William Ellis

Phylogeographic differentiation in the mitochondrial control region in the koala, Phascolarctos cinereus (Goldfuss 1817)

The koala, Phascolarctos cinereus, is a geographically widespread species endemic to Australia, with three currently recognized subspecies: P.c. adustus, P.c. cinereus, and P.c. victor. Intraspecific variation in the mitochondrial DNA (mtDNA) control region was examined in over 200 animals from 16 representative populations throughout the species’ range. Eighteen different haplotypes were defined in the ≈ 860 bp mtDNA control region, as determined by heteroduplex analysis/temperature gradient gel electrophoresis (HDA/TGGE). Any single population typically possessed only one or two haplotypes yielding an average within‐population haplotypic diversity of 0.180 ± 0.003, and nucleotide diversity of 0.16%. Overall, mtDNA control region sequence diversity between populations averaged 0.67%, and ranged from 0% to 1.56%. Nucleotide divergence between populations averaged 0.51%, and ranged from 0% to 1.53%. Neighbour‐joining methods revealed limited phylogenetic distinction between geographically distant populations of koalas, and tentative support for a single evolutionarily significant unit (ESU). This is consistent with previous suggestions that the morphological differences formalized by subspecific taxonomy may be interpreted as clinal variation. Significant differentiation in mtDNA‐haplotype frequencies between localities suggested that little gene flow currently exists among populations. When combined with microsatellite analysis, which has revealed substantial differentiation among koala populations, we conclude that the appropriate short‐term management unit (MU) for koalas is the local population.

Tree use, diet and home range of the koala (Phascolarctos cinereus) at Blair Athol, central Queensland

Free-ranging koalas (Phascolarctos cinereus) were monitored by means of radio-relocation in the area of Blair Athol Coal Mine and surrounding properties. Daytime tree use, home range and diet of these koalas was determined in spring and autumn, as was the leaf moisture composition of potential fodder species. Koalas used on average 93 (male koalas) and 56 (female koalas) trees during the period of observation, occupying home ranges of 135 and 101 ha respectively. Mean sightings per tree were 1.19 for both males and females and home-range sizes were not significantly different between sexes or seasons. Koalas were observed returning to previously used daytime roosting trees infrequently (<12%). Although koalas were observed roosting in trees of the species that they ate, proportional species representation in the diet of these koalas during spring and autumn did not accurately reflect concurrent observations of their daytime tree-roosting behaviour. Koalas were observed to utilise non-fodder species for daytime roosting, and patterns of daytime tree use and diet selection varied between individuals inhabiting adjacent communities within the site. Leaf moisture of tree species represented in koala diets was greater in autumn than spring.

Genetic variation and structuring in the threatened koala populations of Southeast Queensland

Habitat fragmentation can act to cause reproductive isolation between conspecifics and undermine species’ persistence, though most studies have reported the genetic condition of populations that have already declined to a very small size. We examined genetic diversity within the vulnerable, declining koala (Phascolarctos cinereus) population in Southeast Queensland, Australia to determine the genetic impact of ongoing threatening processes. Five hundred and twelve koalas from ten Southeast Queensland Local Government Areas on the mainland and one island were genotyped at six polymorphic microsatellite loci. Based on Bayesian cluster analysis incorporating spatial data, the regional koala population was subdivided into six clusters, with location of major roads and rivers appearing to be consistent with being barriers to gene flow. The distribution of mtDNA control region haplotypes identified distinct coastal and inland clades suggesting that historically there was gene flow between koalas along the coast (though little interchange between coastal and inland animals). In contrast, koalas from the Koala Coast (Brisbane City, Logan City and Redland Shire) were shown by microsatellite analysis to be genetically distinct from adjacent areas. It is likely, therefore, that more recent reductions in population size and restricted gene flow through urbanisation have contributed to the genetic differentiation of koalas in the Koala Coast region.

Spatiotemporal dynamics of habitat use by koalas: the checkerboard model

Animal movement patterns and use of space depend upon food and nonfood resources, as well as conspecific and heterospecific interactions, but models of habitat use often neglect to examine multiple factors and rarely include marsupials. We studied habitat use in an Australian population of koalas (Phascolarctos cinereus) over a 6-year period in order to determine how koalas navigate their environment and partition limited patchy food and nonfood resources. Tree selection among koalas appears to be mediated by folar chemistry, but nonfood tree selection exerts a major impact on home range use due to thermoregulatory constraints. Koalas moved on a daily basis, during both day and night, but daytime resting site was not necessarily in the same location as nighttime feeding site. Koalas had substantial home range overlap in the near absence of resource sharing with less than 1% of trees located in areas of overlap used by multiple koalas. We suggest that koala spatiotemporal distribution and habitat use are probably based upon a community structure of individuals, with a checkerboard model best describing overlap in home range area but not in resource use. Nonfood refugia and social networks should be incorporated into models of animal range and habitat use.

Female koalas prefer bellows in which lower formants indicate larger males

Despite an extensive literature on the role of acoustic cues in mate choice little is known about the specific vocal traits that female mammals prefer. We used resynthesis techniques and playback experiments to examine the behavioural responses of oestrous female koalas, Phascolarctos cinereus, to male bellows in which a specific acoustic cue to body size, the formants, were modified to simulate callers of different body size. Oestrous females looked longer towards, and spent more time in close proximity to, loudspeakers broadcasting bellows simulating larger male koalas. These findings suggest that female koalas use formants (key components of human speech) to select larger males as mating partners, and represent the first evidence of a marsupial mating preference based on a vocal signal. More generally, these results indicate that intersexual selection pressures to lower formants and exaggerate size are present in a marsupial species, raising interesting questions about the evolutionary origins of formant perception.

Climate change and the koala Phascolarctos cinereus: water and energy

We studied two groups of koalas during a drought in central Queensland to investigate potential impacts of climatic variability on the physiology and behaviour of this species. The tree use, water turnover, field metabolic rate and diet of koalas during autumn and spring were compared to a similar study of koalas in summer and winter, also in central Queensland, to generate a seasonal picture of the response of koalas to climatic variation. We also compared the microclimate temperature of a range of food and non-food tree species against daily ambient temperatures, to examine the benefit to koalas of using of non-food species. Field metabolic rate, adjusted for body mass, was significantly higher in spring than autumn and there was no difference between males and females. Neither females with pouch young nor those with back young had significantly different FMR to that of females without young, confirming that koalas may compartmentalize energy demands during lactation. Estimations of theoretical water in...

Breeding dynamics of koalas in open woodlands

The spatial and breeding dynamics of koalas in sub-tropical woodlands at Blair Athol in central Queensland were intensively monitored between 1993 and 1998. Genetic relationships among koalas at this locality were studied to determine the breeding dynamics of males, including whether ‘resident’ or ‘transient’ males dominate as sires. Males and females were radio-collared and tracked periodically throughout each year of the study. Genotypes from hypervariable microsatellite loci identified uniquely all individuals and were used to analyse parentage as well as to determine population genetic parameters when compared with other regional localities. Koalas at Blair Athol comprise a population in genetic equilibrium. Gene diversity estimates show the population to be similar to other populations found in similar habitat in the region, and estimates of genetic differentiation among four regional populations showed that gene flow conforms to a model of isolation by distance. Analysis of parentage found that both resident and transient males sired about equal numbers of offspring. Familial DNA analysis revealed multiple paternities of successive young in this population. The conclusion from this study is that ‘resident’ status among males does not confer any advantage for parentage.

The use of faecal cuticle examination to determine the dietary composition of koalas

Captive koalas were provided with specified proportions of known browse Species. Faecal pellets collected from each koala were analysed for cuticle fragments to determine the accuracy of this method for estimating the species composition of the diet Of koalas. Food species were consistently observed in faecal pellets collected 34 h after presentation of browse and remained represented in pellets collected up to 154h post-feeding. Faecal pellet examination provided an accurate means of describing the species composition of the diet of the koalas for the species of browse offered.

Perception of size-related formant information in male koalas (Phascolarctos cinereus)

Advances in bioacoustics allow us to study the perceptual and functional relevance of individual acoustic parameters. Here, we use re-synthesised male koala bellows and a habituation–dishabituation paradigm to test the hypothesis that male koalas are sensitive to shifts in formant frequencies corresponding to the natural variation in body size between a large and small adult male. We found that males habituated to bellows, in which the formants had been shifted to simulate a large or small male displayed a significant increase in behavioural response (dishabituation) when they were presented with bellows simulating the alternate size variant. The rehabituation control, in which the behavioural response levels returned to that of the last playbacks of the habituation phase, indicates that this was not a chance increase in response levels. Our results provide clear evidence that male koalas perceive and attend to size-related formant information in their own species-specific vocalisations and suggest that formant perception is a widespread ability shared by marsupials and placental mammals, and perhaps by vertebrates more widely.

Body size and sexual selection in the koala

Sexual selection is often characterized by polygynous breeding systems, size dimorphism, and skewed operational sex ratios. Koalas are sexually dimorphic in multiple domains, yet are absent from the literature on sexual selection and the structure of their mating system is unclear. We provide the first documentation of the strength of sexual selection in koalas by using microsatellite markers to identify sires. We combine the genetic data with morphological data in order to assess the role of body size in regulating reproductive output. During our 4-year study, 37% of males were identified as possible sires. Males were significantly larger than females, with sires heavier than non-sires. Male body mass correlated with annual reproductive output, with Crow’s Index of Opportunity for Selection revealing that variation in male reproductive success was threefold higher than that of females. Since it appears that male koalas rarely engage in physical confrontations over access to females, size dimorphism could be based upon non-agonistic competition and/or female mate choice. We propose that size dimorphism in koalas evolved as a consequence of endurance rivalry promoting vocal sexual advertisements that attract females. We suggest that female choice is a key mediator of male reproductive output.