Habitat requirements for the conservation of arboreal marsupials in dry sclerophyll forests of Southeast Queensland, Australia

The habitat requirements of arboreal marsupials were investigated in the dry sclerophyll forests of southeast Queensland, Australia. Species richness and abundance of arboreal marsupials was correlated to the proportion of total stand basal area occupied by lemon-scented gum (Corymbia citriodora), the height of the tallest trees, and density of hollow-bearing trees. The first two factors suggested that the most productive forests were also the most suitable habitats for arboreal marsupials. Importantly, the number of hollow-bearing trees was a significant factor in determining species richness and abundance of arboreal marsupials in this study, with the maximum number of species reached at sites containing greater than or equal to4 hollow-bearing trees/ha, and maximum abundance occurring at sites with :6 hollow-bearingtrees/ha. The proportion of C. citriodora was significant for the presence of the common brushtail possum (Trichosurus vulpecula), greater glider (Petauroides volans), and the yellow-bellied glider (Petaurus australis), while understory Acacia sp. density was important for the presence of the sugar glider (Petaurus breviceps). The yellow-bellied glider was also affected by two other variables: the density of hollow-bearing trees >50 cm diameter at breast height (dbh), and the time since the last logging. Current Codes of Practice regulating the density of hollow-bearing trees and silvicultural practices in state-owned timber production forests appear to provide adequate protection for arboreal marsupials, but the recently introduced increase in timber extraction rates within state forests may be detrimental to the animals. Also, protective prescriptions do not apply to the privately owned and leasehold estates, which contain the majority of the dry sclerophyll forests in southeast Queensland.

Disease, habitat fragmentation and conservation

Habitat loss and the resultant fragmentation of remaining habitat is the primary cause of loss of biological diversity. How do these processes affect the dynamics of parasites and pathogens? Hess has provided some important insights into this problem using metapopulation models for pathogens that exhibit 'S-I' dynamics; for example, pathogens such as rabies in which the host population may be divided into susceptible and infected individuals. A major assumption of Hess's models is that infected patches become extinct, rather than recovering and becoming resistant to future infections. In this paper, we build upon this framework in two different ways: first, we examine the consequences of including patches that are resistant to infection; second, we examine the consequences of including a second species of host that can act as a reservoir for the pathogen. Both of these effects are likely to be important from a conservation perspective. The results of both sets of analysis indicate that the benefits of corridors and other connections that allow species to disperse through the landscape far outweigh the possible risks of increased pathogen transmission. Even in the commonest case, where harmful pathogens are maintained by a common reservoir host, increased landscape connectance still allows greater coexistence and persistence of a threatened or endangered host.

Spatial structure and habitat variation in a grasshopper hybrid zone

A hybrid zone between the grasshoppers Chorthippus brunneus and C. jacobsi (Orthoptera: Acrididae) in northern Spain has been analyzed for variation in morphology and ecology. These species are readily distinguished by the number of stridulatory pegs on the hind femur. Both sexes are fully winged and inhabit disturbed habitats throughout the study area. We develop a maximum-likelihood approach to fitting a two-dimensional cline to geographical variation in quantitative traits and for estimating associations of population mean with local habitat. This method reveals a cline in peg number approximately 30 km south of the Picos de Europa Mountains that shows substantial deviations in population mean compared with the expectations of simple tension zone models. The inclusion of variation in local vegetation in the model explains a significant proportion of the residual variation in peg number, indicating that habitat-genotype associations contribute to the observed spatial pattern. However, this association is weak, and a number of populations continue to show strong deviations in mean even after habitat is included in the final model. These outliers may be the result of long-distance colonization of sites distant from the cline center or may be due to a patchy pattern of initial contact during postglacial expansion. As well as contrasting with the smooth hybrid zones described for Chorthippus parallelus, this situation also contrasts with the mosaic hybrid zones observed in Gryllus crickets and in parts of the hybrid zone between Bombina toad species, where habitat-genotype associations account for substantial amounts of among-site variation.

Mapping glider habitat in dry eucalypt forests for Montreal Process indicator 1.1: Fragmentation of forest types

Accurate habitat mapping is critical to landscape ecological studies such as required for developing and testing Montreal Process indicator 1.1e, fragmentation of forest types. This task poses a major challenge to remote sensing, especially in mixedspecies, variable-age forests such as dry eucalypt forests of subtropical eastern Australia. In this paper, we apply an innovative approach that uses a small section of one-metre resolution airborne data to calibrate a moderate spatial resolution model (30 m resolution; scale 1:50 000) based on Landsat Thematic Mapper data to estimate canopy structural properties in St Marys State Forest, near Maryborough, south-eastern Queensland. The approach applies an image-processing model that assumes each image pixel is significantly larger than individual tree crowns and gaps to estimate crown-cover percentage, stem density and mean crown diameter. These parameters were classified into three discrete habitat classes to match the ecology of four exudivorous arboreal species (yellowbellied glider Petaurus australis, sugar glider P. breviceps, squirrel glider P. norfolcensis , and feathertail glider Acrobates pygmaeus), and one folivorous arboreal marsupial, the greater glider Petauroides volans. These species were targeted due to the known ecological preference for old trees with hollows, and differences in their home range requirements. The overall mapping accuracy, visually assessed against transects (n = 93) interpreted from a digital orthophoto and validated in the field, was 79% (KHAT statistic = 0.72). The KHAT statistic serves as an indicator of the extent that the percentage correct values of the error matrix are due to ‘true’ agreement verses ‘chance’ agreement. This means that we are able to reliably report on the effect of habitat loss on target species, especially those with a large home range size (e.g. yellow-bellied glider). However, the classified habitat map failed to accurately capture the spatial patterning (e.g. patch size and shape) of stands with a trace or sub-dominance of senescent trees. This outcome makes the reporting of the effects of habitat fragmentation more problematic, especially for species with a small home range size (e.g. feathertail glider). With further model refinement and validation, however, this moderateresolution approach offers an important, cost eff e c t i v e advancement in mapping the age of dry eucalypt forests in the region.

Testing landscape metrics as indicators of habitat loss and fragmentation in continuous eucalypt forests (Queensland, Australia)

Landscape metrics are widely applied in landscape ecology to quantify landscape structure. However, many are poorly tested and require rigorous validation if they are to serve as reliable indicators of habitat loss and fragmentation, such as Montreal Process Indicator 1.1e. We apply a landscape ecology theory, supported by exploratory and confirmatory statistical techniques, to empirically test landscape metrics for reporting Montreal Process Indicator 1.1e in continuous dry eucalypt forests of sub-tropical Queensland, Australia. Target biota examined included: the Yellow-bellied Glider (Petaurus australis); the diversity of nectar and sap feeding glider species including P. australis, the Sugar Glider P. breviceps, the Squirrel Glider P. norfolcensis, and the Feathertail Glider Acrobates pygmaeus; six diurnal forest birds species; total diurnal bird species diversity; and the density of nectar-feeding diurnal bird species. Two scales of influence were considered: the stand-scale (2 ha), and a series of radial landscape extents (500 m - 2 km; 78 - 1250 ha) surrounding each fauna transect. For all biota, stand-scale structural and compositional attributes were found to be more influential than landscape metrics. For the Yellow-bellied Glider, the proportion of trace habitats with a residual element of old spotted-gum/ironbark eucalypt trees was a significant landscape metric at the 2 km landscape extent. This is a measure of habitat loss rather than habitat fragmentation. For the diversity of nectar and sap feeding glider species, the proportion of trace habitats with a high coefficient of variation in patch size at the 750 m extent was a significant landscape metric. None of the landscape metrics tested was important for diurnal forest birds. We conclude that no single landscape metric adequately captures the response of the region's forest biota per se. This poses a major challenge to regional reporting of Montreal Process Indicator 1.1e, fragmentation of forest types.

Effects of habitat characteristics and climate on the distribution and colouration of Dusky Moorhens (Gallinula tenebrosa) in south-east Queensland

We determined which factors predict the presence and abundance of Dusky Moorhens (Gallinula tenebrosa) at wetlands by surveying the ecological and habitat characteristics of 62 sites across south-east Queensland. Moorhens were observed in 48 of the sites sampled. They were more likely to be found at sites surrounded by taller terrestrial vegetation and where free-floating and attached aquatic vegetation was more abundant. The number of moorhens found at a site increased in relation to vegetation height, the abundance of attached aquatic vegetation and the number of purple swamphens observed. These results suggest that there are ecological constraints on the distribution of moorhens, and that food abundance and the availability of suitable nesting sites determine the overall distribution and abundance of moorhens in wetlands. Adult moorhens develop brightly coloured fleshy frontal shields, bills and legs when breeding, although in some populations birds maintain year-round colouration. We observed year-round breeding colouration in 23 out of 34 sampling sites that had moorhens and were surveyed in August. Coloured moorhens were found during winter at sites with higher minimum winter temperatures, and more abundant free-floating and submerged leafy vegetation. In addition, higher proportions of moorhens were coloured at sites with higher mean minimum temperatures. The retention of year-round breeding colouration appears to be restricted to areas with warmer winter temperatures and more abundant food. The results suggest that areas not occupied by moorhens are of inadequate quality to support breeding populations. We suggest that ecological constraints on independent breeding in Dusky Moorhens may have favoured the evolution of their unusual cooperative breeding system, which involves frequent mate-sharing by both sexes.

Habitat choice and vigilance behaviour of brush-tailed rock-wallabies (Petrogale penicillata) within their nocturnal foraging ranges

Habitat choice by brush-tailed rock-wallabies (Petrogale penicillata) in south-east Queensland was investigated by comparing the attributes of the nocturnal foraging locations that they selected with those of random locations within a radius of 50 m. Brush-tailed rock-wallabies were shown to select foraging locations on the basis of forage quality and/or their ability to see predators, rather than protection from predators amongst vegetation that could conceal them. Habitat choice may have been affected by limited food availability, as this study was conducted in the winter dry season. The attributes of foraging locations that brush- tailed rock-wallabies perceived as increasing their predation risk were assessed by recording the proportion of time that brush- tailed rock-wallabies spent vigilant while foraging. To measure vigilance, focal animals were observed with a night- vision scope for two minutes and the proportions of time spent vigilant and feeding were recorded. No measured feature of foraging locations was related to higher vigilance levels, suggesting that brush- tailed rock-wallabies did not alter their vigilance whether sheltered amongst grass tussocks or in open habitat, or whether feeding on good quality or poorer quality vegetation. Vigilance levels significantly declined as overnight temperatures decreased, which may have resulted from higher energy requirements of brush- tailed rock-wallabies during winter. The only factors that were found to significantly increase vigilance levels were high winds and moonlit nights. On bright nights, brush- tailed rock-wallabies were very unsettled and during high winds they often did not emerge to feed. More information is needed about how macropods detect predators at night before the effects of wind and light intensity upon vigilance can be fully understood.

Habitat-predator association and avoidance in rainbowfish (Melanotaenia spp.)

The ability to recall the location of a predator and later avoid it was tested in nine populations of rainbowfish (Melanotaenia spp.), representing three species from a variety of environments. Following the introduction of a model predator into a particular microhabitat, the model was removed, the arena rotated and the distribution of the fish recorded again. In this manner it could be determined what cues the fish relied on in order to recall the previous location of the predator model. Fish from all populations but one (Dirran Creek) were capable of avoiding the predator by remembering either the location and/or the microhabitat in which the predator was recently observed. Reliance on different types of visual cues appears to vary between populations but the reason for this variation remains elusive. Of the ecological variables tested (flow variability, predator density and habitat complexity), only the level of predation appeared to be correlated with the orientation technique employed by each population. There was no effect of species identity, which suggests that the habitat that each population occupies plays a strong role in the development of both predator avoidance responses and the cues used to track predators in the wild.

Occurrence of immature Culicoides molestus (Skuse) (Diptera : Ceratopogonidae) in relation to habitat characteristics

Analysis of beach sites on the Gold Coast, Australia, found that 14 physical and chemical habitat characteristics differed significantly between sites where numerous immatures of the canal biting midge, Culicoides molestus (Skuse), were found and sites where no midge immatures occurred. Five of the chemical factors found to reliably distinguish C. molestus habitat are major components of seawater, while another, electrical conductivity, is related to the concentration of seawater components. Calcium was the only one of the six primary components of seawater that was not a statistically significant correlate of C. molestus habitation by sand analysis. It is likely that a causative variable in occurrence of immatures is the concentration of seawater present in canals, because larvae are found where seawater component concentration is low in relation to uninhabited sites of similar appearance.

Low-density koala (Phascolarctos cinereus) populations in the mulgalands of south-west Queensland. III. Broad-scale patterns of habitat use

To date there have been few quantitative studies of the distribution of, and relative habitat utilisation by, koalas in the mulgalands of Queensland. To examine these parameters we applied habitat-accessibility and relative habitat-utilisation indices to estimates of faecal pellet density sampled at 149 sites across the region. Modelling the presence of pellets using logistic regression showed that the potential range of accessible habitats and relative habitat use varied greatly across the region, with rainfall being probably the most important determinant of distribution. Within that distribution, landform and rainfall were both important factors affecting habitat preference. Modelling revealed vastly different probabilities of finding a pellet under trees depending on the tree species, canopy size, and location within the region.

Genetic effects of chronic habitat fragmentation on tree species: the case of Sorbus aucuparia in a deforested Scottish landscape

Sustainable forest restoration and management practices require a thorough understanding of the influence that habitat fragmentation has on the processes shaping genetic variation and its distribution in tree populations. We quantified genetic variation at isozyme markers and chloroplast DNA (cpDNA), analysed by polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) in severely fragmented populations of Sorbus aucuparia (Rosaceae) in a single catchment (Moffat) in southern Scotland. Remnants maintain surprisingly high levels of gene diversity (H-E) for isozymes (H-E = 0.195) and cpDNA markers (H-E = 0.490). Estimates are very similar to those from non-fragmented populations in continental Europe, even though the latter were sampled over a much larger spatial scale. Overall, no genetic bottleneck or departures from random mating were detected in the Moffat fragments. However, genetic differentiation among remnants was detected for both types of marker (isozymes Theta(n) = 0.043, cpDNA Theta(c) = 0.131; G-test, P-value < 0.001). In this self-incompatible, insect-pollinated, bird-dispersed tree species, the estimated ratio of pollen flow to seed flow between fragments is close to 1 (r = 1.36). Reduced pollen-mediated gene flow is a likely consequence of habitat fragmentation, but effective seed dispersal by birds is probably helping to maintain high levels of genetic diversity within remnants and reduce genetic differentiation between them.

Habitat selection and population regulation in temporally fluctuating environments

Understanding and predicting the distribution of organisms in heterogeneous environments lies at the heart of ecology, and the theory of density-dependent habitat selection (DDHS) provides ecologists with an inferential framework linking evolution and population dynamics. Current theory does not allow for temporal variation in habitat quality, a serious limitation when confronted with real ecological systems. We develop both a stochastic equivalent of the ideal free distribution to study how spatial patterns of habitat use depend on the magnitude and spatial correlation of environmental stochasticity and also a stochastic habitat selection rule. The emerging patterns are confronted with deterministic predictions based on isodar analysis, an established empirical approach to the analysis of habitat selection patterns. Our simulations highlight some consistent patterns of habitat use, indicating that it is possible to make inferences about the habitat selection process based on observed patterns of habitat use. However, isodar analysis gives results that are contingent on the magnitude and spatial correlation of environmental stochasticity. Hence, DDHS is better revealed by a measure of habitat selectivity than by empirical isodars. The detection of DDHS is but a small component of isodar theory, which remains an important conceptual framework for linking evolutionary strategies in behavior and population dynamics.

Limited effect of anthropogenic habitat fragmentation on molecular diversity in a rain forest skink, Gnypetoscincus queenslandiae

To examine the effects of recent habitat fragmentation, we assayed genetic diversity in a rain forest endemic lizard, the prickly forest skink (Gnypetoscincus queenslandiae), from seven forest fragments and five sites in continuous forest on the Atherton tableland of northeastern Queensland, Australia. The rain forest in this region was fragmented by logging and clearing for dairy farms in the early 1900s and most forest fragments studied have been isolated for 50-80 years or nine to 12 skink generations. We genotyped 411 individuals at nine microsatellite DNA loci and found fewer alleles per locus in prickly forest skinks from small rain forest fragments and a lower ratio of allele number to allele size range in forest fragments than in continuous forest, indicative of a decrease in effective population size. In contrast, and as expected for populations with small neighbourhood sizes, neither heterozygosity nor variance in allele size differed between fragments and sites in continuous forests. Considering measures of among population differentiation, there was no increase in F-ST among fragments and a significant isolation by distance pattern was identified across all 12 sites. However, the relationship between genetic (F-ST) and geographical distance was significantly stronger for continuous forest sites than for fragments, consistent with disruption of gene flow among the latter. The observed changes in genetic diversity within and among populations are small, but in the direction predicted by the theory of genetic erosion in recently fragmented populations. The results also illustrate the inherent difficulty in detecting genetic consequences of recent habitat fragmentation, even in genetically variable species, and especially when effective population size and dispersal rates are low.