Roger P. Pech

The Puzzles of Population Cycles and Outbreaks of Small Mammals Solved

Abstract Well-known examples of high-amplitude, large-scale fluctuations of small-mammal populations include vole cycles in the boreal zone of Eurasia, lemming cycles in the high-arctic tundra of Eurasia and North America, snowshoe hare cycles in the boreal zone of North America, and outbreaks of house mice in southeastern Australia. We synthesize the recent knowledge of three key aspects of these animals population cyles: (1) periodicity, amplitude, and spatiotemporal synchrony; (2) reproduction and survival; and (3) underlying mechanisms. Survival rather than reproductive rate appears to drive rates of population increase during these fluctuations. Food limitation may stop increases of cyclic vole, lemming, and hare populations, whereas the decline from peak numbers is caused by predation mortality. In house mice, without coevolved predators, outbreaks may be driven by rainfall, food supply, and disease.

Limits to predator regulation of rabbits in Australia : evidence from predator-removal experiments

SummaryPredator-prey studies in semi-arid eastern Australia demonstrated that populations of rabbits (Oryctolagus cuniculus) could be regulated by predators. The functional, numerical and total responses of foxes (Vulpes vulpes) to rabbits and the numerical response of feral cats (Felis catus) to rabbits, are described. Measurement of the rabbit component of foxes stomach contents indicates a Type III functional response. The size of the fox population in summer was dependent on the availability of rabbits over the immediately preceding rabbit breeding season but there appeared to be no density-dependent aggregation of young foxes in areas of surplus food. The total response of foxes, estimated using the short-term numerical response of dispersing foxes, was directly density-dependent for low rabbit densities and inversely density-dependent for high rabbit densities. Two states are possible with this form of total response: a state with low rabbit densities regulated by predators and a state with high rabbit densities which occurs when rabbits escape predator regulation. The boundary between regulation and non-regulation by predators was demonstrated by a predator-removal experiment. In the treated areas, predators were initially culled and rabbits increased to higher densities than in an untreated area where predators were always present. When predators were allowed back into the treated areas, rabbit populations continued to increase and did not decline to the density in the untreated area. This is the critical evidence for a two-state system. When predators were present, rabbits could be maintained at low densities which were in the density-dependent part of the total response curve for foxes. Exceptionally high rabbit recruitment, or artificially reduced predation, could result in rabbits escaping predator-regulation. Under these circumstances, rabbits could move into the inversely density-dependent region of the total response curve for foxes.

Mice, rats, and people: the bio‐economics of agricultural rodent pests

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Heterogeneous landscapes and the role of refuge on the population dynamics of a specialist predator and its prey

How, and where, a prey species survives predation by a specialist predator during low phases of population fluctuations or a cycle, and how the increase phase of prey population is initiated, are much-debated questions in population and theoretical ecology. The persistence of the prey species could be due mainly to habitats that act as refuges from predation and/or due to anti-predatory behaviour of individuals. We present models for the former conjecture in two (and three) habitat systems with a specialist predator and its favoured prey. The model is based on dispersal of prey between habitats with high reproductive output but high risk of predation, and less productive habitats with relatively low risk of predation. We illustrate the predictions of our model using parameters from one of the most intriguing vertebrate predator–prey systems, the multi-annual population cycles of boreal voles and their predators. We suggest that cyclic population dynamics could result from a sequence of extinction and re–colonization events. Field voles (Microtus agrestis), a key vole species in the system, can be hunted to extinction in their preferred meadow habitat, but persist in sub-optimal wet habitats where their main predator, the least weasel (Mustela nivalis nivalis) has a low hunting efficiency. Re–colonization of favourable habitats would occur after the predator population crashes. At the local scale, the model suggests that the periodicity and amplitude of population cycles can be strongly influenced by the relative availability of risky and safe habitats for the prey. Furthermore, factors like intra-guild predation may lead to reduced predation pressure on field voles in sub-optimal habitats, which would act as a refuge for voles during the low phase of their population cycles. Elasticity analysis suggested that our model is quite robust to changes in most parameters but sensitive to changes in the population dynamics of field voles in the optimal grassland habitat, and to the maximum predation rate of weasels.

The influence of predation risk on foraging behaviour of brushtail possums in Australian woodlands

The potential effects of predation risk on common brushtail possums were investigated in south-eastern Australian woodlands. Patterns of habitat use, foraging costs using giving-up density (GUD) experiments, and indices of body condition and reproductive success were examined at eight sites in two habitat types (eucalypt- or cypress-pine–dominated stands), within three areas of different red fox abundance (high, moderate and low fox density). In cypress-pine–dominated stands, possums travelled further on the ground, visited more feeding stations and had lower GUDs at feeders where foxes were removed than did possums in high-fox-density sites. In contrast, there was no effect of fox removal on the behaviour of possums in eucalypt-dominated stands. Fox removal also had no effect on indices of body condition and reproduction. Minor effects of microhabitat were detected with trackplot and GUD experiments, but, overall, the results suggest that habitat at the stand-level was more important. The non-lethal effects of foxes in different habitats may need to be taken into account when developing conservation strategies for native marsupials.

Can outbreaks of house mice in south-eastern Australia be predicted by weather models?

Outbreaks of house mice (Mus domesticus) occur irregularly in the wheat-growing areas of south-eastern Australia, and are thought to be driven by weather variability, particularly rainfall. If rainfall drives grass and seed production, and vegetation production drives mouse dynamics, we should achieve better predictability of mouse outbreaks by the use of plant-production data. On a broader scale, if climatic variability is affected by El Nino-Southern Oscillation (ENSO) events, large-scale weather variables might be associated with mouse outbreaks. We could not find any association of mouse outbreaks over the last century with any ENSO measurements or other large-scale weather variables, indicating that the causal change linking mouse numbers with weather variation is more complex than is commonly assumed. For the 1960-2002 period we were only partly successful in using variation in cereal production to predict outbreaks of mice in nine areas of Victoria and South Australia, and we got better predictability of outbreaks from rainfall data alone. We achieved 70% correct predictions for a qualitative model using rainfall and 58% for a quantitative model using rainfall and spring mouse numbers. Without the detailed specific mechanisms underlying mouse population dynamics, we may not be able to improve on these simple models that link rainfall to mouse outbreaks.

Disease surveillance in wildlife with emphasis on detecting foot and mouth disease in feral pigs

Surveillance strategies for detecting disease in wildlife are reviewed. Opportunistic and structured surveillance schemes are examined, and the detection of foot and mouth disease (FMD) in feral pigs ( Sus scrofa ) in Australia discussed in more detail. The probability of detecting an individual case of FMD in feral pigs in Australia with the current opportunistic surveillance scheme is estimated to be less than 0·0015. A model derived from the binomial distribution and for sampling without replacement estimated that between 28 and 3077 FMD cases would occur before the disease outbreak is detected. The range depends on the level of certainty required for detection. A model of the dynamics of the disease and an equation for estimating sample sizes for sampling without replacement were combined. For a feral pig population occupying 100 km 2 at a population density of 15/km 2 , the estimated time to detection of an FMD outbreak, with opportunistic surveillance, was 23 to 358 days. Alternatively, with a fixed sample size of 200 feral pigs, the minimum time to detection was 29 days, and for 100 pigs, was 35 days. These results are for a 95% confidence level in detecting an outbreak. Methods for ranking areas according to the probability of disease occurrence or detection are discussed and presented as components of possible structured surveillance schemes.

Do Exotic Vertebrates Structure the Biota of Australia? An Experimental Test in New South Wales

From 1993 to 2001, we conducted a series of experiments in a mixed grassland–woodland system in central New South Wales (NSW) to quantify the interactions between red foxes and their prey and competitors. Foxes were removed from two areas around the perimeter of Lake Burrendong, and data were collected from these areas and a nearby untreated area before, during, and after the period of fox control. The arrival of rabbit hemorrhagic disease (RHD) in 1996 provided an opportunity to examine the interactive effects of controlling foxes and rabbits. In this landscape, typical of central NSW, (a) the fox population was not affected by a large reduction in the abundance of rabbits, or vice versa; (b) the cat population declined in areas where foxes were removed after the large RHD-induced reduction in rabbit numbers, but there was no consistent response to the removal of foxes; (c) the abundance of some macropod species increased in response only to the combined removal of rabbits and foxes; (d) there were no consistent changes in the abundances of bird species in response to the removal of either foxes or rabbits, but there were clear habitat differences in bird species richness; and (e) there was likely to be an increase in woody plant species after the large reduction in rabbit populations by RHD. We conclude that (a) long-term field experiments (more than 3 years) are required to quantify the indirect consequences of controlling foxes and rabbits, and (b) single manipulations, such as fox control or rabbit control, are not necessarily sufficient for the conservation of remnant woodland communities in southeastern Australia.

Rats on the run: removal of alien terrestrial predators affects bush rat behaviour

Predators can strongly influence the microhabitat use and foraging behaviour of prey. In a large-scale replicated field experiment in East Gippsland, Australia, we tested the effects of reduced alien red fox (Vulpes vulpes) and alien wild dog (Canis lupus familiaris) abundance (treatment) on native bush rat (Rattus fuscipes) behaviour. Bush rats are exposed to two main guilds of predators, namely mammalian carnivores and birds of prey. Tracking rat movements using the spool-and-line technique revealed that, in treatment sites, rats used ground cover, which provides shelter from predators, less often than at unmanipulated fox and wild dog abundance (non-treatment sites). In treatment sites, rats more frequently moved on logs where they would have been exposed to hunting foxes and dogs than in non-treatment sites. Furthermore, in treatments, rats showed a preference for understorey but not in non-treatments. Hence, bush rats adapted their behaviour to removal of alien terrestrial predators. Giving-up densities (GUDs) indicated no treatment effects on the marginal feeding rate of bush rats. Interestingly, GUDs were higher in open patches than in sheltered patches, suggesting higher perceived predation risk of bush rats during foraging at low versus high cover. The lack of treatment effects on GUDs but the clear response of bush rats to cover may be explained by the impact of predators other than foxes and wild dogs.

Models for the introgression of a transgene into a wild population within a stochastic environment, with applications to pest control

Several forms of control are currently under consideration for pest species in Australia. Perhaps the most novel makes use of either an inducible fatality gene (IFG) or an inducible sterility gene (ISG). The transgene is integrated into the genome of target populations by periodically releasing transgenic animals. When high proportions of the population carry the transgene then the gene may be induced to reduce abundance. A fundamental feasibility issue is how much time and effort is required before the proportion of the population carrying the transgene reaches values close to 1. It is shown that the speed of introgression may be underestimated if models do not allow for year-to-year variability in natural processes, such as the number of young arising from natural breeding or the number of adults surviving to breed. Furthermore, the improvements in speed of introgression due to variability are magnified for species with lower lifespans. We include, as specific examples, application of the model to common carp (Cyprinus carpio) and mosquitofish (Gambusia affinis), which are major freshwater pests in Australia.