PREDATOR-PREY INTERACTIONS IN THE SPINIFEX GRASSLANDS OF CENTRAL AUSTRALIA A thesis submitted in partial fulfilment of the requirements for the award of the degree of DOCTOR OF PHILOSOPHY from UNIVERSITY OF WOLLONGONG by RACHEL M. PALTRIDGE, Bsc (hons.) SCHOOL OF BIOLOGICAL SCIENCES 2005 CERTIFICATION I, Rachel M. Paltridge, declare that this thesis, submitted in partial fulfilment of the requirements for the award of Doctor of Philosophy, in the Department of Biological Sciences, University of Wollongong, is wholly my own work unless otherwise referenced or acknowledged. The document has not been submitted for qualifications at any other academic institution. Rachel M. Paltridge 10 May 2005. i ABSTRACT Predation by exotic predators (cats Felis catus and foxes Vulpes vulpes) is believed to be one of the factors that has contributed to the decline of medium-sized mammals in arid Australia. Other factors include habitat degradation by introduced herbivores (rabbits Oryctolagus cuniculus and grazing stock) and altered fire regimes after Aboriginal people moved into permanent settlements. In general, the impact of exotic predators on arid zone mammals is believed to be significant only when predator numbers have been elevated by increased food availability from exotic prey species (rabbits, house-mice Mus domesticus, cattle carcasses) or when native prey populations have already been dramatically reduced by competition from introduced herbivores. In much of the spinifex grasslands of the central Australian deserts, pastoralism never occurred, rabbit colonisation was extremely patchy and in some areas, traditional burning was still being practised when the extinctions commenced. None of the current models of mammalian extinctions adequately explain the declines in this environment. In this study I examined predator-prey interactions in two areas of the Tanami Desert to investigate whether predation by exotic predators may be a primary agent of extinction in its own right, capable of causing mass declines even in the absence of other human- induced perturbations. If this were the case then the following would be expected: (i) cats and foxes would eat medium-sized mammals when they are available, but be able to survive on alternative prey when mammals are scarce; (ii) populations of cats and foxes would be buffered against the declines of mammals during droughts, or would be able to recover more quickly than medium-sized mammals after droughts; (iii) medium- sized mammals would be more vulnerable to predation by cats and foxes than by dingoes Canis lupus dingo and other native predators, and (iv) there would be a correlation between the timings of the extinctions and the colonisation (or sudden increase) of cats and foxes. These predictions were investigated by monitoring the diets and relative abundance of cats, foxes and dingoes in relation to fluctuating prey availability in two areas of the Tanami Desert at latitudes separated by approximately 400 km. Mean annual rainfall is higher and more reliable in the northern study area which was situated in the centre of bilby Macrotis lagotis distribution within the Northern Territory, whereas the southern study area was located on the southern edge of the bilby’s range. Within each study ii area, monitoring occurred at three sites, approximately 20 km apart. Each site contained a sub-plot in each of two habitat types. Field work was conducted between September 1995 and December 1997. When the study began, the southern study area was experiencing drought conditions, however both study areas received significant rainfall in early 1997. The population dynamics of a variety of potential prey groups were monitored to examine their resilience during droughts, patterns of recolonisation after rainfall, and use of two habitat types: the ubiquitous sandplain, and the moister, nutrient enriched palaeodrainage habitat which is believed to have provided a refuge for medium-sized mammals during droughts. Native mammals were uncommon throughout the study period. Bilbies and macropods were significantly more abundant in the northern study area, and tended to occur more frequently in palaeodrainage habitat than sandplain. However, the palaeodrainage habitat did not appear to provide adequate refuge for the medium and large mammals during drought conditions in the southern study area, as they disappeared from the study sites altogether. Small mammals were significantly more abundant in the southern study area but densities remained low (< 2% trap success) throughout the study, and showed little response to improved seasonal conditions. In contrast, the abundance and species richness of birds showed a marked increase following rainfall in the southern study area. Flocks of nomadic birds arrived within several months of drought-breaking rains, increasing the relative abundance of birds from 9.3 per km of transect in December 1996 to 49/km in July 1997. Reptiles were the most resilient prey group during the drought conditions. Both varanids and smaller reptiles were equally abundant in the wet and dry years and showed no difference in abundance between study areas. However, reptiles showed marked temperature-related patterns in activity, with many species becoming inactive in the winter months. A total of 142 cat scats, 126 fox scats and 75 dingo scats were analysed to investigate predator diets in the two study areas. Unlike cat, fox and dingo diets elsewhere in Australia (and the world), mammalian prey did not dominate. Reptile was the prey category that was most frequently consumed by cats and foxes in ‘summer’ (October- iii April) and by dingoes throughout the year, and was identified as a “seasonal staple” prey type for all three predators in the Tanami Desert. When biomass of prey was taken into account, the varanids (predominantly the sand goanna Varanus gouldii) were the most important prey sustaining predators in the two study areas. Birds were an important part of the diets of cats and foxes in winter when reptiles were less active. Small mammals were consumed by cats and foxes throughout the study, in proportion to their field abundances. Invertebrates were a major component of the diets of foxes, representing 31% of prey items consumed. There was considerable overlap in the diets of the three predator species, but dingoes ate more medium (100-999 g) and large (>1000 g) prey than cats and foxes did. The scarcity of medium-sized mammals in the study areas provided little opportunity to find evidence of predation events on such prey. However, bilby remains were found in two cat scats and one dingo scat in the northern study area, mulgara Dasycercus cristicauda remains occurred in several cat and fox scats from the southern study area, and there were fourteen occurrences of marsupial mole Notoryctes typhlops in predator scats during the study, primarily in fox scats. Elsewhere in Australia, there is ample evidence that cats and foxes regularly consume medium-sized mammalian prey (e.g. rabbits and ringtail possums Pseudocheirus peregrinus) when it is available. Overall cats were the most abundant eutherian predators in the two study areas, and they were significantly more abundant in the northern study area than the southern study area. Surveys revealed that cats can persist into droughts by feeding on reptilian prey. When the study commenced, cats occurred on five of the six sub-plots in the southern study area, despite six consecutive years of below-average rainfall. However, by the end of the first year, they could only be detected on one sub-plot. Recolonisation of the sites rapidly occurred after significant rainfall (260 mm in 2 months), when nomadic birds colonised the sites and provided a plentiful food source. Foxes also declined to very low densities during drought in the southern study area, but they had recolonised all sites by the winter of 1997. This coincided with the increase in abundance of birds, which became their most frequently consumed prey item. Overall, foxes were equally abundant in the two study areas, but statistical analyses revealed a significant interaction between latitude and habitat because in the southern study area foxes tended to utilise the palaeodrainage habitat more than the sandplain, whereas in iv the northern study area the majority of fox sign was detected in the sandplain habitat. This may have been due to the abundance of dingoes in the palaeodrainage habitat in the northern study area. Dingoes were significantly more abundant in the northern study area than the southern, where they were usually only present at one of the three sites. The northern study area had higher densities of macropods (supplementary prey for dingoes) and more reliable access to drinking water, which persisted in the palaeodrainage channels for up to 6 months after significant rain events. Dingo numbers were relatively stable throughout the study and did not increase in response to improved seasonal conditions in the southern study area in 1997. This study revealed that the distribution of foxes extends further north into the Tanami Desert than has previously been reported, and is not necessarily tied to the distribution of rabbits in the Northern Territory. Furthermore, discussion with Aboriginal people who lived a traditional lifestyle in the area until the 1940s, revealed that foxes were already present in the northern Tanami desert at that time, before the disappearance of many medium-sized mammal species. The patterns of medium-sized mammalian extinctions in the northern and western deserts between 1940 and 1960 is thus consistent with the colonisation of the fox. Although cats had been present in central Australia for at least 50 years before the mammalian declines occurred, this does not discount them from contributing to the extinction process. It is postulated that during the early decades of their colonisation of the arid interior, cat populations may have been maintained at low levels by predation from dingoes and also Aboriginal people (for whom cats were a favoured food). But between 1920 and 1960 the western deserts were depopulated of Aboriginal people, and human hunting of cats diminished.