Flowering ecology of a Box-Ironbark Eucalyptus community

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.

Ecological value of riparian zones to birds in forest landscapes

Riparian zones are a characteristic component of many landscapes throughout the world and increasingly are valued as key areas for biodiversity conservation. Their importance for bird communities has been well recognised in semi-arid environments and in modified landscapes where there is a marked contrast between riparian and adjacent non-riparian vegetation. The value of riparian zones in largely intact landscapes with continuous vegetation cover is less well understood. This research examined the importance of riparian habitats for avifauna conservation by investigating the ecological interactions contributing to the pattern of bird assemblages in riparian and adjacent non-riparian habitats. Specifically, the focus is on the bird assemblages of riparian zones and those of adjacent non-riparian vegetation types and the influence that associated differences in resource availabilities, habitat structure and conditions have on observed patterns. This study was conducted in the foothill forests of the Victorian Highlands, south-east Australia. Mixed-species eucalypt (genus Eucalyptus) forests dominate the vegetation of this region. Site selection was based on the occurrence of suitable riparian habitat interspersed within extensive, relatively undisturbed (i.e. no recent timber harvesting or fire events) forest mosaics. A series of 30 paired riparian and non-riparian sites were established among six stream systems in three forest areas (Bunyip State Park, Kinglake National Park and Marysville State Forest). Riparian sites were positioned alongside the stream and the non-riparian partner site was positioned on a facing slope at a distance of approximately 750 m. Bird surveys were carried out during 29 visits to each site between July 2001 and December 2002. Riparian sites were floristically distinct from non-riparian sites and had a more complex vegetation structure, including a mid-storey tree layer mostly absent from non-riparian sites, extensive fine litter and coarse woody debris, and dense ground-layer vegetation (e.g. sedges and ground ferns). The characteristic features of non-riparian habitats included a relatively dense canopy cover, a ground layer dominated by grasses and fine litter, and a high density of canopy-forming trees in the smaller size-classes. Riparian zones supported a significantly greater species richness, abundance and diversity of birds when compared to non-riparian habitats. The composition of bird assemblages differed significantly between riparian and non-riparian habitats, with riparian assemblages displaying a higher level of similarity among sites. The strongest contributors to observed dissimilarities between habitat types included species that occurred exclusively in either habitat type or species with large contrasts in abundance between habitat types. Much of the avifauna (36%) of the study area is composed of species that are common and widespread in south-east Australia (i.e. forest generalists). Riparian habitats were characterised by a suite of species more typical of wetter forest types in south-east Australia and many of these species had a restricted distribution in the forest mosaic. Some species (7%) occurred exclusively in riparian habitats (i.e. riparian selective species) while others (43%) were strongly linked to these habitats (i.e. riparian associated species). A smaller proportion of species occurred exclusively (2%) in non-riparian habitats (i.e. non-riparian selective species) or were strongly linked to these habitats (10%; i.e. non-riparian associated species). To examine the seasonal dynamics of assemblages, the variation through time in species richness, abundance and composition was compared between riparian and non-riparian sites. Riparian assemblages supported greater richness and abundance, and displayed less variation in these parameters, than non-riparian assemblages at all times. The species composition of riparian assemblages was distinct from non-riparian assemblages throughout the annual cycle. An influx of seasonal migrants elevated species richness and abundance in the forest landscape during spring and summer. The large-scale movement pattern (e.g. coastal migrant, inland migrant) adopted by migrating species was associated with their preference for riparian or non-riparian habitats in the landscape. Species which migrate north-south along the east coast of mainland Australia (i.e. coastal migrants) used riparian zones disproportionately; eight of eleven species were riparian associated species. Species which migrate north-south through inland Australia (i.e. inland migrants) were mostly associated with non-riparian habitats. The significant differences in the dynamics of community structure between riparian and non-riparian assemblages shows that there is a disproportionate use of riparian zones across the landscape and that they provide higher quality habitat for birds throughout the annual cycle. To examine the ecological mechanisms by which riparian assemblages are richer and support more individual birds, the number of ecological groups (foraging, nest-type and body mass groups) represented, and the species richness of these groups, was compared between riparian and non-riparian assemblages. The structurally complex vegetation and distinctive habitat features (e.g. aquatic environments, damp sheltered litter) provided in the riparian zone, resulted in the consistent addition of ecological groups to riparian assemblages (e.g. sheltered ground – invertebrates foraging group) compared with non-riparian assemblages. Greater species richness was accommodated in most foraging, nest-type and body mass groups in riparian than non-riparian assemblages. Riparian zones facilitated greater richness within ecological groups by providing conditions (i.e. more types of resources and greater abundance of resources) that promoted ecological segregation between ecologically similar species. For a set of commonly observed species, significant differences in their use of structural features, substrates and heights were registered between riparian and non-riparian habitats. The availability and dynamics of resources in riparian and non-riparian habitats were examined to determine if there is differential availability of particular resources, or in their temporal availability, throughout the annual cycle. Riparian zones supported more abundant and temporally reliable eucalypt flowering (i.e. nectar) than non-riparian habitats throughout the annual cycle. Riparian zones also supported an extensive loose bark resource (an important microhabitat for invertebrates) including more peeling bark and hanging bark throughout the year than at non-riparian sites. The productivity of eucalypts differed between habitat types, being higher in riparian zones at most times for all eucalypts combined, and for some species (e.g. Narrow-leaved Peppermint Eucalyptus radiata). Non-riparian habitats provided an abundant nectar resource (i.e. shrub flowering) at particular periods in the annual cycle. Birds showed clear relationships with the availability of specific food (i.e. nectar) and foraging resources (i.e. loose bark). The demonstration of a greater abundance of resources and higher primary productivity in riparian zones is consistent with the hypothesis that these linear strips that occupy only a small proportion of the landscape have a disproportionately high value for birds. Riparian zones in continuous eucalypt forest provide high quality habitats that contribute to the diversity of habitats and resources available to birds in the forest mosaic, with positive benefits for the landscape-level species pool. Despite riparian and non-riparian habitat supporting distinct assemblages of birds, strong linkages are maintained along the riparian-upslope gradient. Clearly, the maintenance of diverse and sustainable assemblages of birds in forest landscapes depends on complementary management of both riparian and non-riparian vegetation.

Phytogeny of the Quambalariaceae fam. nov., including important Eucalyptus pathogens in South Africa and Australia

The genus Quambalaria consists of plant-pathogenic fungi causing disease on leaves and shoots of species of Eucalyptus and its close relative, Corymbia. The phylogenetic relationship of Quambalaria spp., previously classified in genera such as Sporothrix and Ramularia, has never been addressed. It has, however, been suggested that they belong to the basidiomycete orders Exobasidiales or Ustilaginales. The aim of this study was thus to consider the ordinal relationships of Q. eucalypti and Q. pitereka using ribosomal LSU sequences. Sequence data from the ITS nrDNA were used to determine the phylogenetic relationship of the two Quambalaria species together with Fugomyces (= Cerinosterus) cyanescens. In addition to sequence data, the ultrastructure of the septal pores of the species in question was compared. From the LSU sequence data it was concluded that Quambalaria spp. and F. cyanescens form a monophyletic clade in the Microstromatales, an order of the Ustilaginomycetes. Sequences from the ITS region confirmed that Q. pitereka and Q. eucalypti are distinct species. The ex-type isolate of F. cyanescens, together with another isolate from Eucalyptus in Australia, constitute a third species of Quambalaria, Q. cyanescens (de Hoog & G.A. de Vries) Z.W. de Beer, Begerow & R. Bauer comb. nov. Transmission electron-microscopic studies of the septal pores confirm that all three Quambalaria spp. have dolipores with swollen lips, which differ from other members of the Microstromatales (i.e. the Microstromataceae and Volvocisporiaceae) that have simple pores with more or less rounded pore lips. Based on their unique ultrastructural features and the monophyly of the three Quambalaria spp. in the Microstromatales, a new family, Quambalariaceae Z.W. de Beer, Begerow & R. Bauer fam. nov., is described.

Roost tree characteristics determine use by the white-striped freetail bat (Tadarida australis, Chiroptera : Molossidae) in suburban subtropical Brisbane, Australia

We examined factors affecting roost tree selection by the white-striped freetail bat Tadarida australis (Chiroptera: Molossidae), a large insectivorous bat in suburban Brisbane, Australia. We compared biophysical characteristics associated with 34 roost trees and 170 control trees of similar diameter, height and tree senescence characters. Roost trees used by the white-striped freetail bat had significantly higher numbers of hollows in the trunk and branches (P < 0.003) and were more likely to contain a large trunk cavity with an internal diameter of > 30 cm (P < 0.001) than control trees. These trees also accommodated more species of hollow-using fauna (P = 0.005). When comparing roost trees with control trees of similar diameters and heights, roost trees were on average at a later stage of tree senescence (P < 0.001). None of the roost trees were found in the large forest reserves fringing the Brisbane metropolitan area despite these areas being used for foraging by the white-striped freetail bat. Although all tree locations in this study were in modified landscapes, roost trees tended to be surrounded by groups of trees and undergrowth. Roost trees provide important habitat requirements for hollow-using fauna in suburban, rural and forested environments.

Australia`s box - ironbark forests and woodlands : saving the fragments of a threatened ecosystem

Australia's box-ironbark forests and woodlands once covered about 14 per cent of the State of Victoria on the riverine plains and foothills of the Great Dividing Range. But approximately 83 per cent of the total original habitat has been destroyed and what remains of this significant ecosystem is now highly fragmented and vulnerable to further degradation. Moreover, only 14 per cent of the area remaining is on public land. A 10 year campaign on the part of the environmental movement eventually succeeded in forcing the State government to conduct an independent inquiry into this ecosystem and make recommendations on future management. This paper outlines the innovative public participation process adopted by the Victorian State government and the outcomes of the inquiry. A subsequent compensation package for commercial operations disadvantaged by the proclamation of a series of new national parks is also discussed, as are the shortcomings of a process that can have little or no impact on what happens on private land.

Ecology an Australian perspective

This is the first ecological text that deals comprehensively with ecological principles and practice in an Australian context, with wholly Australian examples. There are four sections, dealing with the basics of climate soils and energy flows, major communities, the discipline itself, and major issues.

Living with Dungalah (Murray River) : the case study of an environmental issue in Australia

This research is a case study of Dungalah, a river Europeans call the Murray which describes and accounts for the past, present and future experiences of Indigenous and non-Indigenous communities in their protection, use and management of the Dungalah and its surrounding land. It provides a snapshot of the experiences of the researcher, her family, friencds and the Yorta Yorta people, living with Dungalah.

Importance of riparian vegetation to terrestrial avifauna along the Murray River, south-eastern Australia

Bird life occurring along the Murray River was distinctly different from surrounding much drier vegetation. It was found that the presence of the Murray River, with it's associated moist Red Gum forests, provide a corridor whereby birds typically of cool climates can expand their range and occur in an arid landscape.

Sooty owl ecology and recent small mammal decline

This thesis investigated the ecology of the threatened sooty owl, which was used to improve understanding of recent small mammal declines and how a top-order predator adapted to changes in ecosystem condition following European settlement. This knowledge will help improve biodiversity conservation and management of forested ecosystems in south-eastern Australia.

Challenges for native grassland conservation on Victoria's Northern Plains

Discusses some of the challenges faced in attempting to retain and conserve grasslands on Victoria's Northern Plains over the past decade. The development of a strategic vision and directions and opportunities for the future are highlighted.