Long-term flowering patterns of melliferous Eucalyptus (Myrtaceae) species

The flowering patterns of 28 Victorian melliferous (honey-producing) eucalypts were investigated by using long-term observations of highly experienced, commercial apiarists. Frequency, timing, duration and intensity of flowering were determined, as were spatial differences within and among species. Data were obtained by face-to-face interviews with 25 Victorian apiarists, each of whom had operated a minimum of 350 hives for a minimum of 30 years. Flowering frequency ranged from 1 to 7 years, and most species flowered once every 2–4 years. Long-term flowering frequency, timing and duration were reported as constant, although short-term perturbations could occur. Most melliferous species flowered during spring and summer for a period of 3 months or more. Only few species had shorter flowering periods. Information provided by apiarists compared well with available published information (e.g. flowering period reported in field guides) and revealed a reliable, largely untapped source of long-term data, the use of which could benefit many ecological research endeavours.

Flowering ecology of honey-producing flora in South-East Australia

This report examines the flowering ecology (flowering patterns and the production of floral resources, i.e. nectar and pollen) of important Australian melliferous (honey-producing) flora. Aspects of flowering ecology that can have a negative impact on invertebrates, including honeybees, were also investigated. The research was based on information sourced by highly experienced, commercial beekeepers and, so, provides a valuable written record of long-term observations relating to flowering ecology which otherwise may be lost following the death of beekeepers. Results of this study are of far-reaching importance, not only to the beekeeping industry, but to land managers, the general public and the future of Australian flora and fauna.

An understanding of flowering ecology is vital for many reasons, including implementing appropriate management practices which ensure the sustainability and growth of natural resources and industries like the beekeeping industry. Despite the importance of such studies, very little research has considered flowering ecology in Australian flora. Furthermore, research often was based on short-term data; long-term data are widely acknowledged as being necessary in such research in order to determine ‘real’ flowering patterns. Thus, studies of flowering ecology which use long-term data are vital.

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.

Flowering ecology of south-east Australian melliferous flora

The flowering ecology of south-east Australian melliferous (honey) flora was studied, using observational data from our most experienced beekeepers. Short-term variation and long-term trends were observed which may have critical ecological implications. The study is a significant contribution to flowering ecology and provides an important foundation to guide future research.

Isolation of sperms from the pollen tube of flowering plants during fertilization

Sperm cells have been isolated from pollen tubes growing in style segments of the dicotlyledon Rhododendron macgregoriae and the monocotyledon Gladiolus gandavensis by the in vivo/in vitro method at various stages of fertilization. Pollen tubes emerged from the cut end of the style into agar medium, and more than 95% contained sperm cells. Sperm cells were released from the pollen tubes by osmotic shock or by placing styles in wall-degrading enzymes: 0.5% macerozyme and 1% cellulase. The isolated sperms were ellipsoidal protoplasts of diameter about 2 × 3 micrometers in Gladiolus and about 3 × 4 micrometers in Rhododendron. After isolation, a proportion of the sperm cells occurred in pairs linked at one end by finger-like connections. The pairs of isolated sperms were dimorphic in terms of surface area and volume. By cutting the styles at various positions and times after pollination, the potential exists to detect changes in sperm gene expression associated with fertilization.

The potential for indirect effects between co-flowering plants via shared pollinators depends on resource abundance, accessibility and relatedness

Co-flowering plant species commonly share flower visitors, and thus have the potential to influence each other's pollination. In this study we analysed 750 quantitative plant-pollinator networks from 28 studies representing diverse biomes worldwide. We show that the potential for one plant species to influence another indirectly via shared pollinators was greater for plants whose resources were more abundant (higher floral unit number and nectar sugar content) and more accessible. The potential indirect influence was also stronger between phylogenetically closer plant species and was independent of plant geographic origin (native vs. non-native). The positive effect of nectar sugar content and phylogenetic proximity was much more accentuated for bees than for other groups. Consequently, the impact of these factors depends on the pollination mode of plants, e.g. bee or fly pollinated. Our findings may help predict which plant species have the greatest importance in the functioning of plant-pollination networks.

Refugees and residents: densities and habitat preferences of lorikeets in urban Melbourne

Lorikeet densities were measured across four habitat types in urban Melbourne. Musk Glossopsitta concinna and Rainbow Lorikeets Trichoglossus haematodus were shown to preferentially use established streetscapes with predominantly native vegetation. The high densities of Musk Lorikeets recorded possibly reflect a paucity of flowering in Victorian BoxIronbark forests during the autumnlwinter of 2002 and the availability of supplementary nectar resources in the urban environment. Future planting decisions in recently developed streetscapes will dictate the long-term resource potential for lorikeets and other nectarivores in urban Melbourne.

Does firefighting foam affect the growth of some Australian native plants?

Firefighting foams (Class A foams) are an effective and widespread firefighting tool, and are frequently used in environmentally sensitive areas. They are known to be ecologically damaging in aquatic environments; however, their impacts at the plant species or ecosystem level are relatively unknown. Reports of shoot damage to plants, suppressed flowering and changes in plant community composition suggest that the environmental damage caused by their use may be unacceptable. Applications of four levels of foam to seedlings of seven Australian plant species, from five representative and widespread families, showed no detectable impacts on a range of vegetative growth characteristics. The results are encouraging for continued use of firefighting foam in sensitive natural habitats.

Surviving suppression : no detectable impacts of Class A foam on soil invertebrates and some Australian native plants

Firefighting foams (Class A foams) are an effective and widespread firefighting tool which are frequently used in environmentally sensitive areas. Firefighting foams are known to be ecologically damaging in aquatic environments, however their impacts at the plant species or ecosystem level are relatively unknown. Reports of shoot damage to plants, suppressed flowering, and changes in plant community composition suggested that the ecological damage caused by their use may be unacceptable. However, applications of foam to seedlings of some Australian plant species from representative and widespread families, showed no detectable impacts on a range of vegetative growth characteristics. Application of 1.0% foam to heathland soils showed no detectable impacts on soil invertebrate Orders sampled over several months. The results are encouraging for the continued use of Class A foam as a fire suppression technique.

Collapse of an avifauna : climate change appears to exacerbate habitat loss and degradation

Aim. We characterized changes in reporting rates and abundances of bird species over a period of severe rainfall deficiency and increasing average temperatures. We also measured flowering in eucalypts, which support large numbers of nectarivores characteristic of the region.

Location.  A 30,000-km2 region of northern Victoria, Australia, consisting of limited amounts of remnant native woodlands embedded in largely agricultural landscapes.

Methods. There were three sets of monitoring studies, pitched at regional  (survey programmes in 1995–97, 2004–05 and 2006–08), landscape (2002–03 and 2006–07) and site (1997–2008 continuously) scales. Bird survey techniques used a standard 2-ha, 20-min count method. We used Bayesian analyses of reporting rates to document statistically changes in the avifauna through time at each spatial scale.

Results. Bird populations in the largest remnants of native vegetation (up to 40,000 ha), some of which have been declared as national parks in the past decade, experienced similar declines to those in heavily cleared andscapes. All categories of birds (guilds based on foraging substrate, diet, nest site; relative mobility; geographical distributions) were affected similarly. We detected virtually no bird breeding in the latest survey periods. Eucalypt flowering has declined significantly over the past 12 years of drought.

Main conclusions. Declines in the largest woodland remnants commensurate with those in cleared landscapes suggest that reserve systems may not be relied upon to sustain species under climate change. We attribute population declines to low breeding success due to reduced food. Resilience of bird populations in this woodland system might be increased by active management to enhance habitat quality in existing vegetation and restoration of woodland in the more fertile parts of landscapes.

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.