A comparison of mark-recapture distance-sampling methods applied to aerial surveys of eastern grey kangaroos

Aerial surveys of kangaroos (Macropus spp.) in Queensland are used to make economically important judgements on the levels of viable commercial harvest. Previous analysis methods for aerial kangaroo surveys have used both mark-recapture methodologies and conventional distance-sampling analyses. Conventional distance sampling has the disadvantage that detection is assumed to be perfect on the transect line, while mark-recapture methods are notoriously sensitive to problems with unmodelled heterogeneity in capture probabilities. We introduce three methodologies for combining together mark-recapture and distance-sampling data, aimed at exploiting the strengths of both methodologies and overcoming the weaknesses. Of these methods, two are based on the assumption of full independence between observers in the mark-recapture component, and this appears to introduce more bias in density estimation than it resolves through allowing uncertain trackline detection. Both of these methods give lower density estimates than conventional distance sampling, indicating a clear failure of the independence assumption. The third method, termed point independence, appears to perform very well, giving credible density estimates and good properties in terms of goodness-of-fit and percentage coefficient of variation. Estimated densities of eastern grey kangaroos range from 21 to 36 individuals km-2, with estimated coefficients of variation between 11% and 14% and estimated trackline detection probabilities primarily between 0.7 and 0.9.

Hydrogen utilising bacteria from the forestomach of eastern grey (Macropus giganteus) and red (Macropus rufus) kangaroos

Reductive acetogenesis is an alternative to methanogenesis for removing hydrogen produced during enteric fermentation. In Australia, kangaroos have evolved an enlarged forestomach analogous to the rumen of sheep and cattle. However, unlike sheep and cattle, kangaroos produce very little methane from enteric fermentation. From samples of gut contents from five eastern grey and three red kangaroos, we were not able to detect methanogens using a PCR protocol, but did detect the formyltetrahydrofolate synthetase (FTHFS) gene (likely to be used for reductive acetogenesis) in all animals. Isolations to recover acetogens resulted in two different classes of hydrogen consuming bacteria being isolated. The first class consisted of acetogens that possessed the FTHFS gene, which except for Clostridium glycolicum, were not closely related to any previously cultured bacteria. The second class were not acetogens but consisted of enterobacteria (Escherichia coli and Shigella) that did not possess FTHFS genes but did utilise hydrogen and produce acetate. Enumeration of the acetogens containing the FTHFS gene by real-time PCR indicated that bacteria of the taxa designated YE257 were common to all the kangaroos whereas YE266/YE273 were only detected in eastern grey kangaroos. When present, both species occurred at densities above *106 cell equivalents per mL. C. glycolicum was not detected in the kangaroos and, unlike YE257 and YE266/273, is unlikely to play a major role in reductive acetogenesis in the foregut of kangaroos.

Increased reproductive success in older female red kangaroos and the impact of harvesting

Wildlife harvesting has a long history in Australia, including obvious examples of overexploitation. Not surprisingly, there is scepticism that commercial harvesting can be undertaken sustainably. Kangaroo harvesting has been challenged regularly at Administrative Appeals Tribunals and elsewhere over the past three decades. Initially, the concern from conservation groups was sustainability of the harvest. This has been addressed through regular, direct monitoring that now spans > 30 years and a conservative harvest regime with a low risk of overharvest in the face of uncertainty. Opposition to the harvest now continues from animal rights groups whose concerns have shifted from overall harvest sustainability to side effects such as animal welfare, and changes to community structure, genetic composition and population age structure. Many of these concerns are speculative and difficult to address, requiring expensive data. One concern is that older females are the more successful breeders and teach their daughters optimal habitat and diet selection. The lack of older animals in a harvested population may reduce the fitness of the remaining individuals; implying population viability would also be compromised. This argument can be countered by the persistence of populations under harvesting without any obvious impairment to reproduction. Nevertheless, an interesting question is how age influences reproductive output. In this study, data collected from a number of red kangaroo populations across eastern Australia indicate that the breeding success of older females is up to 7-20% higher than that of younger females. This effect is smaller than that of body condition and the environment, which can increase breeding success by up to 30% and 60% respectively. Average age of mature females in a population may be reduced from 9 to 6 years old, resulting in a potential reduction in breeding success of 3-4%. This appears to be offset in harvested populations by improved condition of females from a reduction in kangaroo density. There is an important recommendation for management. The best insurance policy against overharvest and unwanted side effects is not research, which could be never-ending. Rather, it is a harvest strategy that includes safeguards against uncertainty such as harvest reserves, conservative quotas and regular monitoring. Research is still important in fine tuning that strategy and is most usefully incorporated as adaptive management where it can address the key questions on how populations respond to harvesting.

Fencing artificial waterpoints failed to influence density and distribution of red kangaroos (Macropus rufus)

Provision of artificial waterpoints in Australian rangelands has resulted in an increase in the range and density of kangaroos. At high densities, kangaroos can inhibit vegetation regeneration, particularly in some protected areas where harvesting is prohibited. Fencing off waterpoints has been proposed to limit these impacts. Our aim was to determine whether fencing off waterpoints during a drought (when kangaroos would be especially water-limited) would influence the density and distribution of red kangaroos (Macropus rufus). Two waterpoints were fenced within the first 6 months of the 27-month study and a further two waterpoints were kept unfenced as controls in Idalia National Park, western Queensland. We estimated kangaroo densities around waterpoints from walked line-transect counts, and their grazing distribution from dung-pellet counts. Fencing off waterpoints failed to influence either the density or distribution up to 4 km from the waterpoints. Our results indicate that food availability, rather than the location of waterpoints, determines kangaroo distribution. Few areas in the rangelands are beyond kangaroos' convenient reach from permanent waterpoints. Therefore, fencing off waterpoints without explicitly considering the spatial context in relation to other available water sources will fail to achieve vegetation regeneration.

Observation of a novel babesia spp. in eastern grey kangaroos (macropus giganteus) in australia

The roles and epidemiological features of tick-borne protozoans are not well elicited in wildlife. Babesia spp. are documented in many domestic animals, including cattle, horses, pigs, dogs and cats. Three cases affecting eastern grey kangaroos are described. The kangaroos exhibited neurological signs, depression and marked anaemia, and microscopic examination of blood smears revealed intraerythrocytic piroplasms. One to seven intraerythrocytic spherical, oval, pyriform and irregularly-shaped parasites consistent with Babesia spp. were seen in the blood smears and the percentage of infected erythrocytes was estimated to be approximately 7% in each case. Data suggest that the tick vector for this kangaroo Babesia sp. is a Haemaphysalis species. For Case 2, ultrastructural examination of the erythrocytes of the renal capillaries showed parasites resembling Babesia spp. and 18 of 33 erythrocytes were infected. DNA sequencing of the amplified 18S rDNA confirmed that the observed intraerythrocytic piroplasms belong to the genus Babesia. The phylogenetic position of this new kangaroo Babesia sp. (de novo Babesia macropus), as a sister species to the new Australian woylie Babesia sp., suggests a close affinity to the described Afro-Eurasian species Babesia orientalis and Babesia occultans suggesting perhaps a common ancestor for the Babesia in kangaroos. © 2012 Australian Society for Parasitology.

Low-density koala (Phascolarctos cinereus) populations in the mulgalands of south-west Queensland. II. Distribution and diet

This study used faecal pellets to investigate the broadscale distribution and diet of koalas in the mulgalands biogeographic region of south-west Queensland. Koala distribution was determined by conducting faecal pellet searches within a 30-cm radius of the base of eucalypts on 149 belt transects, located using a multi-scaled stratified sampling design. Cuticular analysis of pellets collected ffom 22 of these sites was conducted to identify the dietary composition of koalas within the region. Our data suggest that koala distribution is concentrated in the northern and more easterly regions of the study area, and appears to be strongly linked with annual rainfall. Over 50% of our koala records were obtained from non-riverine communities, indicating that koalas in the study area are not primarily restricted to riverine communities, as bas frequently been suggested. Cuticular analysis indicates that more than 90% of koala diet within the region consists of five eucalypt species. Our data highlights the importance of residual Tertiary landforms to koala conservation in the region.

Low-density koala (Phascolarctos cinereus) populations in the mulgalands of south-west Queensland. IV. Abundance and conservation status

Urban encroachment on dense, coastal koala populations has ensured that their management has received increasing government and public attention. The recently developed National Koala Conservation Strategy calls for maintenance of viable populations in the wild. Yet the success of this, and other, conservation initiatives is hampered by lack of reliable and generally accepted national and regional population estimates. In this paper we address this problem in a potentially large, but poorly studied, regional population in the State that is likely to have the largest wild populations. We draw on findings from previous reports in this series and apply the faecal standing-crop method (FSCM) to derive a regional estimate of more than 59 000 individuals. Validation trials in riverine communities showed that estimates of animal density obtained from the FSCM and direct observation were in close agreement. Bootstrapping and Monte Carlo simulations were used to obtain variance estimates for our population estimates in different vegetation associations across the region. The most favoured habitat was riverine vegetation, which covered only 0.9% of the region but supported 45% of the koalas. We also estimated that between 1969 and 1995 -30% of the native vegetation associations that are considered as potential koala habitat were cleared, leading to a decline of perhaps 10% in koala numbers. Management of this large regional population has significant implications for the national conservation of the species: the continued viability of this population is critically dependent on the retention and management of riverine and residual vegetation communities, and future vegetation-management guidelines should be cognisant of the potential impacts of clearing even small areas of critical habitat. We also highlight eight management implications.

Seasonal resource availability and use by an endangered tropical mycophagous marsupial

This study highlights the importance of considering how seasonality of rainfall affects availability of resources and consequently species distributions within tropical ecosystems. The endangered northern bettong, Bettongia tropica Wakefield is thought to be restricted to habitats where seasonal availability of hypogeous fungi, their principal food resource, remains high. To test this hypothesis fungal abundance was quantified in the early wet, late wet, early dry and late dry seasons within known bettong habitat. A relationship was found between precipitation and fungal availability, with the abundance of hypogeous fungi being significantly lower in the late dry season. Fungal availability correlated strongly with the seasonal rainfall pattern determined from 74-year monthly means. This contrasts with a previous study where mycophagy, measured by faecal analysis, remained high across seasons presumably because of aseasonal rainfall during that study period. Alloteropsis semialata R.Br. (cockatoo grass) use by bettongs increased significantly during the period of low fungal availability. This suggests that the importance of cockatoo grass as an alternative food resource during annual and extended dry periods has previously been underestimated. With the frequency and intensity of drought expected to increase with global climate change, these findings have significant implications for bettong management. The important and possibly equivalent dependence of B. tropica on both hypogeous fungi and A. semialata helps to explain their habitat preference and identifies this species as a true ecotonal specialist.

Frequency and precision of aerial surveys for kangaroo management

The appropriate frequency and precision for surveys of wildlife populations represent a trade-off between survey cost and the risk of making suboptimal management decisions because of poor survey data. The commercial harvest of kangaroos is primarily regulated through annual quotas set as proportions of absolute estimates of population size. Stochastic models were used to explore the effects of varying precision, survey frequency and harvest rate on the risk of quasiextinction for an arid-zone and a more mesic-zone kangaroo population. Quasiextinction probability increases in a sigmoidal fashion as survey frequency is reduced. The risk is greater in more arid regions and is highly sensitive to harvest rate. An appropriate management regime involves regular surveys in the major harvest areas where harvest rate can be set close to the maximum sustained yield. Outside these areas, survey frequency can be reduced in relatively mesic areas and reduced in arid regions when combined with lowered harvest rates. Relative to other factors, quasiextinction risk is only affected by survey precision (standard error/mean × 100) when it is >50%, partly reflecting the safety of the strategy of harvesting a proportion of a population estimate.

Stochastic demography and population dynamics in the red kangaroo Macropus rufus

1. Many organisms inhabit strongly fluctuating environments but their demography and population dynamics are often analysed using deterministic models and elasticity analysis, where elasticity is defined as the proportional change in population growth rate caused by a proportional change in a vital rate. Deterministic analyses may not necessarily be informative because large variation in a vital rate with a small deterministic elasticity may affect the population growth rate more than a small change in a less variable vital rate having high deterministic elasticity. 2. We analyse a stochastic environment model of the red kangaroo (Macropus rufus), a species inhabiting an environment characterized by unpredictable and highly variable rainfall, and calculate the elasticity of the stochastic growth rate with respect to the mean and variability in vital rates. 3. Juvenile survival is the most variable vital rate but a proportional change in the mean adult survival rate has a much stronger effect on the stochastic growth rate. 4. Even if changes in average rainfall have a larger impact on population growth rate, increased variability in rainfall may still be important also in long-lived species. The elasticity with respect to the standard deviation of rainfall is comparable to the mean elasticities of all vital rates but the survival in age class 3 because increased variation in rainfall affects both the mean and variability of vital rates. 5. Red kangaroos are harvested and, under the current rainfall pattern, an annual harvest fraction of c. 20% would yield a stochastic growth rate about unity. However, if average rainfall drops by more than c. 10%, any level of harvesting may be unsustainable, emphasizing the need for integrating climate change predictions in population management and increase our understanding of how environmental stochasticity translates into population growth rate.

Using harvest statistics to monitor temporal variation in kangaroo density and harvest rate

Management of the commercial harvest of kangaroos relies on quotas set annually as a proportion of regular estimates of population size. Surveys to generate these estimates are expensive and, in the larger states, logistically difficult; a cheaper alternative is desirable. Rainfall is a disappointingly poor predictor of kangaroo rate of increase in many areas, but harvest statistics (sex ratio, carcass weight, skin size and animals shot per unit time) potentially offer cost-effective indirect monitoring of population abundance (and therefore trend) and status (i.e. under-or overharvest). Furthermore, because harvest data are collected continuously and throughout the harvested areas, they offer the promise of more intensive and more representative coverage of harvest areas than aerial surveys do. To be useful, harvest statistics would need to have a close and known relationship with either population size or harvest rate. We assessed this using longterm (11-22 years) data for three kangaroo species (Macropus rufus, M. giganteus and M. fuliginosus) and common wallaroos (M. robustus) across South Australia, New South Wales and Queensland. Regional variation in kangaroo body size, population composition, shooter efficiency and selectivity required separate analyses in different regions. Two approaches were taken. First, monthly harvest statistics were modelled as a function of a number of explanatory variables, including kangaroo density, harvest rate and rainfall. Second, density and harvest rate were modelled as a function of harvest statistics. Both approaches incorporated a correlated error structure. Many but not all regions had relationships with sufficient precision to be useful for indirect monitoring. However, there was no single relationship that could be applied across an entire state or across species. Combined with rainfall-driven population models and applied at a regional level, these relationships could be used to reduce the frequency of aerial surveys without compromising decisions about harvest management.

Reassessing spatial and temporal dynamics of kangaroo populations

Long-running datasets from aerial surveys of kangaroos (Macropus giganteus, Macropus [uliginosus, Macropus robustus and Macropus rufus) across Queensland, New South Wales and South Australia have been analysed, seeking better predictors of rates of increase which would allow aerial surveys to be undertaken less frequently than annually. Early models of changes in kangaroo numbers in response to rainfall had shown great promise, but much variability. We used normalised difference vegetation index (NDVI) instead, reasoning that changes in pasture condition would provide a better predictor than rainfall. However, except at a fine scale, NDVI proved no better; although two linked periods of rainfall proved useful predictors of rates of increase, this was only in some areas for some species. The good correlations reported in earlier studies were a consequence of data dominated by large droughtinduced adult mortality, whereas over a longer time frame and where changes between years are less dramatic, juvenile survival has the strongest influence on dynamics. Further, harvesting, density dependence and competition with domestic stock are additional and important influences and it is now clear that kangaroo movement has a greater influence on population dynamics than had been assumed. Accordingly, previous conclusions about kangaroo populations as simple systems driven by rainfall need to be reassessed. Examination of this large dataset has permitted descriptions of shifts in distribution of three species across eastern Australia, changes in dispersion in response to rainfall, and an evaluation of using harvest statistics as an index of density and harvest rate. These results have been combined into a risk assessment and decision theory framework to identify optimal monitoring strategies.

Family-level relationships among the Australasian marsupial “herbivores” (Diprotodontia: Koala, wombats, kangaroos and possums)

The marsupial order Diprotodontia includes 10 extant families, which occupy all terrestrial habitats across Australia and New Guinea and have evolved remarkable dietary and locomotory diversity. Despite considerable attention, the interrelations of these families have for the most part remained elusive. In this study, we separately model mitochondrial RNA and protein-coding sequences in addition to nuclear protein-coding sequences to provide near-complete resolution of diprotodontian family-level phylogeny. We show that alternative topologies inferred in some previous studies are likely to be artifactual, resulting from branch-length and compositional biases. Subordinal groupings resolved herein include Vombatiformes (wombats and koala) and Phalangerida, which in turn comprises Petauroidea (petaurid gliders and striped, feathertail, ringtail and honey possums) and a clade whose plesiomorphic members possess blade-like premolars (phalangerid possums, kangaroos and their allies and most likely, pygmy possums). The topology resolved reveals ecological niche structuring among diprotodontians that has likely been maintained for more than 40 million years.

The public's knowledge of and support for conservation of Australia's tree-kangaroos and other animals

After providing background on Dendrolagus species in Australia, two consecutive surveys of Brisbane's residents are used to assess public knowledge of tree-kangaroos and the stated degree of support for their conservation in Australia. The responses of participants in Survey I are based on their pre-survey knowledge of wildlife. The same set of participants completed Survey II after being provided with additional information on all the wildlife species mentioned in Survey I. Changes in the attitudes of respondents and their degree of support for the protection and conservation of Australia's tree-kangaroos are measured, including changes in their contingent valuations and stated willingness to provide financial support for such conservation. Reasons for wanting to protect tree-kangaroos are specified and analysed. Furthermore, changes that occur in the relative importance of these reasons with increased knowledge are also examined. Support for the conservation of tree-kangaroos is found to rise with the additional knowledge supplied about all species and is compared with variations in support for protection of other mammals. Support for the conservation of Australia's less well known tropical mammals is found to increase relative to better known mammals (icons) present in temperate areas, such as koalas and red kangaroos. Possible implications of the results for government conservation policies in Australia are examined.

Evolution of sexual segregation in mammalian herbivores: kangaroos as marsupial models

Sexual segregation is best known in sexually dimorphic ungulates. Many hypotheses have been proposed to explain the evolution of sexual segregation in ungulates, but all are reducible to the influence of two factors: body size and sex-specific reproductive strategy. Definitive tests of these hypotheses are lacking in ungulates because these factors are confounded, all males being somewhat larger than females. Kangaroos represent a parallel radiation of terrestrial herbivores, but their populations are composed of a spectrum of adult body sizes, ranging from small males the same size as females to large males more than twice the size. We exploited this heteromorphism to assess the independent influences of size and sex in these ungulate analogues. We conducted a preliminary study of western grey kangaroos (Macropus fuliginosus) in north-western Victoria, Australia. Adult males predominately occupied grassland habitat, whereas females occurred mostly in lakebed, woodland and shrubland. Single-sex groups occurred more often than expected during the non-mating season. The diet of large males had the highest proportion of grass, and females had the least. These initial results indicate that both size and sex influence segregation in this species, confirming the worth of kangaroos as marsupial models for research into the evolution of sexual segregation.