Sexual segregation in western grey kangaroos: testing alternative evolutionary hypotheses

In sexually dimorphic ungulates, sexual segregation is hypothesized to have evolved because of sex-specific differences in body size and/or reproductive strategies. We tested these alternative hypotheses in kangaroos, which are ecological analogues of ungulates. Kangaroos exhibit a wide range of body sizes, particularly among mature males, and so the effects of body size and sex can be distinguished. We tested predictions derived from these hypotheses by comparing the distribution of three sex–sex size classes of western grey kangaroos Macropus fuliginosus, in different habitats, and the composition of groups of kangaroos, across seasons. In accordance with the predation risk-reproductive strategy hypothesis, during the non-breeding season, females, which were more susceptible to predation than larger males, and were accompanied by vulnerable young-at-foot, were over-represented in secure habitats. Large males, which were essentially immune to predation, occurred more often than expected in nutrient-rich habitat, and small males, which faced competing demands of predator avoidance and feeding, were intermediate between females and large males in their distribution across habitats. During the breeding season, females continued to be over-represented in secure habitats when their newly emerged pouch young were most vulnerable to predation. All males occupied these same habitats to maximize their chances of securing mates. Consistent with the social hypotheses, groups composed of individuals of the same sex, irrespective of body size, were over-represented in the population during the non-breeding season, while during the breeding season all males sought females so that mixed-sex groups predominated. These results indicate that body size and reproductive strategies are both important, yet independent, factors influencing segregation in western grey kangaroos.

A retrospective study of Babesia macropus associated with morbidity and mortality in eastern grey kangaroos (Macropus giganteus) and agile wallabies (Macropus agilis)

This is a retrospective study of 38 cases of infection by Babesia macropus, associated with a syndrome of anaemia and debility in hand-reared or free-ranging juvenile eastern grey kangaroos (Macropus giganteus) from coastal New South Wales and south-eastern Queensland between 1995 and 2013. Infection with B. macropus is recorded for the first time in agile wallabies (Macropus agilis) from far north Queensland. Animals in which B. macropus infection was considered to be the primary cause of morbidity had marked anaemia, lethargy and neurological signs, and often died. In these cases, parasitised erythrocytes were few or undetectable in peripheral blood samples but were sequestered in large numbers within small vessels of visceral organs, particularly in the kidney and brain, associated with distinctive clusters of extraerythrocytic organisms. Initial identification of this piroplasm in peripheral blood smears and in tissue impression smears and histological sections was confirmed using transmission electron microscopy and molecular analysis. Samples of kidney, brain or blood were tested using PCR and DNA sequencing of the 18S ribosomal RNA and heat shock protein 70 gene using primers specific for piroplasms. The piroplasm detected in these samples had 100 sequence identity in the 18S rRNA region with the recently described Babesia macropus in two eastern grey kangaroos from New South Wales and Queensland, and a high degree of similarity to an unnamed Babesia sp. recently detected in three woylies (Bettongia penicillata ogilbyi) in Western Australia.

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.

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.

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.

The impact of grazing by eastern grey kangaroos (macropus giganteus) on vegetation recovery after fire at Reef Hills regional park, Victoria

In southeastern Australia ecological burning is frequently used to maintain a number of plant and animal populations. However, many of these prescribed fires are small, and may focus intense grazing activity on new regrowth. At Reef Hills Regional Park, Victoria shrub species have senesced, presumably due to the absence of fire. Ecological burning may be necessary to promote regeneration, however, the population density of the Eastern Grey Kangaroo (Macropus giganteus) is high (approx 38 per km2), and grazing pressure presents a significant risk to postfire vegetation recovery. An assessment of grazing patterns and their effects on postfire recovery was carried out at Reef Hills Regional Park through grazing exclusion plots. Preferential grazing by Eastern Grey Kangaroos occurred on small burnt plots compared to adjacent unburnt areas as determined by faecal pellet counts. On burnt areas, there was a significant reduction in shrub diversity on grazed plots compared to ungrazed plots. Most observations of kangaroos were of animals grazing on farmland surrounding the Park, and it is likely that any burning might shift grazing from farmland to burnt areas when new growth occurs. This needs to be considered before any ecological burn plan is applied to manage vegetation communities, particularly if the plan requires small areas to be burnt. We recommended that a large area up to 200 ha area be burnt and monitored to determine whether burning larger areas disperses grazing pressure from macropods to a level where impacts on vegetation are reduced and localized plant extinctions do not occur.

Do harvest refuges buffer kangaroos against evolutionary responses to selective harvesting?

There is a wealth of literature documenting a directional change of body size in heavily harvested populations. Most of this work concentrates on aquatic systems, but terrestrial populations are equally at risk. This paper explores the capacity of harvest refuges to counteract potential effects of size-selective harvesting on the allele frequency,of populations. We constructed a stochastic, individual-based model parameterized with data on red kangaroos. Because we do not know which part of individual growth would change in the course of natural selection, we explored the effects of two alternative models of individual growth in which alleles affect either the growth rate or the maximum size. The model results show that size-selective harvesting can result in significantly smaller kangaroos for a given age when the entire population is subject to harvesting. In contrast, in scenarios that include dispersal from harvest refuges, the initial allele frequency remains virtually unchanged.

Estimating the abundance of eastern grey kangaroos (Macropus giganteus) in south-eastern New South Wales, Australia

To provide an estimate of kangaroo numbers for harvest management, a survey was designed for an area of 29500 km(2) encompassing the agricultural and grazing lands of the Braidwood, Cooma, Goulburn, Gundagai and Yass Rural Lands Protection Board (RLPB) districts in south-east New South Wales. An aerial survey using a helicopter was considered more efficient than ground survey because of the size of the area, relatively high relief and dense tree cover, and the need for regular monitoring. Tree cover and landscape relief was used to stratify the five RLPB districts into areas of probable high, medium and low kangaroo density. Kangaroo density estimated from helicopter surveys conducted in the Northern Tablelands of New South Wales was used to suggest densities and thereby allocate survey effort in each stratum. A survey comprising 735 km of transect line was conducted in winter 2003 with a target precision of 20%. The survey returned an estimate of 286600 32300 eastern grey kangaroos for the whole of the proposed south-east New South Wales kangaroo-management zone. In 2004, a trial harvest of slightly less than 15% of this estimate was taken. Success of the trial will be determined by the impact of harvesting on the population's dynamics, by landholder and industry participation, and by the capacity to monitor population size, harvest offtake and compliance with regulations.

The influence of non-climate predictors at local and landscape resolutions depends on the autecology of the species

Species distribution models have come under criticism for being too simplistic for making robust future forecasts, partly because they assume that climate is the main determinant of geographical range at large spatial extents and coarse resolutions, with non-climate predictors being important only at finer scales. We suggest that this paradigm might be obscured by species movement patterns. To explore this we used contrasting kangaroo (family Macropodidae) case studies: two species with relatively small, stable home ranges (Macropus giganteus and M.robustus) and three species with more extensive, adaptive ranging behaviour (M.antilopinus, M.fuliginosus and M.rufus). We predicted that non-climate predictors will be most influential to model fit and predictive performance at local spatial resolution for the former species and at landscape resolution for the latter species. We compared residuals autocovariate - boosted regression tree (RAC-BRT) model statistics with and without species-specific non-climate predictors (habitat, soil, fire, water and topography), at local- and landscape-level spatial resolutions (5 and 50km). As predicted, the influence of non-climate predictors on model fit and predictive performance (compared with climate-only models) was greater at 50 compared with 5km resolution for M.rufus and M.fuliginosus and the opposite trend was observed for M.giganteus. The results for M.robustus and M.antilopinus were inconclusive. Also notable was the difference in inter-scale importance of climate predictors in the presence of non-climate predictors. In conclusion, differences in autecology, particularly relating to space use, may contribute to the importance of non-climate predictors at a given scale, not model scale per se. Further exploration of this concept across a range of species is encouraged and findings may contribute to more effective conservation and management of species at ecologically meaningful scales. © 2014 Ecological Society of Australia.

Of sheep and rain: large-scale population dynamics of the red kangaroo

1. We analysed time-series data from populations of red kangaroos (Macropus rufus, Desmarest) inhabiting four areas in the pastoral zone of South Australia. We formulated a set of a priori models to disentangle the relative effects of the covariates: rainfall, harvesting, intraspecific competition, and domestic herbivores, on kangaroo population-growth rate. 2. The statistical framework allowed for spatial variation in the growth-rate parameters, response to covariates, and environmental variability, as well as spatially correlated error terms due to shared environment. 3. The most parsimonious model included all covariates but no area-specific parameter values, suggesting that kangaroo densities respond in the same way to the covariates across the areas. 4. The temporal dynamics were spatially correlated, even after taking into account the potentially synchronizing effect of rainfall, harvesting and domestic herbivores. 5. Counter-intuitively, we found a positive rather than negative effect of domestic herbivore density on the population-growth rate of kangaroos. We hypothesize that this effect is caused by sheep and cattle acting as a surrogate for resource availability beyond rainfall. 6. Even though our system is well studied, we must conclude that approximating resources by surrogates such as rainfall is more difficult than previously thought. This is an important message for studies of consumer-resource systems and highlights the need to be explicit about population processes when analysing population patterns.

Should managed populations be monitored every year?

We often need to estimate the size of wild populations to determine the appropriate management action, for example, to set a harvest quota. Monitoring is usually planned under the assumption that it must be carried out at fixed intervals in time, typically annually, before the harvest quota is set. However, monitoring can be very expensive, and we should weigh the cost of monitoring against the improvement that it makes in decision making. A less costly alternative to monitoring annually is to predict the population size using a population model and information from previous surveys. In this paper, the problem of monitoring frequency is posed within a decision-theory framework. We discover that a monitoring regime that varies according to the state of the system call outperform fixed-interval monitoring This idea is illustrated using data for a red kangaroo (Macropits rufus) population in South Australia. Whether or not one should monitor in a given year is dependent on the estimated population density in the previous year, the uncertainty in that population estimate, and past rainfall. We discover that monitoring is-important when a model-based prediction of population density is very uncertain. This may occur if monitoring has not taken place for several years, or if rainfall has been above average. Monitoring is also important when prior information suggests that the population is near a critical threshold in population abundance. However, monitoring is less important when the optimal management action would not be altered by new information.

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.

Resource use by the Eastern Grey Kangaroo and the Black Wallaby in a managed remnant woodland community

This work studies the interactions and relationships that exist between Eastern Grey Kangaroos and Black Wallabies in their utilisation of spatial and trophic resources in a managed remnant woodland community. The thesis provides a closer understanding of the way in which these species impact upon their habitats. The Coranderrk Reserve, near Healesville in eastern Victoria, Australia was the study site. A floristic analysis of the communities of the study site was conducted. This consisted of plant biomass measurements, estimates of plant abundance and cover determination. Faecal pellets from Eastern Grey Kangaroos and Black Wallabies were collected from ten vegetation communities during three different plant productivity periods. The spatial and temporal distribution of the animals was identified by analysing the frequency of occurrence of faecal pellets in the various communities. The use of faecal pellet density as a measure of habitat utilisation was examined. Eastern Grey Kangaroos utilised communities which were characterised by the presence of a dense grassy statum. Black Wallabies were able to utilise all of the communities of the study site regardless of their floristic composition. A reference herbarium of the leaf epidermis of 233 possible forage plant species was accumulated. These epidermal specimens were prepared for Confocal Scanning Laser Microscopy. The information gained was enhanced and stored digitally. Diagnostic information critical for the identification of plant epidermal fragments was assembled into a computer database. This was used to assist in the recognition of unknown epidermal fragments in macropodid faeces. These epidermal plant recognition techniques enabled a list of the contents of Eastern Grey Kangaroo and Black Wallaby faeces during the sampling periods in the individual communities, to be accumulated. Eastern Grey Kangaroos utilised forage which consisted largely of grass and their diets were similar regardless of their feeding sites or the time of the year. Black Wallaby diets were heterogenous with wide variations over space and time observed. The implications of these findings for current wildlife management practices were considered. Black Wallaby and Eastern Grey Kangaroo herbivory have significant impacts on ecosystem integrity. Management strategies should seek to establish ecologically sustainable populations of both species in remnant woodlands where conservation values are important.

Interspecific and geographic variation in the diets of sympatric carnivores: dingoes/wild dogs and red foxes in south-eastern Australia

Dingoes/wild dogs (Canis dingo/familiaris) and red foxes (Vulpes vulpes) are widespread carnivores in southern Australia and are controlled to reduce predation on domestic livestock and native fauna. We used the occurrence of food items in 5875 dingo/wild dog scats and 11,569 fox scats to evaluate interspecific and geographic differences in the diets of these species within nine regions of Victoria, south-eastern Australia. The nine regions encompass a wide variety of ecosystems. Diet overlap between dingoes/wild dogs and foxes varied among regions, from low to near complete overlap. The diet of foxes was broader than dingoes/wild dogs in all but three regions, with the former usually containing more insects, reptiles and plant material. By contrast, dingoes/wild dogs more regularly consumed larger mammals, supporting the hypothesis that niche partitioning occurs on the basis of mammalian prey size. The key mammalian food items for dingoes/wild dogs across all regions were black wallaby (Wallabia bicolor), brushtail possum species (Trichosurus spp.), common wombat (Vombatus ursinus), sambar deer (Rusa unicolor), cattle (Bos taurus) and European rabbit (Oryctolagus cuniculus). The key mammalian food items for foxes across all regions were European rabbit, sheep (Ovis aries) and house mouse (Mus musculus). Foxes consumed 6.1 times the number of individuals of threatened Critical Weight Range native mammal species than did dingoes/wild dogs. The occurrence of intraguild predation was asymmetrical; dingoes/wild dogs consumed greater biomass of the smaller fox. The substantial geographic variation in diet indicates that dingoes/wild dogs and foxes alter their diet in accordance with changing food availability. We provide checklists of taxa recorded in the diets of dingoes/wild dogs and foxes as a resource for managers and researchers wishing to understand the potential impacts of policy and management decisions on dingoes/wild dogs, foxes and the food resources they interact with.

The influence of animal mobility on the assumption of uniform distances in aerial line-transect surveys

Line-transect distance sampling is a widely used method for estimating animal density from aerial surveys. Analysis of line-transect distance data usually relies on a requirement that the statistical distribution of distances of animal groups from the transect line is uniform. We show that this requirement is satisfied by the survey design if all other assumptions of distance sampling hold, but it can be violated by consistent survey problems such as responsive movement of the animals towards or away from the observer. We hypothesise that problems with the uniform requirement are unlikely to be encountered for immobile taxa, but might become substantial for species of high mobility. We test evidence for non-uniformity using double-observer distance data from two aerial surveys of five species with a spectrum of mobility capabilities and tendencies. No clear evidence against uniformity was found for crabeater seals or emperor penguins on the pack-ice in East Antarctica, while minor non-uniformity consistent with responsive movement up to 30 m was found for Adelie penguins. Strong evidence of either non-uniformity or a failure of the capture-recapture validating method was found for eastern grey kangaroos and red kangaroos in Queensland.