Tesis sobre el tema "Eucalyptus – Western Australia"

The WA coolibah tree, Eucalyptus victrix L. Johnson & K. Hill forms an unique and pristine woodland in the Fortescue Valley, in the Pilbara district of Western Australia. Until recently, no research had been done on E. victrix ecology and concern had been expressed by pastoral managers and others about the condition of the woodland occupied by this species. This research was an attempt to understand the ecology of coolibah using a multi-disciplinary approach.A population demographic study of stands in E. victrix woodland reveals that the present tree populations occupy larger areas than saplings and seedlings. Soil moisture and warm summer temperatures are believed to be stimulating factors for seedling recruitment of E. victrix. Different size classes (height/diameter) reflect different recruitment events and these reflect past availability of seeds and adequate soil moisture in that particular area of the woodland. Occasional grasshopper outbreaks and drought cause canopy shrinkage. Presence of hollow boles, restrict dendrochronological examination of tree ages.An important population study was of a post-seedling cohort at Roy Hill, where height distribution reflected a typical single recruitment event. In subsequent years (1995 - 1998) height measurement showed several peaks, suggesting that seedlings were now growing at different rates. Uniform and steady height growth was observed on saplings found at the edges of gilgai. During May 1998 several saplings flowered, it was assumed that E. victrix attained its reproductive maturity at mean height of 2.50 m and with a stem diameter of 50 mm.Seedling recruitment and subsequent growth mainly depend on heavy rainfall flooding events. Seedling recruitment occurs only from current seed rain. Seed longevity reduces after 54 days of burial at 50 mm depth. Mortality (4 - 6 leaf stage) of newly recruited seedlings during ++
subsequent dry months is very high. Furthermore, allelopathic effects (root competition from established grass and insect herbivory) are additional causes of seedling mortality in the years of recruitment.Seedlings recruited at a burnt (disturbed) site, grew faster compared with undisturbed sites with less mortality. This suggested that fire created a suitable condition by reducing root competition, increasing soil nutrients and also creating a gap which providing sufficient solar radiation for seedlings to establish and develop into a healthy population. It is suggested that once newly recruited seedlings overcome the first summer, mortality rates during subsequent years are drastically reduced.Long-term waterlogging of E. victrix seedlings significantly increases seedling stem diameter. Large numbers of adventitious roots are developed and lenticels proliferate on the submerged portion of the stem. Flooding reduces each photosynthesis, transpiration and stomatal conductance. Flooding does not increase shoot fresh or dry weight on 4-, 8- or 17- week old seedlings. Leaf emergence may be stimulated on flooded seedlings compared with unflooded seedlings. While root dry weight is greater in 17-week old flooded plants than 13 - week seedlings, this difference is not significant. It is suggested that maintenance of a high root/shoot ratio is a drought adaptation. Furthermore, comparative study of flood tolerance in semi-arid eucalypt species suggests that those species intolerant of flooding seldom express morphological adaptations and fail to recover from physiological damage.The annual grass Setaria dielsii occurs under the canopy of mature E.victrix trees of the coolibah woodland. This species has probably displaced more palatable perennial grasses. Soil moisture under trees is slightly higher and soil temperatures are less extreme than away from the canopy. Growth of ++
S. dielsii appears to be more associated with soil nutrient status. Levels of total N, Mg, K, and of electrical conductivity (EC) under trees are significantly higher than those away from the tree. Levels of Ca, Na, Fe, and organic carbon do not differ. The under story sub-shrub Malvastrum americanum is an important competitor with S. dielsii. Light availability may determine relative biomass contributions of the two species.The effect of the density of grass species, growing space and time of establishment on E. victrix seedlings (inter-specific competition), and the effect of density and growing space within E. victrix seedling populations (intra-specific competition) were studied under controlled conditions. Results indicate resources necessary for growth of individual E. victrix seedlings were more limiting under increase density of neighbouring grass species than under intra-specific competition. In particular photosynthetic area was drastically reduced in mixed culture.Lack of ground cover beneath the canopy of isolated E. victrix trees was ascribed to toxic or phenolic substances present in leaf, bark and leaf litter of E. victrix. Lactuca sativa seed germination subjected to extracts and leachate demonstrate that the fresh leaf of E. victrix possesses toxic substances which cause deleterious effects on both germination and radicle development. Similarly, increasing concentrations of leaf and bark leachate show reduced germination percentage of L. sativa seeds. Shoot and root biomass of grass and eucalyptus treated with leaf leachate were reduced. E. victrix leaf leachate significantly reduced shoot and root biomass of its own seedlings. High Performance Liquid Chromatogram (HPLC) analysis separated 11 and 8 possible toxic substances from leaf and bark extract respectively.

Plantation eucalypts are a recent and rapidly growing industry in Australia, and will eventually replace the logging of old-growth forests. Over 40% of these plantations have been established in south-western Australia, where more than 160 000 ha of Eucalyptus globulus plantations now occur. In the early 1900s, this species was widely planted as an exotic in South Africa, but succumbed to severe pest (Gonipterus sp.) and disease (Mycosphaerella sp.) problems. Similarly, in south-western Australia E. globulus is an exotic species, but with the additional threat that it is planted adjacent to indigenous eucalypts, which increases the possibility of pests and pathogens switching between closely related eucalypt hosts. Over the past ten years, there have been anecdotal reports of increasing levels of Mycosphaerella leaf disease (MLD) in E. globulus plantations in south-western Australia. This increase in disease level is of concern to the industry. To date there have been no comprehensive studies into the taxonomy, biogeography and population genetics of MLD in south-western Australia. This thesis investigated the impact of MLD in south-western Australia with a focus on its impact, taxonomy, biogeography and population genetics. It is the first study worldwide to quantify the relative impact of different Mycosphaerella species in a regional plantation estate. A survey of pest, disease and nutritional disorders (Chapter 2) found that MLD was the most severe and frequently occurring, single taxonomic health threat to 1 and 2-year-old E. globulus plantations in south-western Australia. For the first time, this survey identified and quantified the impact of pest and disease damage to E. globulus plantations in the region. There were differences in the disease levels between plantations and this was due to initial Mycosphaerella species composition and inoculum level, and local climatic conditions favourable for disease, rather than to the provenance planted or the nutritional status of the individual plantations. The survey for Mycosphaerella pathogens of eucalypts (Chapter 3) identified two new species of Mycosphaerella (M. ambiphylla and M. aurantia) and extended the known geographic range of eight other species (M. cryptica, M. gregaria, M. lateralis, M. marksii, M. mexicana , M. nubilosa, M. parva and M. suberosa). Of these: M. lateralis and M. mexicana were new records for Australia; and M. gregaria, M. nubilosa and M. parva were new records for Western Australia. A new anamorph, Phaeophloeospora ambiphylla was described and linked to M. ambiphylla. The occurrence of these new species and disease records in south-western Australia is significant for the plantation-eucalypt industry worldwide. The finding of two new species highlighted the need to quantify the disease impact of these on eucalypt plantations; and the extension of the range the remaining species raised important quarantine issues, concerned with the movement of plant material between regions and countries. The biogeographical investigation of Mycosphaerella (Chapter 4) identified that the most widespread and serious cause of MLD in south-western Australia is M. cryptica. In addition to occurring on the exotic E. globulus, it also occurs on two of the three important indigenous forestry eucalyptus species in this region. That is, on E. diversicolor, and E. marginata, but not on Corymbia calophylla. In terms of the plantation estate of E. globulus, however, M. nubilosa is the most widespread pathogen. The current study found that MLD on E. globulus is a complex of several different species, whereas, on E. diversicolor and E. marginata it is caused by only M. cryptica. Two species, M. cryptica and M. marksii were found commonly on adult E. globulus leaves. Although M. cryptica was the most frequent and serious cause of disease on adult leaves, M. marksii levels appear to be increasing and the future epidemiology of this pathogen should be closely monitored. There is some concern that these two MLD species could become an economically important problem on adult leaves of E. globulus. At present severe levels of MLD is significantly more common on juvenile than on adult foliage. The phylogenetic analysis (Chapter 5), based on ITS rDNA sequences from the present study and those obtained from GenBank accessions, found that Mycosphaerella is an assemblage of largely polyphyletic anamorph genera. Ten distinct clades emerged from the analysis. With the exception of the Dissoconium and the M. recutita clade, which comprised of two and one species respectively, none were comprised entirely of one anamorph genus alone. The anamorph genera represented were often dispersed across more than one clade, indicating that these anamorphs have arisen separately in different phylogenetic lineages. Cercospora, Stenella and Uwebraunia anamorphs each occurred in more than one separate clade. Although on the basis of rDNA sequence data Mycosphaerella appeared mostly monophyletic there was some evidence that the Mycosphaerella genus may be polyphyletic. This was particularly evident from the Dissoconium clade which grouped as closely to the outgroup Botryosphaeria taxon as it did with the remaining Mycosphaerella species. It was argued that a multi-gene phylogeny, which includes sequencing many species in other genera aligned with Mycosphaerella, is required in order to satisfactorily answer the question of whether Mycosphaerella is truly monophyletic. The phylogenetic analysis also showed that the taxonomy of Mycosphaerella based on ITS sequence data needs further clarification. Some species that are morphologically distinct, such as M. vespa and M. molleriana, shared identical ITS sequences. Other morphologically distinct species differed by as little as one or two nucleotides. Yet in other cases, the sequence variation amongst isolates from the same species differed substantially. Much of this variation in M. cryptica and other species was attributed to poorly edited sequences that had been lodged with GenBank. It was postulated that although a part of the remaining variation reflected the existence of cryptic species, some was likely to be genuine intra-species differences. It was concluded that further genes need to be sequenced, and more standardised cultural studies conducted in order to define species boundaries within Mycosphaerella. Based on the ITS rDNA sequence data, two different molecular methods for the identification of Mycosphaerella species from eucalypts were developed (Chapter 6). The first of these was a PCR-RFLP method that enabled the identification of Mycosphaerella species present on eucalypts in south-western Australia. A key is provided, which enabled the identification of species on a combination of PCR-RFLP DNA fragment migration patterns and a small number of morphological features. This key enables the identification of Mycosphaerella species more easily than keys that rely on morphological features alone. Therefore, this has made it easier for non-Mycosphaerella specialists to identify species from this genus. The second molecular method developed for the identification of Mycosphaerella species was that of primers that selectively amplify the DNA of M. cryptica and M. nubilosa, the two most important causes of MLD (Chapter 6). This will allow the rapid identification of these two species by non-specialists in Mycosphaerella taxonomy. The primers from the current study will also enable early diagnosis of the possible causal organism of MLD in a plantation. Once the use of these primers for amplifying DNA from leaf tissue has been optimised, they will also facilitate studies into the early infection process of M cryptica and M. nubilosa. For example, the presence of the pathogen may be detected prior to the appearance of symptoms. Studies may be conducted to determine the length of a hemi-biotrophic phase, and the extent of tissue colonisation both spatially and temporally, beyond the necrotic lesion in these two Mycosphaerella species. Previously, such studies have been hampered by the slow growth rate of these fungi in culture and the lack of media that would allow their selective isolation and detection by directly plating diseased and non-diseased host tissue. This study has clearly indicated that Mycosphaerella species are the major disease threat to E. globulus plantations in Western Australia. It has also shown that over the relatively short period of time of less than ten years the number of species recorded has increased from three to ten, and that disease severity has increased in plantations. It is therefore critical to continue the research on this genus in order to understand the biology, epidemiology and population genetics of this pathogen. This is necessary in order to inform tree selection and silvicultural practise that will minimise the future impact of MLD. This is particularly important if the industry moves towards clonal and hybrid forestry as has occurred elsewhere in the world. This study has laid the foundations for future research on this disease through the elucidation of the taxonomy of Mycosphaerella in south-western Australia and by providing some important molecular tools for its diagnosis and further study.

[Truncated abstract] Removal of native vegetation to facilitate traditional agriculture practices has been shown to reduce ecosystem health, and restricts the native habitat. The subsequent change in the predominant vegetation water use patterns has altered the catchment water balance, and hydrology which results in land degradation through such processes of salinisation and water logging. More recently, moves toward more sustainable farming practices have been taken to help re-establish catchment hydrological equilibrium and improve catchment ecosystem services. Agroforestry is one such vehicle for this reestablishment. Perennial native vegetation has been shown to have a significant effect on catchment processes, mitigating any further degradation of the land. The effect of alternating native perennial tree belts with traditional broad acre agriculture in the alleys, referred to as alley farming, is investigated in this thesis due to the potential environmental and economic benefits that can result. This thesis investigates the impact of tree belts upon the water table and aims to gauge the ability of alley farming at controlling recharge within the low-medium rainfall zone on the valley floor. The basis of this research is the analysis of data collected from the Toolibin Alley Faring Trial. This experiment was established in 1995 to assess the viability of alley farming and incorporates different combinations of belt width, alley width and revegetation density. Transects of piezometers within each design have been monitored from October 1995 to January 2008. The piezometers were sporadically monitored over this period on a total of 39 dates. ... To further understand the response observed in the water table data, in depth hydrograph analysis of the control piezometer water levels was conducted. The statistical analysis demonstrates that the belts are having a very limited impact on the water table morphology, this is associated with the restricted use of groundwater by the perennial tree belts due to the poor quality, has been applied. This explains why there is limited signature of increased water table depth in the statistical analysis; there is evidence that alley farming as a means of reducing recharge may work however the overriding control on the trial are the rainfall trends rather than perennial growth. The low perennial biomass production at the site is an effect of limited water resources; however a significant distinction can be made between the water table depth and variability beneath high and low biomass belts. There are three main controls at the site; climate, development of perennial biomass and development of perennial root systems (both vertically and laterally). The regional climatic trends will influence water table levels creating a greater soil water storage capacity; therefore the contribution of soil water to transpiration rates will enable the tree belts to have some impact on recharge. Of the alley farming designs tested, the optimal planting density and belt/alley design, from an economic perspective, is identified as having a 4m belt width which generated the greatest biomass. As a means of controlling recharge at the site the effectiveness of alley farming is limited due the shallow saline water table limiting perennial growth.

Thesis (Ph.D.)--Murdoch University, 2003.
Thesis submitted to the Division of Science and Engineering. "Supported by CRC for Sustainable Tourism and Dept of Conservation and Land Management (Western Australia)". Bibliography: leaves 334-355.

[Truncated abstract] In order to assess the sustainability of mixed plantings on saline land, this thesis examined the importance of leaf-litter trapping and microsites on recruitment in a salt affected alley-belted (tree/shrub) agricultural system in Western Australia. Located in the low rainfall region (MAR <330 mm) of the wheatbelt, the 60 ha site consists of concentric rows of Eucalyptus sargentii trees with mounded (6 - 11 cm high) 10 -15 m inter-rows of Atriplex spp. Sustainability of this system and fulfilment of productive and ameliorative functions is dependant on successful recruitment (perennials). Although the present study site was conducted on farmland in a Mediterranean-type climate, low annual rainfall and spatial arrangement of perennial shrubs and trees, allow useful comparisons to be made with naturally occurring banded semi-arid systems and vice-versa. Of key interest were leaf-litter redistribution and trapping by tree and shrub rows and whether litter-cover/microsites facilitated/interfered with seedling recruitment (establishment, growth and survival). Litter from the tree row, redistributed by prevailing winds and rain, accumulated adjacent to saltbush seeding mounds, creating a mosaic of bare and littered areas across the site (total litter 10 t/ha over 22 months). Accumulated litter was hypothesized to differentially influence seasonal soil abiotic parameters (depending on litter-cover density) including; salinity, water availability, infiltration rates, water repellency and temperature. These abiotic conditions were also hypothesized to vary between tree and shrub microsites. Biotically, recruitment at this site was also hypothesized to be determined by interactions (positive and negative) between perennial components and understorey annuals/perennial seedlings. Accumulation of litter and resultant heterogeneity was influenced by shrub morphology, microtopography, wind direction and distance from litter source, with increased litter on the leeward sides of hemispherical Atriplex undulata shrubs and shrubs closest to tree rows. ... The importance of tree/shrub microsites varied seasonally, with no influence in winter due to moderate temperatures and increased water availability. In warmer months saltbush mid-row microsites were most favourable for seedling recruitment due to moderate litter-cover; reducing salinity, temperatures and increasing infiltration; and reduced root-competition/shading by the tree row. Tree microsites also directly inhibited seedling recruitment through increased salinities and water repellency. However, trees also indirectly facilitated recruitment in adjacent areas through provision of leaf-litter. As litter-trapping and recruitment patterns at this site mirror those found in semi-arid natural and artificial systems, the results of this study provide useful insights into creating appropriate mimics of low rainfall natural banded woodland and chenopod shrublands. Saltbush seeding mounds, shrub morphology and litter were key components for litter trapping and recruitment heterogeneity at this site. In this tree/shrub alley planting, where litter quantities directly influence vegetation cover densities, future saline plantings need to consider appropriate tree/shrub row spacings and orientation for efficient resource (seeds, litter and water) capture.

[Truncated abstract] Insufficient water use by annual crop and pasture species leading to costly rises in saline watertables has prompted research into potentially profitable deep-rooted perennial species in the Western Australian wheatbelt. Native mallee eucalypts are currently being developed as a short-rotation coppice crop for production of leaf oils, activated carbon and bio-electricity for low rainfall areas (300—450 mm) too dry for many of the traditional timber and forage species. The research in this study was aimed at developing a knowledge base necessary to grow and manage coppiced mallee eucalypts for both high productivity and salinity control. This firstly necessitated identification of suitable species, climatic and site requirements favourable to rapid growth, and understanding of factors likely to affect yield of the desirable leaf oil constituent, 1,8-cineole. This was undertaken using nine mallee taxa at twelve sites with two harvest regimes. E. kochii subsp. plenissima emerged as showing promise in the central and northern wheatbelt, particularly at a deep acid sand site (Gn 2.61; Northcote, 1979), so further studies focussed on physiology of its resprouting, water use and water-use efficiency at a similar site near Kalannie. Young E. kochii trees were well equipped with large numbers of meristematic foci and adequate root starch reserves to endure repeated shoot removal. The cutting season and interval between cuts were then demonstrated to have a strong influence on productivity, since first-year coppice growth was slow and root systems appeared to cease in secondary growth during the first 1.5—2.5 years after cutting. After decapitation, trees altered their physiology to promote rapid replacement of shoots. Compared to uncut trees, leaves of coppices were formed with a low carbon content per unit area, and showed high stomatal conductance accompanied by high leaf photosynthetic rates. Whole-plant water use efficiency of coppiced trees was unusually high due to their fast relative growth rates associated with preferential investments of photosynthates into regenerating canopies rather than roots. Despite relatively small leaf areas on coppice shoots over the two years following decapitation, high leaf transpiration rates resulted in coppices using water at rates far in excess of that falling as rain on the tree belt area. Water budgets showed that 20 % of the study paddock would have been needed as 0—2 year coppices in 5 m wide twin-row belts in order to maintain hydrological balance over the study period. Maximum water use occurred where uncut trees were accessing a fresh perched aquifer, but where this was not present water budgets still showed transpiration of uncut trees occurring at rates equivalent to 3—4 times rainfall incident on the tree belt canopy. In this scenario, only 10 % of the paddock surface would have been required under 5 m wide tree belts to restore hydrological balance, but competition losses in adjacent pasture would have been greater

The impact of a canker disease of Corymbia calophylla (marri) in the southwest of Western Australia (WA) has increased substantially since it was first observed causing decline and death of this species in the 1970s. By the early 1990s there were expressions of concern and calls to determine the cause and management options. Despite this, there has been very little research into the incidence, severity and possible causes of the disease. There are, however, historical reports dating back to the 1920s of a canker disease of amenity planted C. ficifolia caused by Sporotrichum destructor, though the diagnosis and Latin description were never published. It has been suggested that there may be links between this species and the genus Quambalaria, a group containing leaf and shoot pathogens of species of Eucalyptus and Corymbia. This study examined the incidence and symptomology of the disease, the range of fungal species associated with healthy and diseased C. calophylla, and the pathogenicity of isolates obtained from these surveys. Also investigated was the identity of the pathogen, S. destructor, historically attributed to canker disease of C. ficifolia, to determine whether this pathogen is responsible for the current epidemic of C. calophylla canker and if it is synonymous with Quambalaria. Cankers were present on trees across the range of surveyed sites, with lesions occurring on trunks, branches or twigs of 25.7 % of the C. calophylla surveyed. Canker incidence was significantly greater on trees present at remnant sites, such as roadsides and in paddocks, than forest trees being 38 % and 13.3 %, respectively. Tree height, trunk diameter at breast height and crown position, size and health ratings were significant predictors of canker presence, with cankers more common on larger, older trees and trees with poor crown condition. Bark cracks exuding kino were present on 48 % of the surveyed trees, and when dissected, lesions were observed on 40 %, suggesting that these cracks could be the initial stages of canker disease. This was confirmed by the observation of a number of cracks that developed into perennial cankers during the three year study. Monitoring of canker development and the examination of transverse sections showed the circumvention of host defenses by the pathogen and the subsequential walling off response of the host (which typifies perennial cankers) was not necessarily an annual event, with no change observed in some cankers over the three year period, while others progressed rapidly in that time, occasionally to the point of girdling and killing the host. Initial surveys isolated 44 fungal species from healthy and diseased C. calophylla, with opportunistic pathogens including Endothiella eucalypti and Cytospora eucalypticola common. Subsequent surveys foccussed more on a potential pathogen in the genus Quambalaria, which was rarely isolated from active lesions, presumably because of its slow growth rate, but which sporulated consistently on the surface of older sections of the cankers. DNA sequences confirmed that Q. cyanescens and Q. pitereka are present in southwest WA, with the latter associated with leaf and shoot disease. A third and new species of Quambalaria was isolated from cankers. Comparisons of disease symptoms and conidiogenesis indicate this species is synonymous with S. destructor. The species is formally described here as Q. coyrecup T. Paap sp. nov. A pathogenicity trial was unsuccessful in causing disease symptoms in trees inoculated with core plugs taken from canker lesion margins of diseased trees, though the time frame and environmental factors may not have been adequate for disease development. The core plug inoculation method may also have failed because opportunistic pathogens which were frequently isolated from lesions out-competed Q. coyrecup (paralleling the results achieved by culturing from lesions). Quambalaria coyrecup caused symptoms matching those observed in natural infections when suitable hosts were inoculated, confirming it is the fungus responsible for the current canker disease of C. calophylla and C. ficifolia. Endothiella eucalypti also caused significant lesions, though these were not typical of natural infections, which together with its frequent isolation from both healthy and diseased trees suggests it is an opportunistic pathogen, potentially contributing to disease development in trees already infected with Q. coyrecup. Isolates of Q. pitereka from WA and eastern Australia both caused typical shoot blight symptoms in the WA hosts C. calophylla and C. ficifolia, and the eastern Australian host C. maculata, though a larger path trial is required to examine the possibility of host specificity. Quambalaria cyanescens was non-pathogenic in all inoculation trials. The current cause of cankers in C. calophylla is now known to be the same as the fungus historically implicated in the canker disease of C. ficifolia, when at the time it was described as an endophyte doing little or no damage in C. calophylla. Thus, it is of immediate importance to determine the factors driving this decline, and develop control and management options.

Bauxite mining is a major activity in the jarrah (Eucalyptus marginata Donn ex Sm.) forest of south-western Australia. After mining, poor tree growth can occur in some areas. This thesis aimed to determine whether soil constraints, including reduced depth and compaction, were responsible for poor tree growth at low-quality restored bauxite mines. In particular, this study determined the response of jarrah root morphology, leaf-scale physiology and growth/development to soil constraints at two contrasting (low-quality and high-quality) restored bauxite-mine sites. Jarrah root excavations at a low-quality restored site revealed that deep-ripping equipment failed to penetrate the cemented lateritic subsoil, causing coarse roots to be restricted to the top 0.5 m of the soil profile, resulting in fewer and smaller jarrah trees. An adjacent area within the same mine pit (high-quality site) had a kaolinitic clay subsoil, which coarse roots were able to penetrate to the average ripping depth of 1.5 m. Impenetrable subsoil prevented development of taproots at the low-quality site, with trees instead producing multiple lateral and sinker roots. Trees in riplines, made by deep-ripping, at the high-quality site accessed the subsoil via a major taproot, while those on crests developed large lateral and sinker roots. Bauxite mining is a major activity in the jarrah (Eucalyptus marginata Donn ex Sm.) forest of south-western Australia. After mining, poor tree growth can occur in some areas. This thesis aimed to determine whether soil constraints, including reduced depth and compaction, were responsible for poor tree growth at low-quality restored bauxite mines. In particular, this study determined the response of jarrah root morphology, leaf-scale physiology and growth/development to soil constraints at two contrasting (low-quality and high-quality) restored bauxite-mine sites. Jarrah root excavations at a low-quality restored site revealed that deep-ripping equipment failed to penetrate the cemented lateritic subsoil, causing coarse roots to be restricted to the top 0.5 m of the soil profile, resulting in fewer and smaller jarrah trees. An adjacent area within the same mine pit (high-quality site) had a kaolinitic clay subsoil, which coarse roots were able to penetrate to the average ripping depth of 1.5 m. Impenetrable subsoil prevented development of taproots at the low-quality site, with trees instead producing multiple lateral and sinker roots. Trees in riplines, made by deep-ripping, at the high-quality site accessed the subsoil via a major taproot, while those on crests developed large lateral and sinker roots.

Phytophthora cinnamomi is an introduced soilborne phytopathogen to Western Australia (WA) and impacts on 2000 of the approximately 9000 plant species indigenous in the southwest of WA. Amongst these is Eucalyptus marginata (jarrah), the dominant and economically important hardwood timber species of the jarrah forest. This thesis aimed to investigate the morphological, pathogenic and genotypic variation in two local WA populations of P. cinnamomi isolates. The populations were selected from areas where jarrah clonal lines selected for resistance to P. cinnamomi may be used in the rehabilitation of infested jarrah forest and rehabilitated bauxite minesites in the southwest of WA. Resistance against a range of isolates using different inoculation methods. Seventy-three isolates of P. cinnamomi were collected from diseased jarrah and Corymbia calophylla (marri) trees from two populations located 70 km apart and these were examined for phenotypic and genotypic variation. Microsatellite DNA analysis showed that all isolates were of the same clonal lineage. In P. cinnamomi for the first time I show that there is a broad and continuous variation in the morphology and pathology between two populations of one clonal lineage, and that all phenotypes varied independently from one another. No relationship was found between morphological and pathogenic characters. The ability of isolates in both populations to cause deaths ranged from killing all plants within 59 days to plants being symptomless 182 days after inoculation. Single and multiple paragynous antheridia formed along with amphigynous ones in mating studies with all WA isolates and a sample of worldwide isolates. Developmental studies and cytological examination showed fertilisation tubes developed asynchronously or synchronously from both antheridial types and indicated that either antheridial type contributed a nucleus for fertilisation of the oosphere. This is the first report of paragynous antheridial associations in P. cinnamomi. Antheridial variation is a characteristic that needs to be adjusted in the taxonomic Phytophthora identification keys. In underbark and zoospore stem inoculations of three 1.5-year-old jarrah clonal lines (two ranked as resistant (RR) and one as susceptible (SS) to P. cinnamomi in the original selection trials) at 15, 20, 25 and 30°C, it was found that the method of inoculation did not produce comparable results, particularly at 25 and 30°C. At these temperatures, all three clonal lines had 100% mortality when inoculated underbark, but when inoculated with zoospores, one RR line had 60% survival and the SS and remaining RR line had 100% mortality. Generally, the level of resistance of all clonal lines declined with increasing temperature. Lesion development was measured at 20, 25 and 30°C for 4 days in detached branches of an RR and SS clonal line inoculated underbark with four different P. cinnamomi isolates. Detached branches were found to be a potential screen for jarrah resistance to P. cinnamomi and to allow the identification of susceptible and resistant clonal lines at 30°C. Lesion and colonisation development of P. cinnamomi isolates were assessed in situ (late autumn) of seed-grown and clonal lines of 3.5 to 4.5 year-old jarrah trees growing in a rehabilitated minesite jarrah forest in underbark inoculation of lateral branches (1995) or simultaneously in lateral branches and lateral roots (1996). Trees were underbark inoculated in lateral branches and lateral roots. Colonisation was more consistent as a measure of resistance than lesion length over the two trials because it accounted for the recovery of P. cinnamomi from macroscopically symptomless tissue beyond lesions, which on some occasions, was up to 6 cm. In the two trials, one RR clonal line consistently had small lesion and colonisation lengths in branches and roots. In contrast, the remaining two RR clonal lines had similar lesion and colonisation lengths to the SS clonal line and may, therefore, not be suitable for use in the rehabilitation of P. cinnamomi infested areas. The relative rankings of the jarrah clonal lines by colonisation lengths were similar between branch and root inoculations. Branch inoculations are a valid option for testing resistance and susceptibility of young jarrah trees to P. cinnamomi. The pathogen was recovered on Phytophthora selective agar 3–6 months after inoculation from 50% of samples with lesions and 30% of symptomless samples in a series of growth cabinet, glasshouse and field experiments. However, up to 11% of samples with and without lesions and from which P. cinnamomi was not initially isolated contained viable pathogen after leaching the plant material in water over 9 days. This indicates that the pathogen could be present as dormant structures, such as chlamydospores, where dormancy needs to be broken for germination to occur, or fungistatic compounds in the tissue need to be removed to allow the pathogen to grow, or both. These results have important implications for disease diagnosis and management, disease-free certification and quarantine clearance. No clonal line of jarrah was found to be 100% resistant using different inoculation methods, environmental conditions and when challenged by individuals from a large range of P. cinnamomi isolates. Even the most promising RR line had individual replicates that were unable to contain lesions or died with time. This suggests that further screening work may be required using more isolates varying in their capacity to cause disease and a broader range of environmental conditions. Jarrah clonal lines that survive such rigorous screening could then be expected to survive planting out in a range of environments in the jarrah forest and rehabilitated bauxite minesites.

The impact of a canker disease of Corymbia calophylla (marri) in the southwest of Western Australia (WA) has increased substantially since it was first observed causing decline and death of this species in the 1970s. By the early 1990s there were expressions of concern and calls to determine the cause and management options. Despite this, there has been very little research into the incidence, severity and possible causes of the disease. There are, however, historical reports dating back to the 1920s of a canker disease of amenity planted C. ficifolia caused by Sporotrichum destructor, though the diagnosis and Latin description were never published. It has been suggested that there may be links between this species and the genus Quambalaria, a group containing leaf and shoot pathogens of species of Eucalyptus and Corymbia. This study examined the incidence and symptomology of the disease, the range of fungal species associated with healthy and diseased C. calophylla, and the pathogenicity of isolates obtained from these surveys. Also investigated was the identity of the pathogen, S. destructor, historically attributed to canker disease of C. ficifolia, to determine whether this pathogen is responsible for the current epidemic of C. calophylla canker and if it is synonymous with Quambalaria. Cankers were present on trees across the range of surveyed sites, with lesions occurring on trunks, branches or twigs of 25.7 % of the C. calophylla surveyed. Canker incidence was significantly greater on trees present at remnant sites, such as roadsides and in paddocks, than forest trees being 38 % and 13.3 %, respectively. Tree height, trunk diameter at breast height and crown position, size and health ratings were significant predictors of canker presence, with cankers more common on larger, older trees and trees with poor crown condition. Bark cracks exuding kino were present on 48 % of the surveyed trees, and when dissected, lesions were observed on 40 %, suggesting that these cracks could be the initial stages of canker disease. This was confirmed by the observation of a number of cracks that developed into perennial cankers during the three year study. Monitoring of canker development and the examination of transverse sections showed the circumvention of host defenses by the pathogen and the subsequential walling off response of the host (which typifies perennial cankers) was not necessarily an annual event, with no change observed in some cankers over the three year period, while others progressed rapidly in that time, occasionally to the point of girdling and killing the host. Initial surveys isolated 44 fungal species from healthy and diseased C. calophylla, with opportunistic pathogens including Endothiella eucalypti and Cytospora eucalypticola common. Subsequent surveys foccussed more on a potential pathogen in the genus Quambalaria, which was rarely isolated from active lesions, presumably because of its slow growth rate, but which sporulated consistently on the surface of older sections of the cankers. DNA sequences confirmed that Q. cyanescens and Q. pitereka are present in southwest WA, with the latter associated with leaf and shoot disease. A third and new species of Quambalaria was isolated from cankers. Comparisons of disease symptoms and conidiogenesis indicate this species is synonymous with S. destructor. The species is formally described here as Q. coyrecup T. Paap sp. nov. A pathogenicity trial was unsuccessful in causing disease symptoms in trees inoculated with core plugs taken from canker lesion margins of diseased trees, though the time frame and environmental factors may not have been adequate for disease development. The core plug inoculation method may also have failed because opportunistic pathogens which were frequently isolated from lesions out-competed Q. coyrecup (paralleling the results achieved by culturing from lesions). Quambalaria coyrecup caused symptoms matching those observed in natural infections when suitable hosts were inoculated, confirming it is the fungus responsible for the current canker disease of C. calophylla and C. ficifolia. Endothiella eucalypti also caused significant lesions, though these were not typical of natural infections, which together with its frequent isolation from both healthy and diseased trees suggests it is an opportunistic pathogen, potentially contributing to disease development in trees already infected with Q. coyrecup. Isolates of Q. pitereka from WA and eastern Australia both caused typical shoot blight symptoms in the WA hosts C. calophylla and C. ficifolia, and the eastern Australian host C. maculata, though a larger path trial is required to examine the possibility of host specificity. Quambalaria cyanescens was non-pathogenic in all inoculation trials. The current cause of cankers in C. calophylla is now known to be the same as the fungus historically implicated in the canker disease of C. ficifolia, when at the time it was described as an endophyte doing little or no damage in C. calophylla. Thus, it is of immediate importance to determine the factors driving this decline, and develop control and management options.

As the Eucalyptus globulus plantation industry expands and matures in southwestern Australia (WA), the impact of disease within the plantation environment is predicted to increase. This thesis investigated the most abundant canker-causing pathogen associated with branch and stem cankers, Endothiella eucalypti the anamorph of Cryphonectria eucalypti. Endothiella eucalypti was widespread, although at low incidence, throughout the WA plantation estate and was frequently observed sporulating on the bark of healthy hosts in the absence of disease. Regions with a long (approximately 20 years) plantation history, such as Bunbury, had the highest incidence of this pathogen. A high degree of variability in pathogenicity, growth rate and colony morphology was observed between WA isolates of En. eucalypti. In the glasshouse, a significant variation in susceptibility of seven E. globulus provenances to En. eucalypti was observed. Although an interaction between the E. globulus provenance and En. eucalypti isolate was recorded, some provenances were generally more susceptible than others. In two 18-month-old plantations, the susceptibility of three provenances to En. eucalypti was significantly influenced by environmental conditions. Visual assessment of general tree health indicated that less healthy trees had smaller lesions than healthy trees. It is hypothesised that the selection of E. globulus provenances to suit site conditions in the future should decrease the risk of serious disease, especially on marginal sites. Endothiella eucalypti caused disease in intact stems of two-year-old E. globulus under glasshouse conditions. This suggests that En. eucalypti may not require a wound to infect in the field. Vegetative compatibility groupings between WA En. eucalypti isolates indicated a relatively high degree of genotypic diversity within the WA asexual population of En. eucalypti, whereas inter-simple sequence repeats PCR (ISSR-PCR) analysis indicated a lower level of genotypic diversity. Discrepancies between traditional and molecular techniques, such as ISSR-PCR, was attributed to the more specific gene-togene analysis afforded by molecular techniques. ISSR-PCR successfully distinguished variability within the En. eucalypti population and with the teleomorph, isolated in South Africa. It also separated Cryphonectria cubensis isolates from the C. eucalypti isolates. As copper is the micronutrient most limiting growth of E. globulus in WA, its role in the resistance of two E. globulus provenances was examined in a glasshouse trial. Lesion extension or defence responses of E. globulus to En. eucalypti did not differ between Cu-adequate and Cu-deficient plants. It is suggested that constitutive levels of host defence enzymes played a more important role in providing protection for the host against En. eucalypti than the external supply of copper. A reduction in the canopy volume of E. globulus within plantations due to insect herbivory or foliar pathogens, such as Mycosphaerella spp., has been reported to predispose the host to disease caused by non-aggressive canker-causing fungi. Under two separate glasshouse trials, conditions of 100% defoliation and 80% defoliation maintained over six weeks prior to inoculation, were required to significantly increase lesion extension caused by En. eucalypti in E. globulus stems. The ability of defoliated E. globulus to retain a degree of resistance to En. eucalypti was attributed to the rapid replacement of foliage and up-regulation of photosynthesis in remaining leaves. The carbohydrate reserves of the plant were depleted following defoliation and remained depressed regardless of the length of time the trees remained defoliated. In conclusion, the endophytic habit of En. eucalypti poses a threat to highly stressed trees, however it does not appear to be an immediate threat to WA plantation health. Although En. eucalypti has not yet been responsible for a major disease outbreak in WA, the impact of this disease on plantation-grown eucalypts elsewhere in Australia and worldwide serves as an indication of its potential to affect WA plantations.

Phytophthora cinnamomi is an introduced soilborne phytopathogen to Western Australia (WA) and impacts on 2000 of the approximately 9000 plant species indigenous in the southwest of WA. Amongst these is Eucalyptus marginata (jarrah), the dominant and economically important hardwood timber species of the jarrah forest. This thesis aimed to investigate the morphological, pathogenic and genotypic variation in two local WA populations of P. cinnamomi isolates. The populations were selected from areas where jarrah clonal lines selected for resistance to P. cinnamomi may be used in the rehabilitation of infested jarrah forest and rehabilitated bauxite minesites in the southwest of WA. Resistance against a range of isolates using different inoculation methods. Seventy-three isolates of P. cinnamomi were collected from diseased jarrah and Corymbia calophylla (marri) trees from two populations located 70 km apart and these were examined for phenotypic and genotypic variation. Microsatellite DNA analysis showed that all isolates were of the same clonal lineage. In P. cinnamomi for the first time I show that there is a broad and continuous variation in the morphology and pathology between two populations of one clonal lineage, and that all phenotypes varied independently from one another. No relationship was found between morphological and pathogenic characters. The ability of isolates in both populations to cause deaths ranged from killing all plants within 59 days to plants being symptomless 182 days after inoculation. Single and multiple paragynous antheridia formed along with amphigynous ones in mating studies with all WA isolates and a sample of worldwide isolates. Developmental studies and cytological examination showed fertilisation tubes developed asynchronously or synchronously from both antheridial types and indicated that either antheridial type contributed a nucleus for fertilisation of the oosphere. This is the first report of paragynous antheridial associations in P. cinnamomi. Antheridial variation is a characteristic that needs to be adjusted in the taxonomic Phytophthora identification keys. In underbark and zoospore stem inoculations of three 1.5-year-old jarrah clonal lines (two ranked as resistant (RR) and one as susceptible (SS) to P. cinnamomi in the original selection trials) at 15, 20, 25 and 30 degrees C, it was found that the method of inoculation did not produce comparable results, particularly at 25 and 30 degrees C. At these temperatures, all three clonal lines had 100% mortality when inoculated underbark, but when inoculated with zoospores, one RR line had 60% survival and the SS and remaining RR line had 100% mortality. Generally, the level of resistance of all clonal lines declined with increasing temperature. Lesion development was measured at 20, 25 and 30 degrees C for 4 days in detached branches of an RR and SS clonal line inoculated underbark with four different P. cinnamomi isolates. Detached branches were found to be a potential screen for jarrah resistance to P. cinnamomi and to allow the identification of susceptible and resistant clonal lines at 30 degrees C. Lesion and colonisation development of P. cinnamomi isolates were assessed in situ (late autumn) of seed-grown and clonal lines of 3.5 to 4.5 year-old jarrah trees growing in a rehabilitated minesite jarrah forest in underbark inoculation of lateral branches (1995) or simultaneously in lateral branches and lateral roots (1996). Trees were underbark inoculated in lateral branches and lateral roots. Colonisation was more consistent as a measure of resistance than lesion length over the two trials because it accounted for the recovery of P. cinnamomi from macroscopically symptomless tissue beyond lesions, which on some occasions, was up to 6 cm. In the two trials, one RR clonal line consistently had small lesion and colonisation lengths in branches and roots. In contrast, the remaining two RR clonal lines had similar lesion and colonisation lengths to the SS clonal line and may, therefore, not be suitable for use in the rehabilitation of P. cinnamomi infested areas. The relative rankings of the jarrah clonal lines by colonisation lengths were similar between branch and root inoculations. Branch inoculations are a valid option for testing resistance and susceptibility of young jarrah trees to P. cinnamomi. The pathogen was recovered on Phytophthora selective agar 3-6 months after inoculation from 50% of samples with lesions and 30% of symptomless samples in a series of growth cabinet, glasshouse and field experiments. However, up to 11% of samples with and without lesions and from which P. cinnamomi was not initially isolated contained viable pathogen after leaching the plant material in water over 9 days. This indicates that the pathogen could be present as dormant structures, such as chlamydospores, where dormancy needs to be broken for germination to occur, or fungistatic compounds in the tissue need to be removed to allow the pathogen to grow, or both. These results have important implications for disease diagnosis and management, disease-free certification and quarantine clearance. No clonal line of jarrah was found to be 100% resistant using different inoculation methods, environmental conditions and when challenged by individuals from a large range of P. cinnamomi isolates. Even the most promising RR line had individual replicates that were unable to contain lesions or died with time. This suggests that further screening work may be required using more isolates varying in their capacity to cause disease and a broader range of environmental conditions. Jarrah clonal lines that survive such rigorous screening could then be expected to survive planting out in a range of environments in the jarrah forest and rehabilitated bauxite minesites.

[Truncated abstract. Please see the pdf format for the complete text. Also, formulae and special characters can only be approximated here. Please see the pdf version for an accurate reproduction.] This thesis examines the water use, ecophysiology and hydraulic architecture of Eucalyptus marginata (jarrah) growing on bauxite mine rehabilitation sites in the jarrah forest of south-western Australia. The principal objective was to characterise the key environment and plant-based influences on tree water use, and to better understand the dynamics of water use over a range of spatial and temporal scales in this drought-prone ecosystem. A novel sap flow measurement system (based on the use of the heat pulse method) was developed so that a large number of trees could be monitored concurrently in the field. A validation experiment using potted jarrah saplings showed that rates of sap flow (transpiration) obtained using this system agreed with those obtained gravimetrically. Notably, diurnal patterns of transpiration were measured accurately and with precision using the newly developed heat ratio method. Field studies showed that water stress and water use by jarrah saplings on rehabilitation sites were strongly seasonal: being greatest in summer when it was warm and dry, and least in winter when it was cool and wet. At different times, water use was influenced by soil water availability, vapour pressure deficit (VPD) and plant hydraulic conductance. In some areas, there was evidence of a rapid decline in transpiration in response to dry soil conditions. At the end of summer, most saplings on rehabilitation sites were not water stressed, whereas water status in the forest was poor for small saplings but improved with increasing size. It has been recognised that mature jarrah trees avoid drought by having deep root systems, however, it appears that saplings on rehabilitation sites may have not yet developed functional deep roots, and as such, they may be heavily reliant on moisture stored in surface soil horizons. Simple predictive models of tree water use revealed that stand water use was 74 % of annual rainfall at a high density (leaf area index, LAI = 3.1), high rainfall (1200 mm yr-1) site, and 12 % of rainfall at a low density (LAI = 0.4), low rainfall (600 mm yr-1) site, and that water use increased with stand growth. A controlled field experiment confirmed that: (1) sapling transpiration was restricted as root-zone water availability declined, irrespective of VPD; (2) transpiration was correlated with VPD when water was abundant; and (3) transpiration was limited by soil-to-leaf hydraulic conductance when water was abundant and VPD was high (> 2 kPa). Specifically, transpiration was regulated by stomatal conductance. Large stomatal apertures could sustain high transpiration rates, but stomata were sensitive to hydraulic perturbations caused by soil water deficits and/or high evaporative demand. No other physiological mechanisms conferred immediate resistance to drought. Empirical observations were agreeably linked with a current theory suggesting that stomata regulate transpiration and plant water potential in order to prevent hydraulic dysfunction following a reduction in soil-to-leaf hydraulic conductance. Moreover, it was clear that plant hydraulic capacity determined the pattern and extent of stomatal regulation. Differences in hydraulic capacity across a gradient in water availability were a reflection of differences in root-to-leaf hydraulic conductance, and were possibly related to differences in xylem structure. Saplings on rehabilitation sites had greater hydraulic conductance (by 50 %) and greater leaf-specific rates of transpiration at the high rainfall site (1.5 kg m-2 day1) than at the low rainfall site (0.8 kg m-2 day1) under near optimal conditions. Also, rehabilitation-grown saplings had significantly greater leaf area, leaf area to sapwood area ratios and hydraulic conductance (by 30-50 %) compared to forest-grown saplings, a strong indication that soils in rehabilitation sites contained more water than soils in the forest. Results suggested that: (1) the hydraulic structure and function of saplings growing under the same climatic conditions was determined by soil water availability; (2) drought reduced stomatal conductance and transpiration by reducing whole-tree hydraulic conductance; and (3) saplings growing on open rehabilitation sites utilised more abundant water, light and nutrients than saplings growing in the forest understorey. These findings support a paradigm that trees evolve hydraulic equipment and physiological characteristics suited to the most efficient use of water from a particular spatial and temporal niche in the soil environment.

[Truncated abstract] Eleven sites, each with the trio of land uses: Eucalyptus globulus plantation, pasture and natural vegetation, representing from the Mediterranean climate, high rainfall region (<550 mm annually) of south-western Australia were investigated to assess medium-term changes in the P-supplying capacity of soils in eucalypt plantations growing on agricultural land. The natural vegetation soils were a benchmark for comparing soil P change since land clearing and development for agriculture. The experimental framework provided an ideal basis for studying changes in P forms since land clearing and fertilization for agriculture and the ensuing conversion to plantations (on an average 9 years ago). Conventional soil P indices measure plant available P that is more relevant to short duration annual crops and pastures. To predict medium-term P availability, P forms were determined using Hedley et al.’s (1982) fractionation scheme and fractions were grouped using the Guo and Youst (1998) criteria into readily, moderately and sparingly available P. The P species were also determined by 31P NMR spectroscopy of 0.5M NaOH-0.1M EDTA extracts. Hedley et al.’s (1982) inorganic P extracted by anion exchange resin and by NaHCO3 are widely considered to be approximations to the actual plant available P. The availability to plants of other P fractions is less certain and this is examined in an experiment to compare the plant availability of various P fractions in soils from fertilized and unfertilized land uses following exhaustive cropping in the glasshouse. The soil texture for the sites studied included coarse sand, loamy sand, clayey sand, and sandy loam. Surface soils (0-10 cm) have pH(CaCl2) in the acidic range (mean 4.4) and there is no significant difference due to differences in land use (P<0.05). The soils are of low EC (1:5 H2O) - 6 mS m-1. There is an almost 5-fold variation in organic C among sites (from 1.4% to 8%) but organic C values did not show any significant effect (P<0.05) of changes in land use. To evaluate the degree of similarity of soils within each triplet set at a site principal component analysis was carried out on those soil chemical⁄mineralogical characteristics that were least likely to be affected by changes in land use practices. This analysis showed good matching of the triplet of sub-sites on the whole, especially for the duo of pasture and plantation land uses. This degree of matching of the trio of land uses was considered while interpreting the effects of land use on the forms and behaviour of soil P, and variations due to various extents of mismatch were mostly addressed using statistical techniques including regression analysis to interpret sub-site difference

As the Eucalyptus globulus plantation industry expands and matures in southwestern Australia (WA), the impact of disease within the plantation environment is predicted to increase. This thesis investigated the most abundant canker-causing pathogen associated with branch and stem cankers, Endothiella eucalypti the anamorph of Cryphonectria eucalypti. Endothiella eucalypti was widespread, although at low incidence, throughout the WA plantation estate and was frequently observed sporulating on the bark of healthy hosts in the absence of disease. Regions with a long (approximately 20 years) plantation history, such as Bunbury, had the highest incidence of this pathogen. A high degree of variability in pathogenicity, growth rate and colony morphology was observed between WA isolates of En. eucalypti. In the glasshouse, a significant variation in susceptibility of seven E. globulus provenances to En. eucalypti was observed. Although an interaction between the E. globulus provenance and En. eucalypti isolate was recorded, some provenances were generally more susceptible than others. In two 18-month-old plantations, the susceptibility of three provenances to En. eucalypti was significantly influenced by environmental conditions. Visual assessment of general tree health indicated that less healthy trees had smaller lesions than healthy trees. It is hypothesised that the selection of E. globulus provenances to suit site conditions in the future should decrease the risk of serious disease, especially on marginal sites. Endothiella eucalypti caused disease in intact stems of two-year-old E. globulus under glasshouse conditions. This suggests that En. eucalypti may not require a wound to infect in the field. Vegetative compatibility groupings between WA En. eucalypti isolates indicated a relatively high degree of genotypic diversity within the WA asexual population of En. eucalypti, whereas inter-simple sequence repeats PCR (ISSR-PCR) analysis indicated a lower level of genotypic diversity. Discrepancies between traditional and molecular techniques, such as ISSR-PCR, was attributed to the more specific gene-togene analysis afforded by molecular techniques. ISSR-PCR successfully distinguished variability within the En. eucalypti population and with the teleomorph, isolated in South Africa. It also separated Cryphonectria cubensis isolates from the C. eucalypti isolates. As copper is the micronutrient most limiting growth of E. globulus in WA, its role in the resistance of two E. globulus provenances was examined in a glasshouse trial. Lesion extension or defence responses of E. globulus to En. eucalypti did not differ between Cu-adequate and Cu-deficient plants. It is suggested that constitutive levels of host defence enzymes played a more important role in providing protection for the host against En. eucalypti than the external supply of copper. A reduction in the canopy volume of E. globulus within plantations due to insect herbivory or foliar pathogens, such as Mycosphaerella spp., has been reported to predispose the host to disease caused by non-aggressive canker-causing fungi. Under two separate glasshouse trials, conditions of 100% defoliation and 80% defoliation maintained over six weeks prior to inoculation, were required to significantly increase lesion extension caused by En. eucalypti in E. globulus stems. The ability of defoliated E. globulus to retain a degree of resistance to En. eucalypti was attributed to the rapid replacement of foliage and up-regulation of photosynthesis in remaining leaves. The carbohydrate reserves of the plant were depleted following defoliation and remained depressed regardless of the length of time the trees remained defoliated. In conclusion, the endophytic habit of En. eucalypti poses a threat to highly stressed trees, however it does not appear to be an immediate threat to WA plantation health. Although En. eucalypti has not yet been responsible for a major disease outbreak in WA, the impact of this disease on plantation-grown eucalypts elsewhere in Australia and worldwide serves as an indication of its potential to affect WA plantations.

Species prescriptions are developed for revegetating abandoned acidic coal overburden seepage sites in the Collie region of Western Australia. The research involved selecting appropriate plant species and determining successful methods of enhancing revegetation. Candidate species were screened for tolerance to acidic overburden materials, local climate conditions and metal toxicity. Methods tested included improving spoil conditions and trialing an alternative method for seeding.Twelve species of native plants were tested for tolerance in two acid overburden materials in pot and field trials. Eucalyptus robusta is the most tolerant, Eucalyptus camaldulensis and Eucalyptus cladocalyx are highly tolerant, Eucalyptus rudis and Melaleuca hamulosa demonstrate potential, provided adequate soil moisture is available.An important growth restriction factor in acid soils is the presence of free aluminium ions. A glasshouse trial performed on seven species for tolerance to aluminium toxicity revealed E. robusta as most tolerant and E. camaldulensis and Kunzea ericifolia a highly tolerant. E. rudis and M. hamulosa are moderately tolerant, but E. cladocalyx and Eucalyptus diversicolor are very sensitive to aluminium.Various methods were trialed to increase growth of seedlings transplanted on to acidic overburden sites. Both commercial cow manure and slow-release fertiliser tablets increase growth, whereas commercial potting mix and lime do not. Inoculation of plants with the ectomycorrhiza fungus Pisolithus tinctorius increases the amount of infection in roots but does not enhance plant growth.Supplementary fertilisation is necessary to maintain growth (nitrogen) and restore chlorophyll production (phosphorus) in fast growing eucalypt seedlings planted into typical acidic spoils. Poor levels of nutrient availability in such acidic sites appear to be the primary factor in ++
retarding growth. In the absence of supplementation, foliage reddening is observed in several species.An alternative method of seeding dumps is fascining. Prepared dump surfaces may be covered with capsule-laden branchwood of myrtaceous species. Material of the locally available Kunzea ericifolia is effective in producing many seedlings. Subsequent seedling growth is enhanced with fertiliser and lime addition.

Plantation eucalypts are a recent and rapidly growing industry in Australia, and will eventually replace the logging of old-growth forests. Over 40% of these plantations have been established in south-western Australia, where more than 160 000 ha of Eucalyptus globulus plantations now occur. In the early 1900’s, this species was widely planted as an exotic in South Africa, but succumbed to severe pest (Gonipterus sp.) and disease (Mycosphaerella sp.) problems. Similarly, in south-western Australia E. globulus is an exotic species, but with the additional threat that it is planted adjacent to indigenous eucalypts, which increases the possibility of pests and pathogens switching between closely related eucalypt hosts. Over the past ten years, there have been anecdotal reports of increasing levels of Mycosphaerella leaf disease (MLD) in E. globulus plantations in south-western Australia. This increase in disease level is of concern to the industry. To date there have been no comprehensive studies into the taxonomy, biogeography and population genetics of MLD in south-western Australia. This thesis investigated the impact of MLD in south-western Australia with a focus on its impact, taxonomy, biogeography and population genetics. It is the first study worldwide to quantify the relative impact of different Mycosphaerella species in a regional plantation estate. A survey of pest, disease and nutritional disorders (Chapter 2) found that MLD was the most severe and frequently occurring, single taxonomic health threat to 1 and 2-year-old E. globulus plantations in south-western Australia. For the first time, this survey identified and quantified the impact of pest and disease damage to E. globulus plantations in the region. There were differences in the disease levels between plantations and this was due to initial Mycosphaerella species composition and inoculum level, and local climatic conditions favourable for disease, rather than to the provenance planted or the nutritional status of the individual plantations. The survey for Mycosphaerella pathogens of eucalypts (Chapter 3) identified two new species of Mycosphaerella (M. ambiphylla and M. aurantia) and extended the known geographic range of eight other species (M. cryptica, M. gregaria, M. lateralis, M. marksii, M. mexicana , M. nubilosa, M. parva and M. suberosa). Of these: M. lateralis and M. mexicana were new records for Australia; and M. gregaria, M. nubilosa and M. parva were new records for Western Australia. A new anamorph, Phaeophloeospora ambiphylla was described and linked to M. ambiphylla. The occurrence of these new species and disease records in south-western Australia is significant for the plantation-eucalypt industry worldwide. The finding of two new species highlighted the need to quantify the disease impact of these on eucalypt plantations; and the extension of the range the remaining species raised important quarantine issues, concerned with the movement of plant material between regions and countries. The biogeographical investigation of Mycosphaerella (Chapter 4) identified that the most widespread and serious cause of MLD in south-western Australia is M. cryptica. In addition to occurring on the exotic E. globulus, it also occurs on two of the three important indigenous forestry eucalyptus species in this region. That is, on E. diversicolor, and E. marginata, but not on Corymbia calophylla. In terms of the plantation estate of E. globulus, however, M. nubilosa is the most widespread pathogen. The current study found that MLD on E. globulus is a complex of several different species, whereas, on E. diversicolor and E. marginata it is caused by only M. cryptica. Two species, M. cryptica and M. marksii were found commonly on adult E. globulus leaves. Although M. cryptica was the most frequent and serious cause of disease on adult leaves, M. marksii levels appear to be increasing and the future epidemiology of this pathogen should be closely monitored. There is some concern that these two MLD species could become an economically important problem on adult leaves of E. globulus. At present severe levels of MLD is significantly more common on juvenile than on adult foliage. The phylogenetic analysis (Chapter 5), based on ITS rDNA sequences from the present study and those obtained from GenBank accessions, found that Mycosphaerella is an assemblage of largely polyphyletic anamorph genera. Ten distinct clades emerged from the analysis. With the exception of the Dissoconium and the M. recutita clade, which comprised of two and one species respectively, none were comprised entirely of one anamorph genus alone. The anamorph genera represented were often dispersed across more than one clade, indicating that these anamorphs have arisen separately in different phylogenetic lineages. Cercospora, Stenella and Uwebraunia anamorphs each occurred in more than one separate clade. Although on the basis of rDNA sequence data Mycosphaerella appeared mostly monophyletic there was some evidence that the Mycosphaerella genus may be polyphyletic. This was particularly evident from the Dissoconium clade which grouped as closely to the outgroup Botryosphaeria taxon as it did with the remaining Mycosphaerella species. It was argued that a multi-gene phylogeny, which includes sequencing many species in other genera aligned with Mycosphaerella, is required in order to satisfactorily answer the question of whether Mycosphaerella is truly monophyletic. The phylogenetic analysis also showed that the taxonomy of Mycosphaerella based on ITS sequence data needs further clarification. Some species that are morphologically distinct, such as M. vespa and M. molleriana, shared identical ITS sequences. Other morphologically distinct species differed by as little as one or two nucleotides. Yet in other cases, the sequence variation amongst isolates from the same species differed substantially. Much of this variation in M. cryptica and other species was attributed to poorly edited sequences that had been lodged with GenBank. It was postulated that although a part of the remaining variation reflected the existence of cryptic species, some was likely to be genuine intra-species differences. It was concluded that further genes need to be sequenced, and more standardised cultural studies conducted in order to define species boundaries within Mycosphaerella. Based on the ITS rDNA sequence data, two different molecular methods for the identification of Mycosphaerella species from eucalypts were developed (Chapter 6). The first of these was a PCR-RFLP method that enabled the identification of Mycosphaerella species present on eucalypts in south-western Australia. A key is provided, which enabled the identification of species on a combination of PCR-RFLP DNA fragment migration patterns and a small number of morphological features. This key enables the identification of Mycosphaerella species more easily than keys that rely on morphological features alone. Therefore, this has made it easier for non-Mycosphaerella specialists to identify species from this genus. The second molecular method developed for the identification of Mycosphaerella species was that of primers that selectively amplify the DNA of M. cryptica and M. nubilosa, the two most important causes of MLD (Chapter 6). This will allow the rapid identification of these two species by non-specialists in Mycosphaerella taxonomy. The primers from the current study will also enable early diagnosis of the possible causal organism of MLD in a plantation. Once the use of these primers for amplifying DNA from leaf tissue has been optimised, they will also facilitate studies into the early infection process of M cryptica and M. nubilosa. For example, the presence of the pathogen may be detected prior to the appearance of symptoms. Studies may be conducted to determine the length of a hemi-biotrophic phase, and the extent of tissue colonisation both spatially and temporally, beyond the necrotic lesion in these two Mycosphaerella species. Previously, such studies have been hampered by the slow growth rate of these fungi in culture and the lack of media that would allow their selective isolation and detection by directly plating diseased and non-diseased host tissue. This study has clearly indicated that Mycosphaerella species are the major disease threat to E. globulus plantations in Western Australia. It has also shown that over the relatively short period of time of less than ten years the number of species recorded has increased from three to ten, and that disease severity has increased in plantations. It is therefore critical to continue the research on this genus in order to understand the biology, epidemiology and population genetics of this pathogen. This is necessary in order to inform tree selection and silvicultural practise that will minimise the future impact of MLD. This is particularly important if the industry moves towards clonal and hybrid forestry as has occurred elsewhere in the world. This study has laid the foundations for future research on this disease through the elucidation of the taxonomy of Mycosphaerella in south-western Australia and by providing some important molecular tools for its diagnosis and further study.

This study assessed the social and biophysical impacts of camping in the eucalypt forests of southwestern Western Australia. This was an integrated study examining both biophysical and social impacts in designated, developed and informal recreation areas used for camping. Four existing and proposed national parks and a Reserve, comprised of 110 designated and 12 informal campsites, provided the study sites. Previous research has focused on backcountry campsites and trails in wilderness areas in United States. A combined survey approach using multiple indicator ratings and measures was used to assess the biophysical impacts of camping. Adjustments to monitoring procedures used in backcountry areas were made so that the indicators were applicable to designated, developed campsites where a management footprint has been imposed. Visitors were surveyed at the designated campsites to establish how existing recreation opportunities were being used. Further, potential indicators and standards were identified to determine what kinds of social and resource conditions were acceptable to visitors and managers. A rating system was then developed combining biophysical and social indicators of importance to visitors and managers with their perceptions of acceptable change obtained from the surveys. Based on the indices derived from the rating system and results for a suite of associated indicators, designated campsites were significantly less impacted than informal ones. For both campsite types the amount of tree damage and litter exceeded the standards set by 50% of visitors and managers. Both visitors and managers were generally more concerned about biophysical impacts than they were about social ones, although site cleanliness was of concern. Both were generally satisfied with the size and number of groups encountered, in contrast to study findings from the United States. This study has developed and successfully applied an integrated approach to monitoring the impacts of recreational use on forested campsites in southwestern Australia. This system effectively and efficiently uses a combination of multiple indicator ratings and measures to produce an impact index, plus social surveys to provide information on conditions, indicators and standards of importance to managers and visitors. It also provides a means for the first time, of objectively monitoring designated, developed campsites where it is inappropriate to judge impacts against an undisturbed control.

This study assessed the social and biophysical impacts of camping in the eucalypt forests of southwestern Western Australia. This was an integrated study examining both biophysical and social impacts in designated, developed and informal recreation areas used for camping. Four existing and proposed national parks and a Reserve, comprised of 110 designated and 12 informal campsites, provided the study sites. Previous research has focused on backcountry campsites and trails in wilderness areas in United States. A combined survey approach using multiple indicator ratings and measures was used to assess the biophysical impacts of camping. Adjustments to monitoring procedures used in backcountry areas were made so that the indicators were applicable to designated, developed campsites where a management footprint has been imposed. Visitors were surveyed at the designated campsites to establish how existing recreation opportunities were being used. Further, potential indicators and standards were identified to determine what kinds of social and resource conditions were acceptable to visitors and managers. A rating system was then developed combining biophysical and social indicators of importance to visitors and managers with their perceptions of acceptable change obtained from the surveys. Based on the indices derived from the rating system and results for a suite of associated indicators, designated campsites were significantly less impacted than informal ones. For both campsite types the amount of tree damage and litter exceeded the standards set by 50% of visitors and managers. Both visitors and managers were generally more concerned about biophysical impacts than they were about social ones, although site cleanliness was of concern. Both were generally satisfied with the size and number of groups encountered, in contrast to study findings from the United States. This study has developed and successfully applied an integrated approach to monitoring the impacts of recreational use on forested campsites in southwestern Australia. This system effectively and efficiently uses a combination of multiple indicator ratings and measures to produce an impact index, plus social surveys to provide information on conditions, indicators and standards of importance to managers and visitors. It also provides a means for the first time, of objectively monitoring designated, developed campsites where it is inappropriate to judge impacts against an undisturbed control.