Box-Ironbark forests occur on the inland hills of the Great Dividing Range in Australia, from western Victoria to southern Queensland. These dry, open forests are characteristically dominated by Eucalyptus species such as Red Ironbark E. tricarpa, Mugga Ironbark E. sideroxylon and Grey Box E. microcarpa. Within these forests, several Eucalyptus species are a major source of nectar for the blossom-feeding birds and marsupials that form a distinctive component of the fauna. In Victoria, approximately 83% of the original pre - European forests of the Box-Ironbark region have been cleared, and the remaining fragmented forests have been heavily exploited for gold and timber. This exploitation has lead to a change in the structure of these forests, from one dominated by large 80-100 cm diameter, widely -spaced trees to mostly small (≥40 cm DBH), more densely - spaced trees. This thesis examines the flowering ecology of seven Eucalyptus species within a Box-Ironbark community. These species are characteristic of Victorian Box-Ironbark forests; River Red Gum E. camaldulensis, Yellow Gum E. leucoxylon, Red Stringybark E. macrorhyncha, Yellow Box E. melliodora, Grey Box E. microcarpa, Red Box E. polyanthemos and Red Ironbark E. tricarpa. Specifically, the topics examined in this thesis are: (1) the floral character traits of species, and the extent to which these traits can be associated with syndromes of bird or insect pollination; (2) the timing, frequency, duration, intensity, and synchrony of flowering of populations and individual trees; (3) the factors that may explain variation in flowering patterns of individual trees through examination of the relationships between flowering and tree-specific factors of individually marked trees; (4) the influence of tree size on the flowering patterns of individually marked trees, and (5) the spatial and temporal distribution of the floral resources of a dominant species, E. tricarpa. The results are discussed in relation to the evolutionary processes that may have lead to the flowering patterns, and the likely effects of these flowering patterns on blossom-feeding fauna of the Box-Ironbark region. Flowering observations were made for approximately 100 individually marked trees for each species (a total of 754 trees). The flower cover of each tree was assessed at a mean interval of 22 (+ 0.6) days for three years; 1997, 1998 and 1999. The seven species of eucalypt each had characteristic flowering seasons, the timing of which was similar each year. In particular, the timing of peak flowering intensity was consistent between years. Other spatial and temporal aspects of flowering patterns for each species, including the percentage of trees that flowered, frequency of flowering, intensity of flowering and duration of flowering, displayed significant variation between years, between forest stands (sites) and between individual trees within sites. All seven species displayed similar trends in flowering phenology over the study, such that 1997 was a relatively 'poor' flowering year, 1998 a 'good' year and 1999 an 'average' year in this study area. The floral character traits and flowering seasons of the seven Eucalyptus species suggest that each species has traits that can be broadly associated with particular pollinator types. Differences between species in floral traits were most apparent between 'summer' and 'winter' flowering species. Winter - flowering species displayed pollination syndromes associated with bird pollination and summer -flowering species displayed syndromes more associated with insect pollination. Winter - flowering E. tricarpa and E. leucoxylon flowers, for example, were significantly larger, and contained significantly greater volumes of nectar, than those of the summer flowering species, such as E. camaldulensis and E. melliodom. An examination of environmental and tree-specific factors was undertaken to investigate relationships between flowering patterns of individually marked trees of E. microcarpa and E. tricarpa and a range of measures that may influence the observed patterns. A positive association with tree-size was the most consistent explanatory variable for variation between trees in the frequency and intensity of flowering. Competition from near-neighbours, tree health and the number of shrubs within the canopy area were also explanatory variables. The relationship between tree size and flowering phenology was further examined by using the marked trees of all seven species, selected to represent five size-classes. Larger trees (≥40 cm DBH) flowered more frequently, more intensely, and for a greater duration than smaller trees. Larger trees provide more abundant floral resources than smaller trees because they have more flowers per unit area of canopy, they have larger canopies in which more flowers can be supported, and they provide a greater abundance of floral resources over the duration of the flowering season. Heterogeneity in the distribution of floral resources was further highlighted by the study of flowering patterns of E. tricarpa at several spatial and temporal scales. A total of approximately 5,500 trees of different size classes were sampled for flower cover along transects in major forest blocks at each of five sample dates. The abundance of flowers varied between forest blocks, between transects and among tree size - classes. Nectar volumes in flowers of E. tricarpa were sampled. The volume of nectar varied significantly among flowers, between trees, and between forest stands. Mean nectar volume per flower was similar on each sample date. The study of large numbers of individual trees for each of seven species was useful in obtaining quantitative data on flowering patterns of species' populations and individual trees. The timing of flowering for a species is likely to be a result of evolutionary selective forces tempered by environmental conditions. The seven species' populations showed a similar pattern in the frequency and intensity of flowering between years (e.g. 1998 was a 'good' year for most species) suggesting that there is some underlying environmental influence acting on these aspects of flowering. For individual trees, the timing of flowering may be influenced by tree-specific factors that affect the ability of each tree to access soil moisture and nutrients. In turn, local weather patterns, edaphic and biotic associations are likely to influence the available soil moisture. The relationships between the timing of flowering and environmental conditions are likely to be complex. There was no evidence that competition for pollinators has a strong selective influence on the timing of flowering. However, as there is year-round flowering in this community, particular types of pollinators may be differentiated along a temporal gradient (e.g. insects in summer, birds in winter). This type of differentiation may have resulted in the co-evolution of floral traits and pollinator types, with flowers displaying adaptations that match the morphologies and energy requirements of the most abundant pollinators in any particular season. Spatial variation in flowering patterns was evident at several levels. This is likely to occur because of variation in climate, weather patterns, soil types, degrees of disturbance and biotic associations, which vary across the Box-Ironbark region. There was no consistency among sites between years in flowering patterns suggesting that factors affecting flowering at this level are complex. Blossom-feeding animals are confronted with a highly spatially and temporally patchy resource. This patchiness has been increased with human exploitation of these forests leading to a much greater abundance of small trees and fewer large trees. Blossom-feeding birds are likely to respond to this variation in different ways, depending upon diet-breadth, mobility and morphological and behavioural characteristics. Future conservation of the blossom-feeding fauna of Box-Ironbark forests would benefit from the retention of a greater number of large trees, the protection and enhancement of existing remnants, and revegetation with key species, such as E. leucoxylon, E. microcarpa and E. tricarpa. The selective clearing of summer flowering species, which occur on the more fertile areas, may have negatively affected the year-round abundance and distribution of floral resources. The unpredictability of the spatial distribution of flowering patches within the region means that all remnants are likely to be important foraging areas in some years.

Anthropogenic utilization of forest and woodland ecosystems can cause declines in flora and fauna species. It is imperative to restore these ecosystems to mitigate further declines. In this thesis, I focused on a highly degraded region, the Box-Ironbark forests and woodlands of Victoria, Australia. Rather than mature stands with large trees, stands are currently dominated by high densities of small stems. This change has resulted in reduced populations of many flora and fauna species dependent on older-growth forests and woodlands. Managers are interested in restoring mature Box-Ironbark forests and woodlands through three alternative management strategies: allocating land to National Parks and allowing stands to develop naturally without harvesting, modifying timber harvesting regimes to retain more medium and large trees, or a new ecological thinning technique that retains target habitat trees and removes competing trees to encourage growth of retained stems. / The effects of each management strategy are not easy to predict due to complex interactions between intervention and stochastic natural processes. Forest simulation models are often employed to overcome this problem. I constructed state-and-transition simulation models (STSMs) to predict the effects of alternative management actions and natural disturbances on vegetation structure. Due to a lack of empirical data, I relied on the knowledge of experts in Box-Ironbark ecology and management to construct STSMs. Models predicted that the development of mature woodlands under all strategies was minimal over the next 150 years, and neither current harvesting nor ecological thinning is likely to expedite the development of mature stands relative to growth and natural disturbances. However, differences in experts’ opinions led to widely diverging model predictions. / Uncertainty must be acknowledged in model construction because it can affect model predictions. I quantified uncertainty due to four sources – between-expert variation, imperfect expert knowledge, natural stochasticity, and model parameterization – to determine which source caused the most variance in model predictions. I found that models were very uncertain and between-expert uncertainty contributed the majority of variance in model predictions. This brings into question the use of consensus methods in forest management where differences between experts are ignored. / Using uncertain model predictions to make management decisions is problematic because any given action can have many plausible outcomes. I applied several decision criteria to uncertain STSM predictions using a formal decision-making framework to determine the optimal management action in Box-Ironbark forests and woodlands. I found that natural development is the most risk-averse option, while ecological thinning is the most risky option because there is a small likelihood that it will greatly expedite the development of mature woodlands. Rather than selecting one option, managers could rely on a risk-spreading approach where the majority of land is allocated to no-cutting National Parks and a small amount of land is allocated to the other two harvesting strategies. This would allow managers to collect monitoring data for all management strategies in order to learn about effects of harvesting and update model predictions through time using adaptive management.

Many researchers have highlighted the dire prospects for biodiversity in fragmented agricultural landscapes and stressed the need for increasing the area of, and connectivity between, natural ecosystems. Some have advocated the use of naturally regenerating forest ecosystems for sequestering atmospheric carbon, with opportunities for dual restoration and carbon benefits. However, no studies have explicitly explored the feasibility of obtaining such dual benefits from a regenerating woody ecosystem. This thesis aims to provide a detailed assessment of the restoration and carbon potential of Brigalow regrowth, an extensive naturally regenerating ecosystem throughout the pastoral regions of north eastern Australia. It combines observational, experimental and modelling techniques to describe the agricultural legacy of pastoral development, identify constraints to restoration and explore methods to remove these constraints. A review of existing ecological knowledge of Brigalow ecosystems is provided in chapter 3, along with discussion of policy and socio-economic issues that are likely to influence how and to what extent regrowth is utilised for restoration and carbon purposes in the Brigalow Belt. The review found restoring regrowth is likely to have benefits for a wide range of native flora and fauna, including the endangered bridled nailtail wallaby. Knowledge gaps exist relating to the landscape ecology of Brigalow regrowth and the impacts of management and climate change on carbon and restoration potential. Also, a conflict exists between short-term carbon sequestration and long-term restoration goals. Regional demand for high biomass regrowth as a carbon offset is likely to be high but ambiguities in carbon policy threaten to diminish the use of natural regrowth for reforestation projects. A large cross-sectional study of regrowth is presented in chapter 4. Data were analysed using multi-level / hierarchical Bayesian models (HBMs). Firstly, we found that repeated attempts at clearing Brigalow regrowth increases stem densities and densities remain high over the long term, particularly in high rainfall areas and on clay soils with deep gilgais. Secondly, higher density stands have slower biomass accumulation and structural development in the long term. Spatial extrapolations of the HBMs indicated that the central and eastern parts of the study region are most environmentally suitability for biomass accumulation, however these may not correspond to the areas that historically supported the highest biomass Brigalow forests. We conclude that carbon and restoration goals are largely congruent within regions of similar climate. At the regional scale however, spatial prioritisation of restoration and carbon projects may only be aligned in areas with higher carbon potential. Given the importance of stem density in determining restoration and carbon potential, an experimental thinning trial was established in dense Brigalow regrowth in southern Queensland (chapter 5). Four treatments were applied in a randomised block design and growth and mortality of a subset of stems was monitored for two years. Data were analysed using mixed-effects models and HBMs and the latter were subsequently used to parameterise an individual-based simulation model of stand structural development and biomass accumulation over 50 years. The main findings of this study were that growth and mortality of stems is influenced by the amount of space available to each stem (a neighbourhood effect) and that thinning accelerates structural development and increases woody species diversity. The examination of neighbourhood effects is taken further by considering drought-related mortality in a Eucalyptus savanna ecosystem (chapter 6). For this work a multi-faceted approach was employed including spatial pattern analyses and statistical models of stem survival to test three competing hypotheses relating to neighbourhood effects on drought related tree mortality. The main finding of this study was that neighbour density and microsite effects both influence drought-related mortality and the observed patterns can readily be explained by an interaction between these two factors. As a whole, this thesis contributes the following scientific insights: (1) restoration and carbon goals may be aligned for naturally regenerating woody ecosystems, but the degree of goal congruence will vary across the landscape in question, (2) while some woody ecosystems retain an excellent capacity to regenerate naturally, the agricultural legacy may still have long term effects on restoration and carbon potential, (3) neighbourhood effects that operate at the stem scale strongly influence dynamics at the ecosystem scale.