Spatial and temporal drivers of small mammal distributions in a semi-arid environment : the role of rainfall, vegetation and life-history

A key task in ecology is to understand the drivers of animal distributions. In arid and semi-arid environments, this is challenging because animal populations show considerable spatial and temporal variation. An effective approach in such systems is to examine both broad-scale and long-term data. We used this approach to investigate the distribution of small mammal species in semi-arid ‘mallee’ vegetation in south-eastern Australia. First, we examined broad-scale data collected at 280 sites across the Murray Mallee region. We used generalized additive mixed models (GAMMs) to examine four hypotheses concerning factors that influence the distribution of individual mammal species at this scale: vegetation structure, floristic diversity, topography and recent rainfall. Second, we used long-term data from a single conservation reserve (surveyed from 1997 to 2012) to examine small mammal responses to rainfall over a period spanning a broad range of climatic conditions, including record high rainfall in 2011. Small mammal distributions were strongly associated with vegetation structure and rainfall patterns, but the relative importance of these drivers was species-specific. The distribution of the mallee ningaui Ningaui yvonneae, for example, was largely determined by the cover of hummock grass; whereas the occurrence of the western pygmy possum Cercartetus concinnus was most strongly associated with above-average rainfall. Further, the combination of both broad-scale and long-term data provided valuable insights. Bolam's mouse Pseudomys bolami was uncommon during the broad-scale survey, but long-term surveys showed that it responds positively to above-average rainfall. Conceptual models developed for small mammals in temperate and central arid Australia, respectively, were not, on their own, adequate to account for the distributional patterns of species in this semi-arid ecosystem. Species-specific variation in the relative importance of different drivers was more effectively explained by qualitative differences in life-history attributes among species.

Landscape characteristics associated with species richness and occurrence of small native mammals inhabiting a coastal heathland: a spatial modelling approach

In the coastal region of south-western Victoria, Australia, populations of native small mammal species are restricted to patches of suitable habitat in a highly fragmented landscape. The size and spatial arrangement of these patches is likely to influence both the occupancy and richness of species at a location. Geographic Information System (GIS)-based habitat models of the species richness of native small mammals, and individual species  occurrences, were developed to produce maps displaying the spatial  configuration of suitable habitat. Models were generated using either generalised linear Poisson regression (for species richness) or logistic regression (for species occurrences) with species richness or  presence/absence as the dependent variable and landscape variables, extracted from both GIS data layers and multi-spectral digital imagery, as the predictor variables. A multi-model inference approach based on the Akaike Information Criterion was used and the resulting model was applied in a GIS framework to extrapolate predicted richness/likelihood of occurrence across the entire area of the study. A negative association between species  richness and elevation, habitat complexity and sun index indicated that richness within the study area decreases with increasing altitude, vertical vegetation structure and exposure to solar radiation. Landform  characteristics were important (to varying degrees) in determining habitat occupancy for all of the species examined, while the influence of habitat complexity was important for only one of the species. Performance of all but one of the models generated using presence/absence data was high, as indicated by the area under the curve of a receiver-operating characteristic plot. The effective conservation of the small mammal species in the area of concern is likely to depend on management actions that promote the protection of the critical habitats identified in the models.

Plant pathogens causing vegetation dieback: a serious threat to the conservation of small mammals in Australia

The soil-borne plant pathogen Phytophthora cinnamomi occurs in most Australian states. It is pathogenic to many Australian species, particularly the Proteaceae, Fabaceae, Dillineaceae and Epacridaceae. In Western Australia, c. 2000 of the 9000 endemic plant species are directly affected by the disease. The epidemic of plant deaths caused by P. cinnamomi is recognised as one of 11 Key Threatening Processes to the Australian Environment, and is now also acknowledged as a potential threat fauna in a range of communities. The implications of landscape modification due to the effects of P. cinnamomi dieback prompted our research, designed to measure the distribution and abundance of small mammals in disease-affected ecosystems. This study was in the Jarrah (Eucalyptus marginata) forests in the Darling Range, Western Australia and measured the distribution and abundance of one small mammal species, the Mardo (Antechinus flavipes) by Elliott trapping in forests with (1) high, (2) mixed and (3) no evidence of Phytophthora dieback. Trap success was highest in sites with no effect of Phytophthora (7.3 animals per 100 trap nights), whereas the lowest trap success was recorded at the high impact sites (0.67 animals per 100 trap night). There was a significant difference in trap success of Mardos in Elliott trapping over 1800 trap nights (x²= 23.19, d.f = 5, p < 0.001). An examination of the distribution of individuals and sexes suggests that Phytophthora-affected sites act as sinks for Mardos, while source areas are healthy, unaffected Jarrah forest.

Recolonisation of powerline corridor vegetation by small mammals : timing and the influence of vegetation management

Powerline corridors through forested ecosystems have been criticised due their potential to fragment the landscape and facilitate the intrusion of undesirable species into natural areas. This study investigates the effects of vegetation management (slashing), on: (1) timing of small mammal recolonisation; (2) vegetation characteristics that drive small mammal responses; and (3) the point where corridor resources are sufficient to provide functional habitat for native species. Small mammal trapping was undertaken within Bunyip State Park, Australia, across three sites, once a month from January 2001 to May 2002 and every 2 months thereafter until January 2004. Changes in vegetation around each trap station were assessed annually in the forest and bi-annually in the corridor. Principal components analysis on the vegetation structural complexity values produced factors for use in species abundance models. Native small mammal species recolonised the corridor 1.5–3.5 years after management and the corridor supported a breeding population of small mammals around 2.5 years post-management. Males however, generally recolonised the corridor first, resulting in a sex-biased population in these areas. Species corridor habitat models for five native and one introduced species suggested cover and shelter were more important in determining corridor use than plant species per se. Powerline corridors have the potential to create a mixture of different successional stages, enhancing habitat availability for many species. However, the intensity of current management needs to be reduced and an integrated approach to management needs to be undertaken if powerline corridors are to continuously provide habitat for native small mammal species.

The short-term responses of small mammals to wildfire in semiarid mallee shrubland, Australia

Context. Wildfire is a major driver of the structure and function of mallee eucalypt- and spinifex-dominated landscapes. Understanding how fire influences the distribution of biota in these fire-prone environments is essential for effective ecological and conservation-based management.

Aims. We aimed to (1) determine the effects of an extensive wildfire (118 000 ha) on a small mammal community in the mallee shrublands of semiarid Australia and (2) assess the hypothesis that the fire-response patterns of small mammals can be predicted by their life-history characteristics.

Methods. Small-mammal surveys were undertaken concurrently at 26 sites: once before the fire and on four occasions following the fire (including 14 sites that remained unburnt). We documented changes in small-mammal occurrence before and after the fire, and compared burnt and unburnt sites. In addition, key components of vegetation structure were assessed at each site.

Key results. Wildfire had a strong influence on vegetation structure and on the occurrence of small mammals. The mallee ningaui, Ningaui yvonneae, a dasyurid marsupial, showed a marked decline in the immediate post-fire environment, corresponding with a reduction in hummock-grass cover in recently burnt vegetation. Species richness of native small mammals was positively associated with unburnt vegetation, although some species showed no clear response to wildfire.

Conclusions. Our results are consistent with the contention that mammal responses to fire are associated with their known life-history traits. The species most strongly affected by wildfire, N. yvonneae, has the most specific habitat requirements and restricted life history of the small mammals in the study area. The only species positively associated with recently burnt vegetation, the introduced house mouse, Mus domesticus, has a flexible life history and non-specialised resource requirements.

Implications. Maintaining sources for recolonisation after large-scale wildfires will be vital to the conservation of native small mammals in mallee ecosystems.

Powerline corridors: degraded ecosystems or wildlife havens?

Management of powerline corridors in Australia has traditionally focused on the complete removal of vegetation using short rotation times owing to the perceived hazard of fire associated with corridor vegetation. Because of the intense management associated with fire hazards, little thought has been given to use of powerline corridors by wildlife. This has resulted in corridors traditionally being viewed as a source of fragmentation and habitat loss within forested ecosystems. We investigated the responses of small mammal communities living in a powerline corridor to management-induced vegetation changes at different successional stages, to determine whether a compromise could be reached between managing corridors for fire and biodiversity. Habitat modelling in the corridor and adjacent forest for three native and one introduced small mammal species demonstrated that species responded to changes in vegetation structural complexity, rather than time-since-management per se. Early seral stages of vegetation recovery after corridor management encouraged the introduced house mouse (Mus domesticus) into corridors and contributed little to biodiversity. Mid-seral-stage vegetation, however, provided habitat for native species that were rare in adjacent forest habitats. As the structural complexity of the vegetation increased, the small mammal community became similar to that of the forest so that corridor vegetation contributed fewer biodiversity benefits while posing an unacceptable fire risk. If ecologically sensitive management regimes are implemented to encourage mid-seral vegetation and avoid complete vegetation removal, powerline corridors have the potential to improve biodiversity. This would maintain landscape connectivity and provide habitat for native species uncommon in the forest while still limiting fuel loads in the corridor.

Changes in vegetation sand floristics under a powerline easement and implications for vegetation management

Utility corridors such as powerlines are widespread linear easements of highly modified vegetation which often fragment natural areas of conservation significance. Vegetation management along these easements is aimed at modifying vegetation structure by the removal of all tall shrubs and trees, which may have adverse impacts on flora and fauna diversity. Victoria's Bunyip State Park is bisected by a high voltage powerline easement which is managed by a four year slashing cycle. Repeated slashing has altered plant species composition and structure of the drier slope and ridge vegetation compared to unslashed adjacent Open Forest vegetation, but Wet Heath within the management zone has remained largely unmodified. At a broad level, plant species diversity in the easement is increased, and higher vegetation density has created small mammal habitat. The powerline easement did not appear to facilitate weed invasion. Vegetation management by repeated slashing has altered the vegetation, but does not appear to have had adverse conservation impacts on local plant and small mammal diversity.

Effect of Phytophthora cinnamomi on the habitat utilization of Antechinus stuartii in a Victorian forest

Phytophthora cinnamomi (Cinnamon fungus) is a pathogenic soil fungus which infects plant communities along the south-eastern coast of Australia, and the south-western corner of Western Australia. The symptoms of this disease include chlorosis, death of branches (ie. ‘dieback’), retarded growth and the eventual death of infected plants. This leads to devastating effects upon plant communities by altering both the structural and floristic characteristics of these communities. Small mammal species are dependent on specific features of their habitat such as vegetation structure and floristics. This thesis investigated alterations to the habitat of the insectivorous marsupial mouse, Antechinus stuartii, due to the presence of P. cinnamomi. The study was undertaken in an area of an open forest in the Brisbane Ranges, Victoria. Significant changes were found in both the floristic composition and structure of the vegetation at study sites infected with P, cinnamomi, compared to uninfected sites. The habitat utilization by A. stuartii of uninfected and infected vegetation was investigated using live trapping and radio-telemetric techniques. Capture rates were higher at sites uninfected by P. cinnamomi, and both male and females selected areas free from infection. Home range areas of males were significantly larger than those of females as assessed by telemetry. Both sexes spent a high proportion of time in areas dominated by Xanthorrhoea australis (Austral grass tree). There were significant relationships between the abundance of A. stuartii and the denseness of vegetation above 1 metre in height, and in particular, the proportion of cover afforded by X. australis. There were no significant differences in the cover of Eucalyptus spp. between uninfected and infected sites, but there were significantly more nest hollows in infected areas. The abundance of invertebrates was examined using pitfall traps. There were no significant differences in the abundance of the larger invertebrate taxa at infected and uninfected sites, but higher abundances of some micro-invertebrate groups in infected areas were recorded. The most likely factors considered to be influential in the habitat selection of A. stuartii were vegetation structure, and the presence of X. australis. To assess whether these factors were important the leaves of X. australis were removed with a brushcutter, to mimic the early effects of infection with P. cinnamomi. Animals did not respond to the alteration of vegetation structure in the short term (3-4 days). Longer-term experiments are required to assess the habitat utilization of A. stuartii at different periods following habitat manipulation. The implications of the presence of P. cinnamomi on the conservation of fauna are discussed. The destructive nature of the pathogen, and the slow rate of recovery from the disease, means that P. cinnamomi can be considered a threatening process to plant communities and the fauna that reside within that habitat. Future management of this disease within natural areas must therefore be cognisant of the potential of P. cinnamomi to significantly affect faunal as well as vegetative communities.

Effects of phytophthora cinnamomi on heathland flora and fauna of the Eastern Otway Ranges

The plant pathogen, Phytophthora dnnamomi, is a cause of dieback disease observed in sclerophyll vegetation in Australia, The effects of P. dnnamomi on flora and fauna were studied at two locations in heathland vegetation near the coastal town of Anglesea, Victoria. The pathogen was isolated from soils beneath diseased heathland plants. The extent of diseased vegetation was assessed by the presence and absence of highly sensitive indicator species, Xanthorrhoea australis and hopogon ceratophyllus. The characteristics of heathland vegetation exhibiting dieback disease associated with the presence of P. dnnamomi were investigated. Plant species richness was similar between diseased and non-diseased areas however diseased areas were characterised by significant declines in the cover and frequency of susceptible species, increases in resistant species and increases in percent cover of open ground. Compared to non-diseased areas, diseased areas exhibited fewer shrub species and decreased shrub cover. The percentage cover and number of species of sedges, lilies and grasses were higher in diseased areas. Structural differences were significant between 0-0.6 m with decreased cover of vegetation in diseased areas. Differences in structure between diseased and non-diseased areas were not as great as expected due to increases in the cover of resistant species. A number of regenerating X australis were observed in post-disease areas. Cluster analysis of floristic data could clearly separate diseased and non-diseased trap stations. The population dynamics and habitat use of eight small mammal species present were compared in diseased and non-diseased areas using trapping and radio-tracking techniques. The number of small mammal species captured in post-disease areas was significantly lower than non-diseased areas. Mean captures of Antechinus stuartii and Rattus fiisdpes were significantly lower in diseased areas on Grid B. Mean captures of Rattus lutreolus were significantly lower in diseased areas on both study grids. Significant differences were not observed in every season over the two year study period. Radio tracking revealed more observations of Sminthopsis leucopus in non-diseased vegetation than in diseased. Cercartetus nanus was frequently observed to utilise the disease susceptible X. australis for nesting. At one location, the recovery of vegetation and small mammal communities in non-diseased and diseased vegetation after fuel reduction burning was monitored for three years post-fire. Return of plant species after fire in both disease classes were similar, reaching 75% of pre-fire richness after three years. Vegetation cover was slower to return after fire in diseased areas. Of the seven small mammal species captured pre-fire, five were regularly captured in the three years after fire. General linear model analysis revealed a significant influence of disease on capture rates for total small mammals before fire and a significant influence of fire on capture rates for total small mammals after fire. After three years, the influence of fire on capture rates was reduced no significant difference was detected between disease classes. Measurements of microclimate indicate that diseased, burnt heathland was likely to experience greater extremes of temperature and wind speed. Seeding of diseased heathland with X. australis resulted in the establishment of seedlings of this sensitive species. The reported distributions of the mamma] species in Victoria were analysed to determine which species were associated with the reported distribution of dieback disease. Twenty-two species have more than 20% of their known distribution in diseased areas. Five of these species, Pseudomys novaehollandiae, Pseudomys fumeust Pseudomys shortridgei, Potorous longipes and Petrogale pencillata are rare or endangered in Victoria. Four of the twenty-two species, Sminthopsis leucopus, Isoodon obesulus, Cercartetus nanus and Rottus lutreolus am observed in Victorian heathlands. Phytophthora cinnamomi changes both the structure and floristics of heathland vegetation in the eastern Qtway Ranges. Small mammals respond to these changes through decreased utilisation of diseased heathland. The pathogen threatens the diversity of species present and future research efforts should be directed towards limiting its spread and rehabilitating diseased areas.

Influence of disturbance on threatened fauna and associated habitat, Victoria

Results of fauna and vegetation surveys conducted around Portland Aluminium smelter between 1979 and 2004 found small mammal abundance and diversity had declined and changes in vegetation communities were related to changes in fire patterns, vegetation fragmentation and weed invasion. Small mammal numbers were greater in nearby National Parks.

Reversed sexual dimorphism and altered prey base : the effect on sooty owl (Tyto tenebricosa tenebricosa) diet

The ecology and function of many Australian predators has likely been disrupted following major changes in prey base due to declines in distribution and abundance of small mammals following European settlement. This study investigated various aspects of the dietary ecology of sooty owls (Tyto tenebricosa tenebricosa), including sexual variation as they potentially exhibit the greatest degree of reversed sexual dimorphism of any owl species worldwide. Sooty owls are highly opportunistic predators of non-volant small mammals, consuming most species known to exist in the region, so their diet fluctuates seasonally and spatially due to varying prey availability, and is particularly influenced by the breeding cycles of prey. Significant intersexual dietary differences existed with female sooty owls predominantly consuming much larger prey items than males, with dietary overlap at 0.62. The current reliance on relatively few native mammalian species is of conservation concern, especially when mammal declines are unlikely to have ceased as many threatening processes still persist in the landscape. Sooty owl conservation appears inextricably linked with small mammal conservation. Conservation efforts should be focussed towards improving prey densities and prey habitat, primarily by implementing control programs for feral predators and preventing the loss of hollow-bearing trees throughout the landscape

Why do some species have geographically varying responses to fire history?

A capacity to predict the effects of fire on biota is critical for conservation in fire-prone regions as it assists managers to anticipate the outcomes of different approaches to fire management. The task is complicated because species' responses to fire can vary geographically. This poses challenges, both for conceptual understanding of post-fire succession and fire management. We examine two hypotheses for why species may display geographically varying responses to fire. 1) Species' post-fire responses are driven by vegetation structure, but vegetation - fire relationships vary spatially (the 'dynamic vegetation' hypothesis). 2) Regional variation in ecological conditions leads species to select different post-fire ages as habitat (the 'dynamic habitat' hypothesis). Our case study uses data on lizards at 280 sites in a ~ 100 000 km2 region of south-eastern Australia. We compared the predictive capacity of models based on 1) habitat associations, with models based on 2) fire history and vegetation type, and 3) fire history alone, for four species of lizards. Habitat association models generally out-performed fire history models in terms of predictive capacity. For two species, habitat association models provided good discrimination capacity even though the species showed geographically varying post-fire responses. Our results support the dynamic vegetation hypothesis, that spatial variation in relationships between fire and vegetation structure results in regional variation in fauna-fire relationships. These observations explain how the widely recognised 'habitat accommodation' model of animal succession can be conceptually accurate yet predictively weak. © 2014 The Authors.

Fire affects microhabitat selection, movement patterns, and body condition of an Australian rodent (Rattus fuscipes)

Resource selection by animals influences individual fitness, the abundance of local populations, and the distribution of species. Further, the degree to which individuals select particular resources can be altered by numerous factors including competition, predation, and both natural- and human-induced environmental change. Understanding the influence of such factors on the way animals use resources can guide species conservation and management in changing environments. In this study, we investigated the effects of a prescribed fire on small-scale (microhabitat) resource selection, abundance, body condition, and movement pathways of a native Australian rodent, the bush rat (Rattus fuscipes). Using a before-after, control-impact design, we gathered data from 60 individuals fitted with spool and line tracking devices. In unburnt forest, selection of resources by bush rats was positively related to rushes, logs and complex habitat, and negatively related to ferns and litter. Fire caused selection for spreading grass, rushes, and complex habitat to increase relative to an unburnt control location. At the burnt location after the fire, rats selected patches of unburnt vegetation, and no rats were caught at a trapping site where most of the understory had been burnt. The fire also reduced bush rat abundance and body condition and caused movement pathways to become more convoluted. After the fire, some individuals moved through burnt areas but the majority of movements occurred within unburnt patches. The effects of fire on bush rat resource selection, movement, body condition, and abundance were likely driven by several linked factors including limited access to shelter and food due to the loss of understory vegetation and heightened levels of perceived predation risk. Our findings suggest the influence of prescribed fire on small mammals will depend on the resulting mosaic of burnt and unburnt patches and how well this corresponds to the resource requirements of particular species.