School of Ecosystem and Forest Sciences - Theses
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Regime resistance and accommodation in sustainable energy transitions
To help accelerate the decarbonisation of electricity generation and meaningfully mitigate climate change, a more nuanced understanding of the power and influence of incumbent electricity firms in government policymaking is required. Building on sustainability transitions literature, concepts from neo-Gramscian political economy and insights from strategic management, this study investigated the ways in which incumbent electricity generators, network operators, retailers, and their industry organisations sought to influence residential solar power policy. The single case study is focused on Victoria, Australia, using data obtained from multiple sources, including: legislation; policy submissions, reports and documents; organisational documents and materials; media releases and articles; and transcripts of in-depth, semi-structured interviews with key informants from electricity utilities and their industry peak bodies. The study examined federal and state government policy instruments that supported the installation of residential solar power before it explored attempts by incumbent electricity firms and their industry organisations to influence the development of Victorian feed-in tariffs, the State Government’s primary policy instrument. The research found that incumbent actors contributed to the (re)production of a socio-technical regime by drawing on material, institutional and discursive forms of power to execute strategies of resistance and accommodation. Incumbent actors resisted feed-in tariffs by discussing them in negative terms, building alliances with opposed civil society organisations, lobbying policymakers and reminding them of their mutually dependent relationship. However, incumbent actors accommodated feed-in tariffs by working with government to implement its policy initiatives. Incumbent firms accommodated renewables more generally by operating renewable energy business units and joining renewable energy industry organisations, although the later served to moderate support for feed-in tariffs. The study also found that a number of external and internal factors shaped the strategic approach of incumbent electricity firms and limited the extent of their resistance. High public support for residential solar power and strong government determination to implement feed-in tariffs were key external factors that discouraged greater resistance from incumbent firms. Key internal factors included the assessment of feed-in tariffs as low risk to the ongoing profitability of incumbent firms, and higher policy priorities, the achievement of which necessitated good relations with government and the sacrifice of lower policy priorities. These findings suggest that a neo-Gramscian approach to understanding politics and power in sustainable energy transitions offers useful insights for policymakers, incumbent and renewable energy firms, and civil society organisations who seek to accelerate the decarbonisation of electricity generation.
Using LiDAR for landscape-scale mapping of potential habitat for the critically endangered Leadbeater's Possum
Leadbeater’s possum (LBP) (Gymnobelideus leadbeateri) is a critically endangered arboreal marsupial located in Victoria’s Central Highlands in southeastern Australia. Populations of hollow-bearing trees, the key nesting habitat resource for Leadbeater’s Possum, are rapidly declining. Disturbances such as bushfires and timber harvesting are believed to be the major causes in the reduction of hollow-bearing trees and thus the extent, quality, and connectivity of LBP habitat. LBP also requires a connected understorey or midstorey for foraging and movement within the forest. Changes in LBP habitat at the landscape-scale are interpolated from plot-level measures and assumed distribution of habitat resources based on forest type and disturbance history. Long-term monitoring at the plot-level has reduced uncertainty around habitat resources required by LBP within the montane ash (Eucalyptus regnans, E. delegatensis, E. nitens) forests of the region; however, it has not addressed uncertainty around the landscape distribution of nesting and foraging habitat resources. Developing effective conservation strategies for LBP requires an understanding of the distribution and quality of available habitat across its potential range. This currently does not exist. I used Light Detection and Ranging (LiDAR) data to develop empirical models that quantify the distribution, quality, and connectivity of LBP habitat in the montane ash forests within a multi-scale framework. LiDAR-derived topographic and vegetation structural metrics were integrated with different habitat analysis approaches to model the distribution of critical LBP habitat features (hollow-bearing trees, foraging and midstorey stratum connectivity) at multiple scales across ~340,000 ha of forest in Victoria’s Central Highlands. In Chapter 2, tree-level individual tree delineation (ITD) algorithms were developed to identify individual trees and analyse tree crown attributes. ITD algorithms for overstorey trees were applied to the whole study area to estimate the abundance and distributions of large live trees with big crowns that have high probability of containing hollows. Based on field surveys in 1939 regrowth mountain ash, the dominant overstorey crowns identified by the ITD algorithms had an 86% success rate in identifying mapped trees in mature forest plots. Our landscape analyses estimated 405,000 large live trees (crown width >15m) and 572,000 (95%CI: 318,000-808,000) potential large live tree with hollows (DBH>150cm), which equate to a mean density of one live large old tree per ha across the Central Highlands. ITD algorithms were also used to map canopies and to calculate the proportion of projection area of midstorey stratum, a surrogate for midstorey coverage, for use in assessments of foraging habitat. Hollow-bearing trees are a key feature of mature forests around the world. They provide critical habitat resources for hollow-dependent animals, many of which are threatened, due to a range of natural and human disturbances. In Chapter 3, to develop more accurate inventories of hollow-bearing trees (both live and dead form classes) across a large landscape, I used LiDAR-based metrics of forest structure and topography, coupled with datasets on environmental conditions to develop statistical models of abundance of hollow-bearing trees using machine learning tools. These provided empirical estimates of HBT density for every hectare of the Central Highlands within the LiDAR footprint. I identified 1,519,000 (95%CI: 1,306,000 – 1,730,000) hollow-bearing trees may occur across the full extent of Victoria’s Central Highlands. This included 833,000 (95%CI: 742,000-923,000) live HBTs and 686,000 (95%CI: 564,000-807,000) dead HBTs. The predictive models provided rigorous, repeatable estimates of tree abundance across a wide range of vegetation classes and forest management zones (with appropriate estimates of uncertainty), as well as a new understanding of the complexity of the structural, topographic, and environmental features associated with abundance of hollow-bearing trees and their spatial variability over large areas. In addition to HBTs for nesting, LBP require a well-connected midstorey stratum to facilitate movement amongst nesting trees and foraging. In Chapter 4, the association between field-assessed multi-storey vegetation connectivity and LiDAR structural metrics was evaluated to develop landscape-scale predictions of connectivity and foraging habitat (density of live wattle) for LBP. Forest structural types with high midstorey connectivity, which are a critical habitat feature for LBP, occupied 17.2% of the study landscape. The landscape-scale predictions of vegetation connectivity provided an understanding of the factors (e.g., topography, fire, logging and interactions among strata) that help shape connectivity and foraging habitat. This understanding is critical for improving the management of the region’s forests and, in particular, ensuring that both key habitat elements, HBTs and dense midstorey vegetation, either co-occur in the same stand or are in close proximity to ensure sufficient high-quality habitat for LBP over space and time. In Chapter 5 I used the landscape-level mapping of habitat resources developed in Chapters 2-4 to predict landscape-scale LBP habitat suitability based on fields records of LBP occurrence. The validated habitat suitability model for LBP had an overall accuracy of 87.3% and an AUC of 0.889. The model predicted that 28,000 ha (95%CI: 17,000ha-40,000ha) of the study landscape (i.e., 9.6%) supported suitable habitat for LBP. Twelve variables were found to have the strongest influence on estimating habitat suitability for LBP. Of these, four related to foraging habitat, three to nesting habitat, three to climatic and topographic factors, and two to disturbance history. Our approach highlights the potential for using high-resolution, spatially explicit data on forest structure at a landscape-scale to map the distribution and abundance of suitable habitat for a critically endangered species. Habitat suitability models generally account for the selection of suitable habitat and their appropriate geographical extents. However, they rarely account for the accessibility of this habitat and connectivity among habitat patches. Effective conservation of species requires that patches of their habitat are connected in space and time. In Chapter 6, graph-theoretic connectivity networks based on resistance surfaces were generated from the LiDAR mapped cover of vertical stratum. The least-cost links between patches from networks were integrated with a kernel density estimator to identify functionally connected regions. The mapping of landscape connectedness identified a potential landscape-scale metapopulation structure for LBP within the Central Highlands. This provides a tool that could be used to expand the existing protected area network to support the metapopulation processes of LBP. The identification of functionally connected regions could enhance the conservation planning for long-term population persistence. The conservation and management of endangered species is major focus of forest and land managers around the world. High-quality data on the distribution and quality of habitat is critical to the development of effective conservation and recovery strategies for species. This thesis developed a set of habitat analysis models for quantifying habitat resources, habitat suitability and connectedness for the critically endangered Leadbeater’s Possum across a range of spatial scales. These analyses highlight the benefits of using high-resolution, spatially explicit LiDAR data from the whole landscape to identify and map (1) the distribution and abundance of hollow-bearing trees, (2) midstorey stratum connectivity, (3) the distribution and abundance of suitable habitat, and (4) metapopulation structures within functionally connected regions. Dynamic modelling, informed by these LiDAR-derived models, can be used to forecast the likely consequences of changing habitat abundance under different management scenarios and evaluate both the short-term and long-term effectiveness of different conservation strategies.
Aspects of the ecology of Astelia australiana, Tall Astelia
The conservation of threatened species requires an understanding of their ecology to identify the factors that are contributing to their decline and to assess their extinction risk. The objective of this thesis was to examine the ecology of Astelia australiana, a threatened herb endemic to Victoria, Australia to inform its conservation management. To achieve this objective, I examined how the distribution and abundance of A. australiana has changed over its entire known range in the 20- year period since its demographic monitoring program began. I also examined its habitat niche and regeneration ecology and response to potential threats including drought, wildfire, disease and low light availability. I then used this new understanding to develop a population viability model to explore the viability of A. australiana to current conditions, threats and limitations. The key findings of my research include a better understanding of the regeneration ecology of A. australiana including that it involves both sexual and asexual reproduction which occur at the same time. This reproduction occurs once in an individual’s lifetime and results in the production of flowers and fruit and three clones. Previously A. australiana had been variously described as dioecious and gynodioecious; however, it is trioecious, having male, female and hermaphrodite flowers on different plants and male and hermaphrodite flowers on the same plants. Various native bird and mammal species are involved in the regeneration ecology of A. australiana as pollinators and dispersers of fruit. Reproduction of A. australiana requires a minimum amount of available light from canopy gaps in the rainforest understorey to occur. A. australiana has declined in abundance by 57% over the 20-year period. Herbivory, and disease have contributed to mortality of A. australiana over this period but do not appear to be the only factors contributing to the observed decline. Although, A. australiana was found to be highly sensitive to drought, multiple lines of evidence including: the pattern of decline and recruitment of A. australiana; comparison of the physiological response of A. australiana to drought in an experimental trial with physiology of individuals in the rainforest; examination of the microclimate of the sites; and assessment of the growth response using dendrochronological techniques, of the dominant tree in the cool temperate rainforest, Nothofagus cunninghamii (Myrtle Beech) to drought periods, all indicate that drought is not driving the observed decline as had been previously concluded by others. Wildfire also resulted in a loss of individuals in one site that was burnt during the study period however, decline from this site was not included in the 57% change in abundance total. Changes in forest structure due to growth in the canopy reduce the light availability in the understory which limits A. australiana recruitment and increases mortality and thus contributes to the species decline. Population viability analysis of A. australiana, predicted that the species populations are at a high risk of large decline (86%) across the species range over the next 50 years under current climate. This research is significant for the conservation management of A. australiana as it highlights new threats to the species and removes some others that were thought to be key threats. It is also significant because the magnitude of the observed decline warrants an up listing of the conservation status of A. australiana from Vulnerable to Endangered at the State and National level. This research has already been used to inform conservation management of the species populations including: fencing, to prevent further decline in one site that had been heavily browsed and disturbed by an introduced herbivore; facilitating targeted surveys for additional sites; increasing its reproductive rate within four sites through manipulation of the light environment; establishing new populations through translocation of individuals; identifying suitable sites for further translocation; establishing the monitoring program at new sites; reviewing the ex situ storage of seed; guiding and prioritising future conservation management actions of A. australiana through modelling and expert advice; and raising the species profile to obtain funding for further research and management of the species.
Between localism and internationalism: multi-level governance, land use decision-making, and REDD+ in Central Kalimantan province, Indonesia
While there has been considerable progress in developing global environmental policies and goals, implementing these at a local level has proven extremely challenging. This study explores how internationally formulated objectives for Reducing Emissions from Deforestation and Forest Degradation (REDD+) are implemented in frontier landscapes undergoing rapid transition. I focus on subnational implementation of REDD+ in relation to land use decision-making in Indonesia. International donors and agencies have actively promoted REDD+ to reduce carbon emissions, improve forest management and broaden livelihood options in developing countries. Indonesia has been a priority country for REDD+ due to high rates of deforestation and continuing land use change. In this thesis, I use multi-level governance (MLG) concepts to investigate how internationally formulated objectives for REDD+ interact with existing systems of land use governance at multiple levels. I incorporate actor-network theory (ANT) within an overall practice orientation to address the following questions: (i) how are land use decisions made, (ii) how is REDD+ being implemented within interacting multi-level governance arrangements, and (iii) how do decision-making processes in these arrangements relate to multiple outcomes? These questions are underscored by a concern with social and environmental justice, and related to this, a concern with ethics in global environmental governance and novel mechanisms such as REDD+. In 2010, Central Kalimantan was selected as the official REDD+ priority province of Indonesia. The province has had a long and complicated history of governance interventions and is currently undergoing rapid land use changes, such as the establishment of large industrial oil palm plantations. I used a case study approach, centring on the southern tropical peatlands of Central Kalimantan, where a variety of REDD+ projects have been implemented. Data was collected during 10 months of field research conducted between 2013 and 2015 and combined ethnographic techniques with document analysis, interviews and field observations. Data collection spread across government levels, actors, and locations. A total of 194 interviews and other field observations and documents were used to explore concepts of negotiation, translation, and learning. The findings show how successive interventions and negotiations over land have transformed the remote forested peat ecosystems into a complex, and contested, mix of land uses. Competing interests and interactions among land use changes are diminishing access to land for villagers and other local people. Processes to translate internationally formulated objectives through multiple layers of governance were difficult to control and lacked flexibility. REDD+ objectives were reinterpreted at each level, and discrepancies and disconnects emerged between higher level objectives and local realities. While learning was mentioned in REDD+ project documents, a reliance on experts, and an emphasis on training and livelihoods to villagers, allowed limited local input into project design. Due to competition for land, REDD+ becomes another powerful force for villagers to contend with. Rather than achieving any desired new status quo, attempts to implement pre-determined plans become entangled in these fluid and messy frontier processes. Overall, this thesis contributes to understanding how interventions for implementing global environmental goals relate to pre-existing, dynamic and emerging land use systems that I conceptualise as ‘frontiers’. It adds new insights into how negotiation, translation, and learning are linked in interactive networked processes and identifies implications for pursuing environmental reforms. I develop a new concept, the entry point, to describe political opportunities and limitations for how those implementing, and being subjected to, global environmental policies can navigate local land allocation systems. The thesis demonstrates the need for social learning in international environmental governance and suggests strategies to move beyond policy design and technical coordination to encourage long-term collaboration and integration of learning at all levels. A focus of REDD+ is carbon rich regions in developing countries that are undergoing rapid changes. Findings from this thesis have implications beyond Indonesia to frontier landscapes in other developing countries where there is competition between large-scale forest conservation and land conversion for local or national economic development.
Impacts of short-interval wildfires on tree demography and forest structure in temperate Australia
Fire is a powerful agent of disturbance in terrestrial ecosystems, and it shapes vegetation composition and patterns globally. This is particularly true in south-eastern Australia where forests are dominated by species of the genus Eucalyptus, many of which have the capacity to recover from high-severity fire by resprouting from epicormic or basal buds. Climate change is predicted to yield more severe fire weather and lengthen fire seasons in temperate Australia, leading to increased wildfire frequency in these forests. While increased fire frequency – resulting in wildfire intervals of under a decade – are known to negatively affect fire-sensitive eucalypts (obligate seeders which have a juvenile period of 10 – 15 years) less is known about how such changes impact fire-tolerant, resprouting eucalypts. This Thesis examines the impacts of recent wildfires in south-eastern Australia, where a series of large wildfires burned over four million hectares of land, leading to the burning of different types of fire-tolerant eucalypt forests (basal resprouters, epicormic resprouters) by high-severity wildfires once, twice, and sometimes three times between 2003 and 2013. In the context of this massive natural experiment in the landscape, my overarching aim was to quantify the impacts of short-interval wildfire on eucalypt tree demography and regeneration, to improve understanding of potential fire-related changes to the structure and resilience of fire-tolerant forests. Short-interval high-severity wildfires significantly increased whole-tree mortality and decreased the abundance of both resprouts and seedlings in basally resprouting eucalypt forests. In these sub-alpine forests, dominated by snow gum (E. pauciflora), more frequent wildfire (two and particularly three short-interval fires) also increased the cover of grasses at the expense of shrubs. In mixed-species eucalypt forests, which occupy extensive tracts of low elevation landscapes in south-eastern Australia, resprouting occurs from both basal and epicormic buds. In these forests, the dynamics of both topkill (i.e. stem, but not whole-tree, mortality) and whole-tree mortality have important ramifications for forest structure. After a single high-severity wildfire, small-diameter stems were typically topkilled; after two short-interval wildfires, the diameter of stems topkilled increased. Additionally, the overall likelihood of either basal or epicormic resprouting decreased after two short-interval wildfires. This decline in resprouting capacity indicated that the size class most vulnerable to ‘resprout failure’ after multiple wildfires was intermediate sized stems (in the vicinity of 20 - 30 cm DBHUB), rather than smaller or larger stems. Seedling regeneration also decreased in these forest types after short-interval wildfires, suggesting that, as for fire-sensitive forests, immaturity risk may be a relevant factor for fire-tolerant forests. Short-interval wildfires reduced the total and aboveground carbon stocks of mixed-species forests, while also increasing the proportion of carbon mass in the dead pool, indicating that resprouter forests might not be perpetually secure carbon stocks under emerging fire regimes. My Thesis highlights that fire-tolerant forests may not be as invulnerable to changes in fire frequency as widely assumed, and that management interventions will likely be required to counteract increasing tree mortality and decreasing tree regeneration if predictions of more frequent and severe wildfires in temperate Australia are realised.
The impact of wood characteristics and properties on the market price of plantation grown teak logs (Tectona grandis Linn.F) in Laos
Teak is considered one of the highest value hardwood species. The Lao wood processing industry has been using teak wood harvested from different ages, ranging from 10 to 35 years, with broadly different prices. However, the current grading rules used for teak logs are unsuitable for development of a fair and transparent pricing market. To date, there has been no regulation of market requirements for wood characteristics and/or wood properties. The key terms of wood characteristics and wood properties used in this study were defined as: a) wood characteristics refer to the visual appearance or features of logs and square logs; for example, heartwood proportion, wood colour, presence of knots, bend, pipe, buttress, decay, end checks and splits; b) wood physical and mechanical properties describe the quantitative characteristics of wood and its behaviour in response to external influences and externally applied forces; for example, wood density, dimensional stability, stiffness, bending strength, compression strength, shear strength, and hardness. The overall aim of this study was to identify the key characteristics and properties of wood that affect the selling price of teak logs grown in Laos. More specifically the objectives of the study were to: 1. define the characteristics and properties of plantation grown teak logs that are valued by wood processors and manufacturers; 2. understand the influences of wood characteristics and properties on the current selling price of plantation grown teak logs used in Lao wood processing and manufacturing industry, and; 3. determine the key characteristics and properties that could be used if a log grading system was to be applied in Laos, and that is based on the results identified from the buyers’ requirements. Both semi-structured interviews of teak buyers and qualitative and quantitative data of wood characteristics and properties of plantation grown teak in Laos that were valued by teak buyers were obtained and analyzed. The results showed that while the harvesting age of plantation teak was reported to be between 10-35-year-old, the dominant harvesting age was between 21-25 years. Three types of teak timber were commonly sold in Laos: 24.7% as standing trees, 39.6% as round logs and 35.7% as square logs. From the marketing supply chain it was found that the end buyers of teak timber were wood sawmillers and wood products manufacturers. The local traders (middlemen) who purchased teak logs from the growers and in turn supply to the end buyers account for 49% of sales. A further 41% was sourced directly from teak growers and the remaining 10% was sourced via local small wood sawmillers and manufacturers. The main wood products produced from plantation teak included joinery products, door and window frames, and furniture (57.9%, 26.3% and 15.8%, respectively). Currently, 83% of teak products produced were consumed on the domestic market. The remaining account for 17% were destined for international markets. The prices of teak showed to be significantly different based on timber types, log sizes, and distance from plantation site to the road in the case of standing trees. However, the price variation was not based on the factors of wood quality and properties. A round log price of US$250/m3, based on both log diameter size (D26-30 cm) and log quality grade (A grade), was much lower than the US$321/m3 amount paid for an ungraded round log of the same size based on log diameter alone. It can be concluded that there is a need to develop a clear system or standard for teak log grading and pricing in Laos. The knot was found to be the major characteristic appearing on all teak wood products, square and round logs. White sapwood, end splits, and buttress were the next most predominant characteristics in teak wood products, square and round logs, respectively. Buyers were found to prefer a larger heartwood proportion and diameter of the logs. On the other hand, decay, pipe, insect holes, bend, and knot holes were found to be the wood defects that the buyers were most concerned about when they bought timbers. Log diameter, bend, buttress and insect holes were only the wood characteristics that have significant impact on log price. Buyers were unwilling to buy any logs containing wood decay or rot. Wood property values were not important for the buyers. Lao teak buyers didn’t offer a different price for teak timbers that have different wood property values. This study concluded that wood properties have no impact on log price. If a grading system is to work, then it must be of use to those who buy logs and square logs, as they would be the main beneficiaries of such a system. Laos already uses a grading scheme; however, the question remains about how well it works. A good grading system can be said to work if the different characteristics; for example, wood characteristics valued by buyers are reflected in the prices for the wood. In a simple test, the following questions could be asked: 1. Is there a significant difference between the prices of graded logs? 2. Do the prices for each grade move independently of one another (thus reflecting the changing nature of how buyers value each characteristic)? and 3. Are the prices of graded logs higher than ungraded logs (at least those that are considered to be of a superior quality). The answers to these questions, when applied to the current grading system, led to the conclusion that such system did not work. All prices tended to move together, and ungraded log prices were found to be greater than graded logs. The next question to address was how a better grading system could be determined. Clearly, any system should reflect the wood characteristics that buyers demand. To that end, it was necessary to survey buyers to determine what they considered to be the most valuable characteristics. These identified characteristics need to be collated into a system that is both clear and usable for the buyers. The following characteristics were found to be important: log diameter, heartwood proportion, presence of insect holes, log bend, presence of knots, end splits, wane/buttress, and wood colour. It should be noted that as the buyers' responses are not universal (for example, some buyers prefer one characteristic, whereas other buyers prefer another), a grading system is preferred. The grading system reflects a set of tolerances within which a range of wood characteristics can be valued by the buyer. In this way, they can pay a price for logs according the characteristics they value and not on the basis of log diameter alone (as they do now). Based on the key finding from the results mentioned above, a guideline for a teak timber grading system was developed. This grading system is a new log grading method, which can be used in Laos and similar developing countries. Implementation of this grading system would benefit both teak buyers and growers. The buyers can minimise timber waste, as buyers can buy only the qualities of timber they want if the timber is graded. Teak growers could also employ practices that result in them growing trees that are more highly valued by buyers because they will have the characteristics buyers pay a premium for. In the long-run, growers would minimise the production of inferior quality logs. The growers will supply only the timber that is required by the industry for specified uses. What should be noted is that in a market where the products are ungraded, average prices for all qualities apply. Thus, it is in the interests of growers to supply the lowest quality product, as this is the one that costs the least to produce and for which they receive the average price as a superior product. On the other hand, a grower who produces a superior product receives the same average price as that of the grower of an inferior product. In a graded product market, the grower receives the price the buyer is willing to pay for the characteristics that are valued by the manufacturers of finished products. Thus, growers are rewarded for improving the quality of the logs that will result in an improved industry.
In defence of flowers: The ecology and biochemistry of a chemical defence, cyanogenesis, in Proteaceae
As sessile organisms, plants produce a remarkable diversity of chemicals to defend their tissues against herbivory. The large body of research into plant defences has primarily focused on plant-herbivore interactions and foliar defences, whereas floral chemical defences have received little attention. This is despite the fact that flowers of many species are known to contain defence metabolites, the sizeable investment of resources in floral structures, the significance of flowers to fitness, and the importance of floral chemistry for both florivores and pollinators. This thesis investigates patterns in floral chemical defence in the Proteaceae, a dominant plant family in Australia characterised by diverse floral traits and a high frequency of floral cyanogenesis - a constitutive nitrogen-based chemical defence involving the release of toxic hydrogen cyanide from endogenous cyanogenic glycosides (CNglycs) upon tissue damage. There were two main aspects to this research, (1) investigations into patterns in the distribution of CNglycs within flowers of different species, and (2) investigations into patterns in whole-plant chemical defence using Telopea speciosissima. In Chapters 2 and 3 I quantified CNglycs in dissected floral tissues and used a new MALDI-MS imaging method to investigate and compare fine-scale patterns in the distribution of CNglycs within flowers of eight congeneric Lomatia species (Chapter 2), and 11 species from different genera from across the family phylogeny (Chapter 3). Together these studies identified: (1) variation in floral CNglyc concentrations over three orders of magnitude across species, (2) unprecedented concentrations of CNglycs in some floral tissues (e.g. Lomatia stigmatic cells and pollen were 11% CNglyc by mass), (3) highly tissue-specific distributions of CNglycs within flowers, (4) substantial interspecific differences in patterns of within-flower CNglyc distribution, and (5) that differences in floral CNglycs across species were not correlated with other traits (e.g. colour, disturbance response) or phylogenetic position. Findings demonstrate the power of MALDI-MSI to reveal dynamic and fine-scale spatial patterns in the distribution of different CNglycs. Data are discussed in relation to defence theories and potential multiple functions of CNglycs in plants. In Chapter 4 I used T. speciosissima, which has CNglycs in all tissues, to compare floral and foliar CNglycs in seven populations from three distinct genetic groups, which also differ in environmental conditions. There was no consistent relationship between floral and foliar cyanogenic capacity within populations; however, mean floral and foliar CNglyc concentrations differed between genetic groups. Correlations between CNglycs and δ13C, N and physical leaf traits suggest these differences reflect both genetic and environmental effects. In Chapter 5, a glasshouse study investigating the effects of N and P supply on above- and below-ground CNglycs in T. speciosissima seedlings identified that root and shoot chemistry responded differently to N and P treatments. The higher CNglyc concentrations in roots may reflect greater importance of root chemical defence in a non-mycorrhizal species that relies on basal resprouting following fire. Together this research provides evidence that foliar chemistry cannot be assumed to reflect whole plant chemistry (floral, root) and that both whole plant and fine-scale tissue-specific studies are required to understand the regulation and roles of CNglycs in plants.
The ecology and evolution of plant-pollinator interactions in Australian Typhonium (Araceae)
Pollination is one of the most important interactions in nature, with the vast majority of flowering plants reliant upon biotic vectors for out-crossing and reproduction. Despite this, the pollination systems of most plants are unknown, limiting our understanding of the evolutionary processes that gave rise to these interactions, dependency on specific pollinators for plant reproduction, and the resilience of interactions in the context of environmental change. The species-rich family Araceae is used worldwide as a model to investigate the ecology and evolution of plant-pollinator interactions, yet detailed studies of pollination in the majority of genera remain scarce. One such sizeable genus is Typhonium Schott, the most diverse genus of Araceae in Australia. Using field studies and a range of analyses (morphological, chemical, molecular), this thesis aimed to investigate the diversity, ecology, and evolution of Typhonium plant-pollinator interactions in Australia. This thesis is divided into four research chapters. Chapter 2 identified significant intra and interspecific variation in floral traits and pollinating insects between allopatric populations of the only species with a broad geographic range, Typhonium brownii, and its geographically restricted sister species T. eliosurum. Both mimic dung, but they are functionally specialised to deceive saprophagous beetle or fly pollinators, respectively. Interspecific trait divergence provided insight into the pollinator-mediated selection of floral traits and the importance of morphological filters. Significant intraspecific variation in T. brownii thermogenesis, anthesis rhythms and scent indicates the occurrence of a T. brownii species complex, comprised of more taxa than currently recognised. The function of floral scent for pollinator attraction was confirmed in field bioassays and is discussed in the context of adaptation to the pre-existing biases of pollinators. Chapter 3 investigated pollination systems of tropical T. angustilobum and T. wilbertii, from north Queensland. T. angustilobum and T. wilbertii were similarly identified as brood-site mimics, share similar thermogenic and anthesis rhythms, but are functionally specialised to beetle and fly pollination, respectively, associated with distinct floral scents and morphological filters. Molecular analyses provide the first evidence of heating via the alternative oxidase (AOX) in Typhonium. Chapter 4 combined pollination data on 14 Typhonium species from across their geographic range to explore trends in plant-pollinator interactions. It presents further evidence for functional specialisation to pollination by saprophagous beetle and fly families in a genus characterised by rewardless brood-site mimicry, floral traps and a high frequency of floral thermogenesis. Differences in pollinator type and climatic region were not related to anthesis rhythms or thermogenic properties. Despite significant scent divergence, some broad differences in scent with pollinator type were identified. Given the prevalence of Staphylinidae-flower interactions in Australian Typhonium, Chapter 5 systematically analysed the worldwide extent of rove beetle floral interactions and pollination. At least 108 staphylinid-plant species interactions were identified across 27 seed plant families, the first assessment of the diverse and complex relationships between staphylinids and plants which were previously overlooked. As concluded in Chapter 6, this thesis develops Typhonium as a new model system to investigate the ecology and evolution of brood-site mimicry, an underappreciated yet widespread form of floral mimicry.
Improving degraded and compacted urban soils to support tree establishment and growth
The creation and expansion of human habitats alters the natural properties of urban soils, not only through direct anthropogenic soil disturbance, but also indirectly through impacts from the anthroposphere. This creates soils that are novel and different to those found in non-urbanised settings. These urban soils are often compacted and degraded and not be suitable for the establishment and growth of deep-rooted tree species that make urban habitats liveable. In this thesis, I have studied methods to rehabilitate compacted and degraded urban soil to improve the successful establishment and growth of trees planted into those soils by improving key physical and biological properties through tillage and the incorporation of OM amendments. My objectives for the study were: 1. To investigate the effects that the incorporation of organic amendments and tillage have on improving the soil properties of degraded and compacted urban soils and their ability to support improved tree performance. 2. To develop an understanding of the interaction between organic amendments and soil type when improving degraded urban soils. 3. To understand the interaction between tree water strategies in determining tree growth response and levels of water stress in degraded urban soils amended with different organic amendments. For the first objective, I found that the use of a ‘scoop and dump’ tillage technique, with the incorporation of organic matter (compost and/or biochar), was effective at improving key physical and biological properties of compacted and degraded urban soils. However, the successful growth of trees in remediated soils was partly determined by the initial site conditions found at each site, and improved tree growth was not always apparent. In both sandy and clay soils, improvements to physical and biological properties, achieved with the use of tillage, were maintained for longer, when organic matter was also incorporated into soils during the tillage. The addition of organic matter improved the biological activity of tilled sand and clay soils, and this may have been a result of improved soil water availability. In addition, I found that there were very few differences in soil and tree growth outcomes between the compost and biochar used in either soil type. Although the incorporation of biochar was effective in the remediation of urban soils, it was no more effective than the incorporation of compost or of compost and biochar combinations. There were no synergistic tree growth benefits from using a combination of compost and biochar. The second objective of this project was to investigate the interaction between organic amendments and soil types when improving degraded urban soils. The water holding properties of sandy soils were improved with the addition of organic matter where the plant available water content was increased in both a greenhouse pot study and a field study. Alternatively, the plant available water content was decreased in a clay soil amended with organic matter in a greenhouse experiment and unchanged in the clay soils of a field experiment. Although the water characteristics were not changed in the clay soil, tree growth was nevertheless improved due to improvements in other soil hydrological properties achieved with the addition of organic matter. These changes allowed for faster infiltration and decreased the bulk density of the clay soils. The third objective of this thesis evaluated the interaction between tree water use strategies, soil types and the incorporation of different types of organic matter. Compared to an unamended control, a xeric tree species accumulated more biomass when grown in a sandy soil amended with compost, biochar and a combination of compost and biochar, under both well-watered and water deficit irrigation regimes. There were no biomass changes in the clay soil. There were no differences measured in the growth of the mesic species in either the sand or clay soil or with the compost, biochar or the combination of compost and biochar. This thesis suggests that while urban soils can be successfully remediated with the application of tillage, the incorporation of organic matter will prolong the benefits achieved by the tillage. However, successful remediation of compacted and degraded urban soils will ultimately depend on applying solutions based on an understanding of existing soil properties and the tree resource acquisition strategies required for a particular site. Management practices utilising the incorporation of organic matter should assess the cost/benefits of different types of organic matter amendments as the outcomes of using different organic matter types may not differ.
The role of plants in green roof rainfall retention
Increasing urbanisation and associated impervious surfaces have negative impacts on humans and the environment. Therefore, more resilient and sustainable stormwater management that mimics the natural predevelopment hydrology cycle to protect urban water sources and reduce urban flooding is required. Contributing to this approach, green roofs can partly compensate the loss of natural landscapes and mitigate runoff. Green roof plant selection has mostly been limited to species with the ability to survive extreme environmental conditions, typically Sedum species. However, green roofs with these species planted in monoculture may not be efficient in terms of rainfall retention. Selecting plants with water use plasticity and increasing substrate depth may improve green roof hydrological performance in these highly variable environments, without substantially introducing drought risk. To examine this, two experiments were conducted. The first, a 17-month controlled rainfall experiment, was examined in two studies: 1) to understand the overall effect of plant water use strategies and of the mixture of plants within water use plasticity on retention, and 2) to understand the mechanisms driving rainfall retention. The second experiment, a 27-month green roof field study, examined the effect of increasing substrate depth on rainfall retention and plant drought stress. Contrary to my initial hypotheses, plant species with water use plasticity did not always provide greater retention performance, compared with conservative succulent species. Retention performance of a plant mixture was dominated by key species and was not always greater than that of monocultures. The effects of plant treatment on retention performance were significant for medium and large rainfall events, but minor for small rainfall events. Rainfall characteristics and available storage of substrates before rainfall events strongly drove retention performance. Although the effects of ET were well understood and shown to be as the key factor for green roof retention, ‘Non-ET’ plant effects (i.e., the reduction in storage capacity, either via preferential flow, or roots occupying storage volume) can change the relative importance of ET. That is, the maximum storage capacity becomes a more important driver of available storage in the substrate compared with the ability of plants to deplete that stored moisture. Increasing substrate depth only marginally improved retention performance, however, deeper substrates significantly reduced the period of time plants were exposed to drought stress. The results showed that green roofs perform well for retention (average ~73% in total study period; average ~91% in per-event), particularly in warm and dry climates with a large proportion of small rainfall events. Selection of plants with water use plasticity and the use of mixtures was less important for improving retention than climate. However, when planting treatments were compared, root-induced preferential flow paths were shown to reduce the water storage capacity of substrates, reducing the water available for evapotranspiration and therefore reducing retention. Therefore, future plant selection needs to consider root traits in addition to water use strategies. In Melbourne, green roof substrates did not need to be much deeper than 150 mm, as the increase in retention was minimal beyond this depth and it was sufficient for maintaining plant performance.
Drivers of drought-related physiological, morphological and anatomical trait expression in a temperate eucalypt
Increasing drought-induced tree mortality is a global problem and current scientific efforts aim to understand why and how trees succumb to drought. Nevertheless, our current understanding of the mechanisms leading to drought-induced tree mortality is limited. Functional traits have emerged as useful indicators of drought susceptibility, but little research explores their underlying drivers. Traits expression can be under strong genetic control or phenotypically plastic. While traits under strong genetic control can only adapt through new generations, phenotypic plasticity facilitates adjustment within the lifetime of a tree which will be essential under rapid climate change. Hence, the ability to phenotypically adjust to drought is an important factor influencing drought resistance and therefore survival. However, differences in population genetics or phenotypic plasticity can result in diverse trait expressions even within a species. Hence, drought resistance may differ, even among populations of the same species. Furthermore, observations on drought mortality indicate that drought resistance likely differs between different developmental stages. However, we currently lack insights on how drought resistance in trees changes with developmental stage, which limits our understanding of species vulnerability under climate change. Therefore, this thesis analyses the expression of drought-related traits to determine which traits are phenotypically plastic as opposed to being under strong genetic control. To investigate phenotypic plasticity in detail, I conducted a set of experiments with different drought intensities and durations. These studies were performed on trees at different stages of development (i.e., seedlings, saplings, and mature trees) to also investigate how trait expression changes as trees develop. Eucalyptus obliqua L.Hér. was selected as the study species due to its ecological importance and wide distribution in south-eastern Australia, increasing the likelihood of trait plasticity. Two experimental chapters of my thesis (Chapter II & IV) were conducted in young (~15 y.o.) E. obliqua forest stands in south-eastern Australia. The first study aimed to investigate whether key drought-related traits are phenotypically plastic or under strong genetic control for E. obliqua populations growing across a precipitation gradient (Chapter II). Phenotypic plasticity was investigated by measuring the drought-related trait expression in five E. obliqua populations once during the summer and winter seasons. Further, seeds were collected at the five sites and seedlings were grown under uniform conditions to assess which traits are under strong genetic control. In the field, drier populations had smaller leaves with thicker xylem vessel walls, a lower vulnerability to embolism and a lower water potential at turgor loss point, which likely confers greater hydraulic safety. Significant phenotypic adjustments in physiological and morphological traits were observed in all E. obliqua populations. The Huber Value (sapwood to leaf area ratio; HV) increased during dry summer conditions while the water potential at turgor loss point decreased, likely to increase the populations’ hydraulic safety in the short-term. Under uniform conditions, seedlings from dry-origin populations express smaller leaves, higher anatomical cavitation resistance and a lower vulnerability to embolism, indicating that these traits are under strong genetic control. Overall, the study indicates that the phenotypic plasticity in physiological and morphological traits may facilitate adjustment to future climatic conditions. In contrast, anatomical traits related to hydraulic vulnerability are under strong genetic control and increase the drought resistance of dry-origin populations. A drought experiment was conducted on E. obliqua seedlings and saplings in a controlled glasshouse experiment (Chapter III). To investigate the potential of E. obliqua to acclimate to drier conditions, trees were first exposed to a mild long-term drought (drought conditioning). Following the mild drought, trees were subjected to a subsequent severe drought to induce mortality. Here, the aim was to investigate if drought conditioning can improve tree function and survival and if trait expression changes as trees transition from seedling into sapling stage. The results suggest that mild long-term drought stress induces mainly tree-level morphological trait adjustments, with drought conditioned trees having less leaf area and biomass compared to well-watered controls. Drought conditioning reduced total leaf area which reduced daily water demand, enabling trees to survive for longer during the severe drought event. However, no drought conditioning effect was observed on the water potential at turgor loss point, leaf xylem vulnerability to embolism, leaf size, maximum vessel diameter or wall thickness, indicating that leaf-level physiology, morphology or anatomy have a limited capacity to acclimate through long-term drought stress. Comparing seedling and sapling trait expression, most traits were observed to undergo substantial adjustments with developmental stages, suggesting that drought susceptibility likely changes significantly as trees develop. Seedlings likely confer drought resistance through a high anatomical cavitation resistance, smaller xylem vessel diameters and a lower vulnerability to embolism. In contrast, saplings express a lower water potential at turgor loss point and leaf mass fraction. Hence, drought resistance is likely to change during tree development. Further, depending on the developmental stage, drought resistance is likely to be conferred through different traits. The second field study (Chapter IV) was designed to analyse seasonal plasticity in drought-related traits as well as adjustments to severe drought in the field. To initialise the prolonged drought, we installed rainfall exclusion gutters, which covered 50% of the surface area, in a E. obliqua forest stand. While no treatment effect between control and treatment sites was observed, the three-years of seasonal trait measurement allowed conclusions on the responses of young trees to different drought intensities in a natural forest stand. In addition to the seasonal observations along the gradient (Chapter II), the data suggest that seasonal morphological adjustments, in form of leaf shedding, occur only in years with more severe drought stress, while physiological traits adjusted also to mild drought stress. In consideration of the observations in Chapter III, the results also suggest that the trees in the field were not severely drought stressed as the full plastic capacity was not implemented. As the field site, with 1000 mm mean annual precipitation, represents nearly E. obliqua’s distribution centre (Fig. 1.2b), this result would further suggest that most E. obliqua trees are under no imminent risk of succumbing to drought-induced tree mortality. In summary, my studies determined the contribution of physiological, morphological and leaf anatomical traits to the drought resistance of E. obliqua. The results highlight that leaf anatomy (i.e., cavitation resistance, xylem vessel walls) and vulnerability to embolism are under strong genetic control and confers a higher drought resistance for dry-origin populations. In contrast, most physiological and morphological traits demonstrated substantial potential to adjust towards drier conditions in all populations. As a general pattern, osmoregulation was observed under all drought durations and intensities, indicating that it is a main contributor to the drought response in E. obliqua. Intense droughts also induce morphological trait adjustments, in particular leaf shedding, which reduce the trees’ total leaf area and thus their water demand. Similarly, mild long-term droughts also induced leaf shedding as a key drought acclimation response. While the mild long-term drought exposure had only limited influence on the leaf physiology and no influence on leaf-level anatomy and morphology, the effectiveness of phenotypic adjustment in tree-level morphology became apparent during the subsequent severe drought. Trees that were previously exposed to the mild, long-term drought were able to delay canopy collapse and plant death during the subsequent severe drought from 27 to 39 days. In conclusion, my studies confirm that most physiological and morphological traits are phenotypically plastic and provide fundamental survival benefit in various drought situations. Phenotypic trait adjustments during tree development were also apparent for most traits and suggested that drought resistance differs during tree development. Seedlings conferred their drought resistance mainly through a high anatomical drought resistance and a low vulnerability to embolism. In contrast, saplings likely use physiological and morphological traits and their phenotypic plasticity to provide drought resistance and mature trees use a high anatomical drought resistance as well as physiological and morphological traits.
Assessing the impact of urbanization on streams using ecohydraulics
Urban stormwater runoff is a primary degrader of stream ecosystems. Excess stormwater runoff causes altered flow regimes, reduced in-stream water quality, and modified channel form. Recognition of such impacts has renewed interest in the protection or restoration of the hydrologic cycle in urban stormwater management. One of the greatest unknowns, however, is how the urban stormwater impacts on the flow regime translates to hydraulic conditions in the channel. This understanding is critical to identifying how the hydraulic environment supports complex and dynamic ecosystem functioning. My thesis investigates how the flow regime and channel form, and their interactions, deliver in-stream hydraulic outcomes for urban streams. I use hydrodynamic modelling (TUFLOW) to predict hydraulic outcomes, with the models developed using hydrographic and topographic data from two streams in the Melbourne region. I firstly predicted a range of ecologically relevant in-stream hydraulic metrics for a natural reach and compared the metrics to those predicted for a downstream urban reach of the same stream. I found that compared to the natural reach, the urban reach experienced much greater channel bed disturbance (~4 times higher), refuge habitat loss (~2 times smaller) and limited floodplain connectivity. This work thus supports with evidence casual physical mechanisms which likely drive urban stream degradation and habitat quality. I disentangled the relative role that the flow regime and channel form play in influencing in-stream hydraulic conditions. I did this through modelling different combinations of flow regime and channel form, - e.g. altered flow regime in a natural channel versus natural flow regime in a modified channel. I found that both flow regime and channel form play key roles in setting the hydraulic conditions. The work revealed that the hydraulic regime is sensitive to the channel morphology which controlled key aspects of the hydraulic regime (e.g. magnitude, frequency and duration). After establishing that both flow regime and channel form play important roles in regulating in-stream hydraulics, I investigated potential management actions which could be applied in both developed- and developing landscapes. I tested the benefit of alternative channel rehabilitation design configurations applied to a (synthetic) degraded urban stream. This work showed that channels designed to increase morphological complexity yield ecohydraulic benefits, but not to levels which approach those predicted under natural flow- and channel conditions. With altered flow regime limiting the effectiveness of such channel designs, I tested if this stressor could be mitigated in developing landscapes using alternative approaches towards urban stormwater management. It demonstrates that widespread application of Stormwater Control Measures (SCMs) could influence the flow regime in ways which translate to ecohydraulic conditions remaining at natural levels. This work suggests that for urbanized catchments, management of in-stream hydraulics requires attention to both the flow regime (using SCMs) and channel form (e.g. through sediment regime management to allow appropriate levels of sediment supply and transport). For developing catchments, SCMs should be implemented as part of the construction phase in order to minimize downstream hydrologic impact and thus geomorphic degradation. My thesis provides an improved mechanistic understanding of why streams draining urban landscapes are commonly physically and ecologically degraded. Consideration of ecohydraulic indicators in urban stream management could provide targets, and help inform efforts towards stream protection or restoration, particularly when ecological objectives are central.