Dasyurid dilemmas: problems and solutions for conserving Australia's small carnivorous marsupials

Dasyurid marsupials are distributed throughout the major terrestrial environments of Australia but since European settlement have suffered local and regional extinctions, range reductions and population declines. In this paper we examine the conservation status of small dasyurids (<500 g) and the threats they face. We also evaluate recovery procedures for threatened taxa and assess their success. Twenty-four percent of smaller dasyurids are classified as vulnerable, endangered or data deficient. Large body size and occupancy of one or two habitat types are correlated strongly with  endangerment species currently considered as 'low risk, near threatened' group closely with vulnerable and endangered species, indicating a risk of further declines. The processes contributing most to declines include habitat loss and fragmentation, altered fire regimes and predation. As of April 200 I, no Recovery Plans had been adopted by the Commonwealth Govemment for any small dasyund species. There is much information on the reproduction and development of smaller dasyurids, making them suitable for captive breeding. However, captive breeding programs have been limited. the  dibbler Paranrechinus apicalis being the only species bred systematically for reintroductions. There is a need for integration between captive breeding programs and recovery planning. as well as for more information on the population viability and metapopulation structures of small dasyurids genetic diversity of populations and inbreeding depression. We suggest a program of survey. research. management and education to Improve conservation outcomes for all small dasyurids.

The influence of time since fire and distance from fire boundary on the distribution and abundance of arboreal marsupials in Eucalyptus regnans-dominated forest in the Central Highlands of Victoria

The impact of time since fire after two consecutive wildfires 44 years apart (1939 and 1983) within the same area, and the distance from the fire boundary «100 m or 500-2000 m), were investigated in relation to the distribution and abundance of arboreal marsupials in 1994. Arboreal marsupials were censused by stagwatching and spotlighting in two relatively young age classes of mountain ash (Eucalyptus regnans) dominated forest in the Central Highlands of Victoria. Five species of arboreal marsupial were detected, but only three were detected in sufficient numbers to determine habitat preferences. Petauroides volans (greater glider) was statistically more abundant in 1939 regrowth forests, while Trichosurus caninus (mountain brushtail possum) showed no significant preference for either age class of forest. All but one record of Gymnobelideus leadbeateri (Leadbeater's possum) came from young forest, though the effect of age-class was not statistically significant. Distance from fire boundary explained little or no variation in mammal distribution or abundance. While the actual number of hollow-bearing trees was similar in both age classes of forest, the long-term lifespan of hollow-bearing trees in more recently burnt forest is predicted to be lower than in unburnt or not recently burnt forest. Post-fire salvage logging following the 1983 wildfires appears to have reduced the number of hollow-bearing trees at sites burnt in 1983.

Ecology of arboreal marsupials in a network of remnant linear habitats

Linear strips of vegetation set within a less-hospitable matrix are common features of landscapes throughout the world. Depending on location, form and function, these linear landscape elements include hedgerows, fencerows, shelterbelts, roadside or streamside strips and wildlife corridors. In many anthropogenically-modified landscapes, linear strips are important components for conservation because they provide a large proportion of the remaining wooded or shrubby habitat for fauna. They may also function to provide connectivity across the landscape. In some districts, the linear strips form an interconnected network of habitat. The spatial configuration of remnant habitat (size, shape and arrangement) may influence habitat suitability, and hence survival, of many species of plant and animal in modified landscapes. Near Euroa in south-eastern Australia, the clearing and fragmentation of temperate woodlands for agriculture has been extensive and, at present, less than 5% tree cover remains, most of which (83%) occurs as linear strips along roads and streams. The remainder of the woodland occurs as relatively small patches and single isolated trees scattered across the landscape. As an assemblage, arboreal marsupials are woodland dependent and vary in their sensitivity to habitat loss and fragmentation. This thesis focusses on determining the conservation status of arboreal marsupials in the linear network and understanding how they utilise the landscape mosaic. Specifically, the topics examined in this thesis are: (1) the composition of the arboreal marsupial assemblage in linear and non-linear woodland remnants; (2) the status and habitat preferences of species of arboreal marsupial within linear remnants; and (3) the ecology of a population of the Squirrel Glider Petaurus norfolcensis in the linear network, focusing on population dynamics, spatial organisation, and use of den trees. The arboreal marsupial fauna in the linear network was diverse, and comprised seven out of eight species known to occur in the district. The species detected within the strips were P. norfolcensis, the Sugar Glider Petaurus breviceps, Common Brushtail Possum Trichosums vulpecula, Common Ringtail Possum Pseudocheirus peregrinus, Brush-tailed Phascogale Phascogale tapoatafa, Koala Phascolarctos cinereus and Yellow-footed Antechinus Antechinus flavipes. The species not detected was the Feathertail Glider Acrabates pygmaeus. Survey sites in linear remnants (strips of woodland along roads and streams) supported a similar richness and density of arboreal mammals to sites in non-linear remnants (large patches or continuous tracts of woodland nearby). Furthermore, the combined abundance of all species of arboreal marsupials was significantly greater in sites in the linear remnants than in the non-linear remnants. This initial phase of the study provided no evidence that linear woodland remnants support a degraded or impoverished arboreal marsupial fauna in comparison with the nonlinear remnants surveyed. Intensive trapping of arboreal marsupials within a 15 km linear network between February 1997 and June 1998 showed that all species of arboreal marsupial (except A. pygmaeus) were present within the linear strips. Further analyses related trap-based abundance estimates to measures of habitat quality and landscape structure. Width of the linear habitat was significantly positively correlated with the combined abundance of all arboreal marsupials, as well as with the abundance of P. norfolcensis and T. vulpecula. The abundance of T. vulpecula was also significantly positively correlated with variation in overstorey species composition, Acacia density and the number of hollow-bearing trees. The abundance of P. norfolcensis was positively correlated with Acacia density and canopy width, and negatively correlated with distance to the nearest intersection with another linear remnant. No significant variables were identified to explain the abundance of P. tapoatafa, and there were insufficient captures of the remaining species to investigate habitat preferences. Petaurus norfolcensis were resident within the linear network and their density (0.95 -1.54 ha-1) was equal to the maximum densities recorded for this species in continuous forest elsewhere in south-eastern Australia. Rates of reproduction were also similar to those in continuous forest, with births occurring between May and December, a mean natality rate of 1.9, and a mean litter size of 1.7. Sex ratios never differed significantly from parity. Overall, the population dynamics of P. norfolcensis were comparable with published results for the species in contiguous forest, clearly suggesting that the linear remnants currently support a self-sustaining, viable population. Fifty-one P. norfolcensis were fitted with radio transmitters and tracked intermittently between December 1997 and November 1998. Home ranges were small (1.3 - 2.8 ha), narrow (20 - 40 m) and elongated (322 - 839 m). Home ranges were mostly confined to the linear remnants, although 80% of gliders also utilised small clumps of adjacent woodland within farm paddocks for foraging or denning. Home range size was significantly larger at intersections between two or more linear remnants than within straight sections of linear remnants. Intersections appeared to be important sites for social interaction because the overlap of home ranges of members of adjacent social groups was significantly greater at intersections than straight sections. Intersections provided the only opportunity for members of three or more social groups to interact, while still maintaining their territories. The 51 gliders were radiotracked to 143 different hollow-bearing trees on 2081 occasions. On average, gliders used 5.3 den trees during the study (range 1-15), and changed den trees every 4.9 days. The number of den trees used by each glider is likely to be conservative because the cumulative number of den trees continued to increase over the full duration of the study. When gliders shifted between den trees, the mean distance between consecutive den sites was 247 m. Den trees were located throughout a glider's home range, thereby reducing the need to return to a central den site and potentially minimising energy expenditure. Dens were usually located in large trees (mean diameter 88.5 cm) and were selected significantly more often than expected based on their occurrence within the landscape. The overall conclusion of this thesis is that the linear network I studied provides high quality habitat for resident populations of arboreal marsupials. Important factors influencing the suitability of the linear remnants appear to be the high level of network connectivity, the location on soils of high nutrient status, the high density of large trees and an acacia understorey. In highly fragmented landscapes, linear habitats as part of the remaining woodland mosaic have the potential to be an integral component in the conservation of woodland-dependent fauna. The habitat value of linear strips of vegetation should not be underestimated.

Fire severity and landscape context effects on arboreal marsupials

Although fire is a major form of natural disturbance worldwide, both fire-derived landscape context effects and the impacts of fire severity are poorly known for many species. To address this knowledge gap, we quantified the response of Australian arboreal marsupials to: (1) the spatial effects of fire, (2) fire severity, and (3) fire impacts on the availability of critical nesting resources - hollow-bearing trees.We identified substantial differences among species in response to fire severity and landscape-scale fire. The Sugar Glider (Petaurus breviceps) and the endangered Leadbeater's Possum (Gymnobelideus leadbeateri) were extremely rare on burned sites irrespective of fire severity. In addition, these two species declined with the amount of burned forest in the surrounding landscape even when their habitat remained unburnt. The Mountain Brushtail Possum (Trichosurus cunninghami) and the Greater Glider (Petauroides volans) both occurred on burned and unburned sites. The Greater Glider responded negatively to fire severity at the site level and also negatively to the amount of forest burned in the surrounding landscape. The abundance of the Mountain Brushtail Possum was lowest on sites subject to moderate severity fire.On unburned sites, the presence and abundance of virtually all species was characterised by a common positive response to the availability of nesting resources in hollow-bearing trees.Our findings underscore the importance of management practices to better protect species that decline after fire. These include conserving areas of unburned forest, particularly those with hollow-bearing trees which are critical nest sites for arboreal marsupials. These recommendations are currently the opposite of existing management practices.

One size does not fit all : monitoring faecal glucocorticoid metabolites in marsupials

Marsupial research, conservation, and management can benefit greatly from knowledge about glucocorticoid (GC) secretion patterns because GCs influence numerous aspects of physiology and play a crucial role in regulating an animal's response to stressors. Faecal glucocorticoid metabolites (FGM) offer a non-invasive tool for tracking changes in GCs over time. To date, there are relatively few validated assays for marsupials compared with other taxa, and those that have been published generally test only one assay. However, different assays can yield very different signals of adrenal activity. The goal of this study was to compare the performance of five different enzyme immunoassays (EIAs) for monitoring adrenocortical activity via FGM in 13 marsupial species. We monitored FGM response to two types of events: biological stressors (e.g., transport, novel environment) and pharmacological stimulation (ACTH injection). For each individual animal and assay, FGM peaks were identified using the iterative baseline approach. Performance of the EIAs for each species was evaluated by determining (1) the percent of individuals with a detectable peak 0.125-4.5days post-event, and (2) the biological sensitivity of the assay as measured by strength of the post-event response relative to baseline variability (Z-score). Assays were defined as successful if they detected a peak in at least 50% of the individuals and the mean species response had a Z⩾2. By this criterion, at least one assay was successful in 10 of the 13 species, but the best-performing assay varied among species, even those species that were closely related. Furthermore, the ability to confidently assess assay performance was influenced by the experimental protocols used. We discuss the implications of our findings for biological validation studies.

Milk: milk of monotremes and marsupials

Lactation has evolved from an ancient reproductive strategy which appears to have been present long before the evolution of extantmammals. The ability to lactate is a feature only found among mammals and involves a facet of maternal care where mothers secretea nutrient-rich milk which is delivered to the young by the mammary gland. Evolutionary studies indicate that lactation was establishedprior to divergence of extant mammalian lineages. It also seems that lactation evolved long before gestation in utero. Secretionsof ancestral mammary glands may have had antimicrobial properties that protected either eggs or hatchlings and organiccomponents that supplemented offspring nutrition. Over the course of evolution, lactation became a highly efficient, effectiveand adaptable means of providing maternal care for neonates. The evolution of a placenta in eutherian mammals resulted inmore extensive intrauterine development of an embryo and the ability to lactate after birth became a critical part of the reproductivestrategy of mammalian species. Following development of highly nutritious milks, evolution produced diversity in milk compositionand function, quantity of milk output, length of lactation, length of intervals between nursing and contributions of lactation tooffspring nutrition.

Conservation management and diets of Powerful Owls (Ninox strenua) in outer urban Melbourne, Australia

The Powerful Owl Ninox strenua is Australia’s largest owl, and is mainly found east of the Great Dividing Range on the mainland in tall-open forests. The species is considered rare, both nationally and in the State of Victoria; and threatened in the Greater Melbourne area. Recovery plans for the future conservation management of N. strenua are being prepared in 2 states.

Historically, Powerful Owls have been thought to require large homes ranges (about 1000 ha per pair) in suitable old-growth forest, which provides nest hollows for the owls and their arboreal marsupial prey. Recent research, however, has found N. strenua may be more numerous and breed more successfully in a wider range of habitats than previously believed. In particular, the birds have been found living in forests and woodlands within the greater metropolitan areas of cities. The most extreme case is where a nest tree has been found within 800m of urban settlement and 6km from the centre of Brisbane.

In this paper we report on the diet, habitat use, and conservation management by a number of breeding pairs of owls in outer urban Melbourne. Study sites range from a relatively undisturbed rainforest habitat 80km from central Melbourne, through dry sclerophyll, eucalyptus-dominated open forest with some disturbance to a site 8km from central Melbourne in highly disturbed urban parkland.

Diets of the families of owls were determined by analyzing remains in regurgitated pellets. The data confirm that arboreal marsupials constitute the major prey items, especially the Common Ringtail Possum Pseudocheirus peregrinus. There were differences in diets depending on the availability of prey species, which suggest a level of opportunism not previously suspected. Our study is also the first to confirm the owls capture adult Common Brushtail Possums Trichosurus vulpecula (15% of pellets containing the remains of this large opossum have bones of mature adults at 1 site) and thus take prey up to two and a half times their own weight. As well our data suggest Powerful Owls are not restricted to hollow-dwelling prey, as in some sites the marsupials rested during the day either in leafy nests called dreys (P. peregrinus) or in house roofs (T. vulpecula).

In the most heavily disturbed sites, breeding success has been reduced, and we have evidence that in one particular year the young were eaten by one of the parents. This followed construction of a bicycle track under the nest during the breeding season. Recommendations are made for the future conservation and habitat management of Powerful Owls in the Yarra Valley corridor.

Are day-active small mammals rare and small birds abundant in Australian desert environments because small mammals are inferior thermoregulators?

Small desert birds are typically diurnal and highly mobile (hence conspicuous) whereas small non-volant mammals are generally nocturnal and less mobile (hence inconspicuous). Birds are more mobile than terrestrial mammals on a local and geographic scale, and most desert birds are not endemic but simply move to avoid the extremes of desert conditions. Many small desert mammals are relatively sedentary and regularly use physiological adjustments to cope with their desert environment (e.g., aestivation or hibernation). It seems likely that prey activity patterns and reduced conspicuousness to predators have reinforced nocturnality in small desert mammals. Differences such as nocturnality and mobility simply reflect differing life-history traits of birds and mammals rather than being a direct result of their differences in physiological capacity for tolerating daytime desert conditions.

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.

Succession of small mammal communities after fire and reintroduction of the swamp antechinus 'Antechinus minimus'

This thesis is involved with changes that have occurred to small mammal populations following a major disturbance in the Anglesea region as a result of the 1983 Ash Wednesday fires. Fire, with its effects on spatial and temporal heterogeneity, was found to be an important factor in the maintenance of vegetation and small mammal community structure and diversity in the region. Successional changes in vegetation and small mammal communities were described by multivariate analyses, using data collected annually from 22 study sites. The use of factor analysis techniques, in reducing the annual capture data content, enabled long-term changes in the structure of mammal communities to be interpreted. The small mammal communities in the coastal heath and forest vegetation in the Anglesea region show evidence of a general resilience, (the degree and speed of recovery), to disturbance. Two phases of successional response to fire by mammal species have been proposed; a ‘re-establishment’ phase which occurs in the initial 5-6 years post-fire and is accompanied by rapid increase in species’ abundance, and a subsequent ‘maintenance’ phase accompanied by relatively minor changes in abundance. Habitat Suitability Indices were produced relating to these phases. Vertical density measures of understorey shrubs and herb layers showed significant relationships with small mammal species abundance at the study sites. Long term studies following major disturbances are needed to distinguish between short term recovery of plant and animal species and long term changes in these species. Studies extending over a number of years enable a better directional view of changes in small mammal communities than can be determined from . observations made over a short period. As a part of the investigation into temporal change, it was proposed to undertake trial reintroductions of the Swamp antechinus, Ant echinus minimus, a marsupial dasyurid species which was trapped in the area prior to the 1983 fire, but rarely subsequently. Other more commonly observed native small mammal species (e.g. Rattus fuscipes,R. lutreolus, Antechinus stuartii, Sminthopsis leucopus) had re-invaded the proposed reintroduction site after this fire. Failure of A. minimus to re-establish may have been due to spatial separation of the pre-fire populations coupled with the extensive area burnt in 1983, A source population of the species was located about 100km to the west and habitat utilization and interspecific and niche relationships between the species making the small mammal community explored. Discriminant analysis revealed some spatial separation of species within a habitat based on structural vegetation factors rather than floristic factors. Temporal separation of species was observed, asA. minimus were more active than Rattus species during daylight periods. There was evidence of micro-habitat selection by species, and structural vegetation factors were most commonly identified in statistical analyses as contributing towards selection by small mammal species. Following a theoretical modelling study three reintroduction trials were carried out near Anglesea during 1992-94. Individuals were subsequently radio tracked, and habitat relationships between the species in the small mammal community investigated. Although successful breeding of A, minimus occurred during the latter two trials, the subsequent fate of offspring was not determined. Invasive techniques required to adequately monitor young animals were considered potentially too damaging. Telemetry studies indicated a preference of A. minimus for short, wet heath vegetation. Structural vegetation factors were identified as being significant in discriminating between capture locations of species. Small scale and inexpensive trial reintroductions have yielded valuable additional data on this species and may be viewed as a useful tool in the conservation of other small native mammals.