock; and levels were highest in E. nitens, GN 3 and GN 7 only 24h after the cold shock exposure. PA levels changed marginally under cold shock conditions, with levels of GN 4 increasing the most by 58%. Starch levels of GN 6 were the most affected by the cold shock, where a 33% increase in levels was recorded. TSS levels of E. grandis and GN 6 increased by 201% and 409% respectively, while TSS levels of GN 2 and GN 3 decreased by 41% and 76% respectively. CF levels of E. nitens and two GNs were most affected by the cold shock, however, all the eucalypts tested, except GN 2, GN 3 and GN 6, displayed a high recovery potential to the cold shock. REC levels fluctuated slightly between unfrozen and frozen samples under standard and cold shock conditions and it was found that E. grandis, GN 1 and GN 3 were the least frost tolerant; and GN 4, GN 7 and GN 8 were the most frost tolerant according to REC levels under cold shock and simulated frost conditions. The results indicate that of all the tested eucalypts, only three GNs were not tolerant to the cold shock and E. grandis and two GNs were not tolerant to the simulated frost. Therefore, it was concluded that all of the eucalypts investigated, apart from E. grandis, GN 1 and GN 3, may be suitable for plantation establishment in areas prone to frost in South Africa. / GR2016

The environmental factors affecting wood property formation of Eucalyptus spp. trees in two distinct geographic areas within South Africa were studied. Wood prop~rty data for trees from 43 sites (26 in Zululand and 17 in Mpumalanga) were collected from work conducted at the Forest and Forest Products Research Centre (FFPRC) at the CSIR (Council for Scientific and Industrial Research) in Durban. The wood properties considered included screened pulp yield, fibre length, wood density and active alkali chemical consumption during pulping. The effect of environmental factors on growth rate (expressed as Site Index at a base age of 5 years) was also measured. A detailed site description for each forest compartment was carried out. Detailed annual and monthly rainfall and minimum and maximum temperature estimates were calculated for each site by interpolating long term means of these variables by splining using the software package Anusplin. These estimates of climatic factors were validated by comparison to the data published in the South African Atlas of Agrohydrology and -Climatology. The outputs of the Anusplin model were used to derive surrogate bioclimatic parameters for each site using the computer program Bioclim. These parameters are considered as better descriptors of the energy-water balance experienced by the plant than normal measures of climate such as mean monthly or annual precipitation. Soil characteristics were measured on samples taken from the individual sites. The effects of these environmental and bioclimatic variables on wood properties were analysed using appropriate statistical techniques. Multiple regression models were used to predict wood properties and it is suggested that this approach could form part of a fibre management system. Wood property prediction models incorporating climate (and bioclimate) alone were preferred to those including soil data as no further site data are required. The effect of edaphic factors was considered to describe any further variation not accounted for by bioclimate alone. Particle size distribution of the soil, as an indication of the water holding capacity of that soil, was not found to effect wood properties or growth significantly. A weak influence of organic matter content in the topsoil on wood density was noted in Mpumalanga. In Zululand, a multiple linear regression using both rainfall of the wettest quarter and mean diurnal temperature range as inputs yielded the best predictive model for growth rate. In this region a combination of precipitation seasonality and mean diurnal temperature range gave the best linear regressi'o,n model describing variation in screened pUlp' yield and fibre length. In Mpumalanga effective rooting depth was found to have a pervasive effect on plant development. Solar radiation (as a measure of energy supply), calculated from a function of latitude, aspect, slope and time of year, was also found to significantly affect the growth rate and SPY of plant material in Mpumalanga. Measures of temperature in both geographic regions were found to significantly affect wood density. / Thesis (M.Sc.)-University of Natal, Durban, 2003.

Caption: “Stump of Blue Gum Tree at Burgersdorp. 1965. The plate bears the inscription: Blougomboom 90 Jaar oud. Grootste boom in N.O. Kaapland. Onder hierdie boom waterskema geopen 1898. Doodvonnis oor Burger P. Klopper uitgespreek in Anglo Boere oorlog. Gesamentlike Diens gehou met vorming van Unie van Suid Afrika. Gedenkplaat onthul met eeufees ossewatrek op Burgersdorp, 12 Oktober 1938.”

Thesis (MScFor)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: The study aims at developing models for predicting aboveground biomass for selected drought tolerant Eucalyptus (E) species (E. cladocalyx, E. gomphocephala and E. grandis x camaldulensis) from the dry west coast. Biomass models were fit for each of the species and a cross-species model was parameterised based on pooled data for all the three species. Data was based on destructive sampling of 28 eucalypt trees which were 20 years of age and additional five five-year old E. gomphocephala trees. Preliminary measurements on diameter at breast height (dbh), height (h) and crown height were recorded in the field. The sampled trees were then felled and samples of discs, branches and foliage were collected. Density of the wood discs and the bark was determined by a water displacement method and computer tomography scanning (CT-scanner). Stem biomass was reconstructed using Smalian’s formula for volume determination and the calculated densities. Upscaling of the crown was carried out by regression equations formulated by employing the sampled branches. Further assessment was carried out on a sub-sample by subjecting the samples to different drying temperatures in a series between 60 and 105ºC. Linear models were parameterised by a simultaneous regression approach based on Seemingly Unrelated Regression (SUR) using the “Systemfit” R statistical package. The predictor variables employed in the study were dbh, d2h and h in which the coefficient of determination (R2), Mean Standard Error (MSE) and Root Mean Standard Error (RMSE) were used to determine the goodness of fit for the models. Akaike Information Criteria (AIC) was also used in the selection of the best fitting model. A system of equations consisting of five models was formulated for each Eucalyptus species. The biomass prediction models had degrees of determination (R2) ranging from 0.65 to 0.98 in which dbh and d2h were the main predictor variable while h improved the model fit. The total biomass models were the best fitting models in most cases while foliage biomass had the least good fit when compared to other models. When the samples were subjected to different drying temperatures, stem wood had the largest percentage change of 6% when drying from 60ºC to 105ºC while foliage had the lowest percentage change of less than 2%. / AFRIKAANSE OPSOMMING: Die doel met hierdie studie is om modelle vir die voorspelling van die bogrondse biomassa van drie droogte-bestande Eucalyptus (E) spesies (E. cladocalyx, E. gomphocephala en E. grandis x camaldulensis), gekweek op die droë kusvlakte in Wes-Kaapland, te ontwikkel. Biomassa modelle vir elk van die spesies is gepas en ’n model gegrond op die gekombineerde data van al drie die spesies, is geparameteriseer. Verder is die biomassa variasie onder verskeie droogingstemperature vasgestel. Die data versameling is uitgevoer gegrond op die destruktiewe mostering van 28 Eucalyptus bome wat 20 jaar oud was en ’n bykomende vyf vyfjarige E. gomphocephala bome. Die aanvanklike mates, naamlik deursnee op borshoogte (dbh), boomhoogte (h) en kroonhoogte is in die veld opgemeet. Die gemonsterde bome is afgesaag en monsters van stamhout skywe, takke en die bas is versamel. Die digtheid van die skywe en die bas is deur die waterverplasing metode, en Rekenaar Tomografie skandering (“CT-scanning”) vasgestel. Stam biomassa is rekonstrukteer deur gebruik te maak van Smalian se formule vir die vasstelling van volume en berekende digtheid. Die opskaal van die kroon biomassa is gedoen met behulp van regressie vergelykings van gekose takmonsters. Submonsters is onderwerp aan ’n reeks van verskillende drogingstemperature tussen 60 en 105ºC. Lineêre modelle is deur ’n gelyktydige regressie benadering gegrond op die Seemingly Unrelated Regression (SUR) wat ’n“Systemfit” R statistiese pakket gebruik, parameteriseer. Die voorspeller veranderlikes wat in hierdie studie gebruik is, is dbh, d2h en h waarin die koëffisient van bepaling (R2), gemiddelde standaardfout (MSE) en vierkantswortel van die gemiddelde standaardfout (RMSE) gebruik is om vas te stel hoe goed die model pas. Akaike Inligting Kriteria is gebruik vir die seleksie van die gepaste model. ’n Reeks vergelykings wat bestaan uit vyf modelle is vir elke Eucalyptus spesie geformuleer. Die biomassa voorspelling model het waardes vir die koëffisiente van bepaling (R2) opgelewer wat strek van 0.65 to 0.98% en waarin dbh en d2h die hoof voorspelling veranderlikes is, terwyl h die pas van die model verbeter. Die totale biomassa model het in die meeste gevalle die beste gepas en die blaarbiomassa die swakste as dit met die ander modelle vergelyk word. Tydens droging vind die grootste persentasie verandering van 6% by stamhout plaas tussen temperature van 60ºC tot 105ºC, en die kleinste persentasie verandering van minder as 2% by blare.

Thesis (PhD(For))--Stellenbosch University, 2012. / The subtropical ecosystem of the Zululand coastal plain is prized by the South African commercial plantation forestry industry for its rapid clonal Eucalyptus growth, short rotations (6 to 7 years) and high yields. This region is typified by sandy soils that are low in clay and organic matter, have small nutrient reserves and are poorly buffered against nutrient loss. The subtropical climate induces rapid decomposition of residues and tree litter resulting in small litter nutrient pools and rapid nutrient release into the soil, particularly after clearfelling. A combination of large nutrient demands through rapid growth, rapid nutrient turnover and small soil nutrient reserves implies that sites in this region are sensitive and may be at risk of nutrient decline under intensive management. The work in this study set out to determine the risk of nutrient depletion through harvesting and residue management on a site within the Zululand region, to assess nutritional sustainability and the risk of yield decline in successive rotations. Some bulk biogeochemical cycling processes of macro-nutrients nitrogen (N), phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) were assessed, and assessments also included sodium (Na). An existing Eucalyptus stand was clearfelled and treatments were imposed on the residues after broadcasting to simulate various levels of nutrient loss through levels of harvesting intensity and residue management. These included residue burning (Burn), residue retention (No-Burn), fertilisation (stem wood nutrient replacement), whole tree harvesting and residue doubling. Outer blocks of the stand were not felled, but included as replicates of an undisturbed standing crop treatment. Biogeochemical nutrient cycling processes were assessed primarily in the standing crop, Burn and No-Burn treatments, in the assumption that these represented the furthest extremes of nutrient loss. Data collection commenced a year prior to clearfelling and continued to two years and six months after planting with key data collection over a 20.1 month period from clearfelling to canopy closure (one year after planting). Water related nutrient pools and fluxes were assessed as atmospheric deposition (bulk rainfall, throughfall and stemflow) and gravitational leaching to 1m soil depth. Drainage fluxes were predicted using the Hydrus model and real-time soil moisture data. Zero tension lysimeters collected soil solution for chemical analysis. Sequential coring in the 0 to 30cm soil layer was used to determine in situ soil N mineralisation. Soil chemical and physical properties were assessed over the first meter of soil at clearfelling and new crop canopy closure to determine soil nutrient pools sizes. Biomass nutrient fluxes were assessed from litterfall, residue and litter decomposition, and above ground accretion into the tree biomass. Leaching and N mineralisation were monitored in the No-Burn, Burn and standing crop treatments only. Atmospheric deposition, while variable, was shown to be responsible for large quantities of nutrients added to the Eucalyptus stand. Nitrogen and K additions were relatively high, but within ranges reported in previous studies. Rapid tree canopy expansion and subsequent soil water utilisation in the standing crop permitted little water to drain beyond 1m resulting in small leaching losses despite a sandy well drained soil. Further leaching beyond this depth was unlikely under the conditions during the study period. Mineralisation and immobilisation of N also remained low with net immobilisation occurring. The standing crop was shown to be a relatively stable system that, outside of extreme climatic events, had a relatively balanced or positive nutrient budget (i.e. nutrient inputs minus outputs). Large quantities of nutrients were removed with stem-wood-only harvesting in the No-Burn treatment leaving substantial amounts on the soil surface in the harvest residues. Whole tree removal increased losses of all nutrients resulting in the largest losses of P and base cations compared to all other treatments. This was mostly due to high nutrient concentrations in the removed bark. Loss of N in the Burn treatment exceeded whole tree N losses through combustion of N held in the harvest residues and litter layer. The majority of K leached from the residues prior to burning and a relatively small fraction of the base cations were lost from the partially decomposed residues during burning. Ash containing substantial amounts of Ca and relatively large amounts of N and Mg remained after burning. Surface soil Ca and Mg was significantly increased by the ash which moved into the soil with rainfall directly after burning. Rapid soil moisture recharge occurred within a few months after clearfelling, increasing leaching from the upper 50cm of soil. Clearfelling increased net N mineralisation rates, increasing mobile NO3-N ions in the soil surface layers. Nitrate concentration peaked and K concentration dipped in the upper soil layers of the Burn treatment directly after burning. Deep drainage and leaching (beyond 1m depth) over the 20.1 month period was, however, not significantly different between the Burn and No-Burn treatments. Rapid soil moisture depletion and nutrient uptake with new crop growth reduced leaching fluxes to levels similar to the standing crop by six months after planting. Taking the full rotation into account, clearfelling induced a short-lived spike in N and cation leaching compared with the low leaching losses in the undisturbed standing crop. Soil N mineralisation over the 20.1 month period in the burnt treatment was half that of the No-Burn treatment. Growth and nutrient accumulation was significantly higher in the fertilised treatment than in other treatments up to 2.5 years of age. Growth in the Burn treatment was greatest compared to other treatments during the first few months, but slowed thereafter. No significant growth differences were found between all other treatments from a year to 2.5 years after planting. Early growth was therefore apparently not limited by N supply despite large differences in N mineralisation between Burn and No-Burn. Foliar vector analysis indicated that fertilisation improved growth initially through increased foliar N and P at six months after planting followed by Mg and Ca at one year. The Burn treatment was not nutrient limited. These growth results contrasted with similar international research on sandy tropical sites where growth was reduced after residue removal and increased after residue doubling. The combined nutrients released from pools in the litter layer or ash and soil in addition to atmospheric inputs were sufficient to provide most nutrients required to maintain similar growth rates across all treatments. This demonstrated the importance of residue derived nutrients to early growth nutrient supply. Reduced N mineralisation through a lack of substrate may limit N supply later in the rotation where residue had been removed. Construction of a nutrient budget for the system revealed that high levels of atmospheric inputs have the potential to partially replenish a large proportion N, K, and Ca lost during clearfelling, provided losses are constrained to stemwood removal only. However, loss of Mg that occurred primarily through leaching may not be replaced under the low Mg inputs recorded in this study. Larger nutrient removals (i.e. stemwood plus other plant parts) placed a heavier reliance on the small soil nutrient pools at this site which can limit future productivity. More intense harvesting and residue management practices dramatically increased the risk of nutrient depletion. Losses of specific nutrients depended on a combination of clearfelling biomass removal, residue burning and subsequent leaching. Nitrogen losses due to harvesting and burning were more substantial than those due to leaching. Mg and K losses depended most strongly on the time after clearfelling before re-establishment of the new crop and rainfall patterns, while Ca and P losses depended directly on the amount of biomass removed. Depletion risk was the greatest for Mg and K through rapid leaching, even after stem wood only removal. Deep root uptake and deep drainage with associated cation loss needs to be investigated further to quantify ecosystem losses and recovery of cations displaced beyond 1m. Atmospheric deposition is one of major factors countering nutrient losses. However, atmospheric inputs may not be reliable as these may lessen in future through pollution control legislation and climate change. Changes in growth rate under poor nutrient management practices are small and difficult to detect relative to the large impacts of changing weather patterns (drought), wildfire and pest and disease. This makes it difficult to prove nutrient related growth decline. It may be possible that improvements in genetics, silvicultural technologies and atmospheric inputs may also be masking site decline (in general) and in part explain the lack of evidence of a growth reduction in the region. As the poorly buffered sandy soils on the Zululand Coast are at risk of nutrient depletion under the short rotation, high productivity stands, it may be necessary to stipulate more conservative harvesting and residue management practices. A more conservative stem-wood only harvesting regime is recommended, retaining all residues on site. Residue burning should be avoided if N losses become a concern. The length of the inter-rotation period must be kept short to reduce cation leaching losses. Site nutrient pools need to be monitored and cations may eventually need to be replenished through application of fertilisers or ash residues from pulp mills. Management practices therefore need to be chosen based on the specific high risk nutrients in order to maintain a sustainable nutrient supply to current and future plantation grown Eucalyptus.