Niche partitioning in a guild of invasive mammalian predators.

Predators compete aggressively for resources, establishing trophic hierarchies that influence ecosystem structure. Competitive interactions are particularly important in invaded ecosystems where introduced predators can suppress native prey species. We investigated whether niche partitioning exists within a guild of invasive mammalian predators and determined the consequences for native species. Over 4405 camera-trap days, we assessed interactions among three invasive predators: two apex predators (feral cats Felis catus and ferrets Mustela furo) and a mesopredator (stoats Mustela erminea), in relation to their primary prey (lagomorphs, rodents and birds) and habitat use. Further, we tested for mesopredator release by selectively removing cats and ferrets in a pulse perturbation experiment. We found compelling evidence of niche partitioning; spatiotemporal activity of apex predators maximized access to abundant invasive prey, with ferrets targeting lagomorphs and cats targeting rodents. Mesopredators adjusted their behavior to reduce the risk of interference competition, thereby restricting access to abundant prey but increasing predation pressure on diurnal native birds. Stoats were only recorded at the treatment site after both larger predators were removed, becoming the most frequently detected predator at 6 months post-perturbation. We suggest there is spatial and resource partitioning within the invasive predator guild, but that this is incomplete, and avoidance is achieved by temporal partitioning within overlapping areas. Niche partitioning among invasive predators facilitates coexistence, but simultaneously intensifies predation pressure on vulnerable native species.

Invasive mammalian predators habituate to and generalize avian prey cues: a mechanism for conserving native prey.

Invasive mammalian predators can cause the decline and extinction of vulnerable native species. Many invasive mammalian predators are dietary generalists that hunt a variety of prey. These predators often rely upon olfaction when foraging, particularly at night. Little is understood about how prey odor cues are used to inform foraging decisions. Prey cues can vary spatially and temporally in their association with prey and can either reveal the location of prey or lead to unsuccessful foraging. Here we examine how two wild-caught invasive mammalian bird predator species (European hedgehogs Erinaceus europaeus and ferrets Mustela putorius furo) respond to unrewarded bird odors over successive exposures, first demonstrating that the odors are perceptually different using house mice (Mus musculus) as a biological olfactometer. We aim to test if introduced predators categorize odor cues of similar prey together, a tactic that could increase foraging efficiency. We exposed house mice to the odors using a standard habituation/dishabituation test in a laboratory setting, and wild-caught European hedgehogs and ferrets in an outdoor enclosure using a similar procedure. Mice discriminated among all bird odors presented, showing more interest in chicken odor than quail or gull odor. Both predator species showed a decline in interest toward unrewarded prey odor (i.e., habituation), but only ferrets generalized their response from one unrewarded bird odor to another bird odor. Hedgehog responses to unrewarded bird odors were highly variable between individuals. Taken together, our results reveal interspecific and intraspecific differences in response to prey odors, which we argue are a consequence of different diet breadth, life and evolutionary histories, and the conditions in each experiment. Generalization of prey odors may have enabled some species of invasive predators to efficiently hunt a range of intraguild prey species, for example, ground-nesting shorebirds. Olfactory manipulation of predators may be a useful conservation tool for threatened prey if it reduces the conspicuousness of vulnerable prey.

Multiple ecological processes underpin the eruptive dynamics of small mammals: House mice in a semi-arid agricultural environment.

Mouse plagues are a regular feature of grain-growing regions, particularly in southern and eastern Australia, yet it is not clear what role various ecological processes play in the eruptive dynamics generating these outbreaks.This research was designed to assess the impact of adding food, water, and cover in all combinations on breeding performance, abundance, and survival of mouse populations on a typical cereal growing farm in northwestern Victoria.Supplementary food, water, and cover were applied in a 2 × 2 × 2 factorial design to 240 m sections of internal fence lines between wheat or barley crops and stubble/pasture fields over an 11-month period to assess the impact on mouse populations.We confirmed that mice were eating the additional food and were accessing the water provided. We did not generate an outbreak of mice, but there were some significant effects from the experimental treatments. Additional food increased population size twofold and improved apparent survival. Both water and cover improved breeding performance. Food and cover increased apparent survival.Our findings confirm that access to food, water, and cover are necessary for outbreaks, but are not sufficient. There remain additional factors that are important in generating mouse plagues, particularly in a climatically variable agricultural environment.

Landowners' Perspectives on Coordinated, Landscape-Level Invasive Species Control: The Role of Social and Ecological Context.

To achieve biodiversity gains, landowner engagement in coordinated invasive species control programs across private lands is needed. Understanding landowners' perspectives toward such coordinated control efforts is crucial to facilitating engagement. We conducted in person and mail surveys of 68 landowners in and adjacent to the area of a proposed invasive predator control program in New Zealand. We find that, similar to previous studies, landowners consider the potential socioeconomic and ecological benefits of invasive species control and express a strong desire to enhance native biodiversity. However, we also find that landowners take into account the complexity of the local social and ecological context in which a program will unfold in three ways: they consider (1) the level of contribution by other landowners and urban residents who are benefiting from collective control efforts; (2) the potential for the program to upset the local "ecological balance", leading to increases in other pests; and (3) the probability that the program will be successful given the likelihood of others participating and control tactics being effective. We suggest that managers of coordinated invasive species control efforts may benefit from devoting time and resources toward addressing beliefs about social and ecological context, rather than solely providing financial subsidies and information about control tactics or the impacts of invasive species.

Exploiting interspecific olfactory communication to monitor predators.

Olfaction is the primary sense of many mammals and subordinate predators use this sense to detect dominant species, thereby reducing the risk of an encounter and facilitating coexistence. Chemical signals can act as repellents or attractants and may therefore have applications for wildlife management. We devised a field experiment to investigate whether dominant predator (ferret Mustela furo) body odor would alter the behavior of three common mesopredators: stoats (Mustela erminea), hedgehogs (Erinaceus europaeus), and ship rats (Rattus rattus). We predicted that apex predator odor would lead to increased detections, and our results support this hypothesis as predator kairomones (interspecific olfactory messages that benefit the receiver) provoked "eavesdropping" behavior by mesopredators. Stoats exhibited the most pronounced responses, with kairomones significantly increasing the number of observations and the time spent at a site, so that their occupancy estimates changed from rare to widespread. Behavioral responses to predator odors can therefore be exploited for conservation and this avenue of research has not yet been extensively explored. A long-life lure derived from apex predator kairomones could have practical value, especially when there are plentiful resources that reduce the efficiency of food-based lures. Our results have application for pest management in New Zealand and the technique of using kairomones to monitor predators could have applications for conservation efforts worldwide.

Species- and site-specific impacts of an invasive herbivore on tree survival in mixed forests.

Invasive herbivores are often managed to limit their negative impact on plant populations, but herbivore density - plant damage relationships are notoriously spatially and temporally variable. Site and species characteristics (both plant and herbivore) must be considered when assessing the potential for herbivore damage, making it difficult to set thresholds for efficient management. Using the invasive brushtail possum Trichosurus vulpecula in New Zealand as a case study, we parameterized a generic model to predict annual probability of browse-induced mortality of five tree species at 12 sites. We compared predicted and observed tree mortality for each species + site combination to establish herbivore abundance - tree mortality thresholds for each site on a single and combined tree species basis. Model results indicated it is likely that possum browse was the primary cause of all tree mortality at nine of the 12 species-site combinations, allowing us to estimate site-specific thresholds below which possum population numbers should be reduced and maintained to keep tree mortality under a predetermined level, for example 0.5% per year. The browse model can be used to set site- and species-specific management action thresholds, and can be adapted easily for other plant or herbivore species. Results for multiple plant or herbivore species at a single site can be combined to create conservative, site-wide management strategies, and used to: determine which sites will be affected most by changes in herbivore abundance; quantify thresholds for herbivore management; and justify expenditure on herbivore control.

Assessing Movements of Brushtail Possums (Trichosurus vulpecula) in Relation to Depopulated Buffer Zones for the Management of Wildlife Tuberculosis in New Zealand.

In New Zealand, managing the threat of bovine tuberculosis (TB) to livestock includes population reduction of potentially infectious wildlife, primarily the brushtail possum (Trichosurus vulpecula). Population control is often targeted on forested buffer zones adjacent to farmland, in order to limit movements of possums across the buffer and reduce the risk of disease transmission to livestock. To assess the effectiveness of buffers in protecting livestock we analysed GPS telemetry data from possums located in untreated forest adjacent to buffers, and used these data to characterise patterns of movement that could lead to possums reaching farmland during the season when most dispersal occurs. Analyses of movement data showed that the direction of dispersal by sub-adult and adult possums and the extent of long exploratory movements were not biased toward forest buffers, even though these provided vacant habitat as suitable for possums as untreated forest. Instead, dispersal and exploratory movements were uncommon even for sub-adult possums and such events typically lasted <10 days. Dispersing possums settled predominantly in river valleys. A simulation model was developed for the 3-6-month dispersal season; it demonstrated a probability of <0.001 that an infected possum, originating from a low-density population with low disease prevalence in untreated forest, would move across 3 km of recently controlled forest buffer to reach farmland. Our results indicate short-term reduction in the risk of TB transmission from possums to livestock in New Zealand by the use of depopulated buffer zones, while acknowledging that the threat of disease spread from untreated forest is likely to increase over time as possum population density and, potentially, TB prevalence among those possums, increase in the buffer zone.

Climate-based models for pulsed resources improve predictability of consumer population dynamics: outbreaks of house mice in forest ecosystems.

Accurate predictions of the timing and magnitude of consumer responses to episodic seeding events (masts) are important for understanding ecosystem dynamics and for managing outbreaks of invasive species generated by masts. While models relating consumer populations to resource fluctuations have been developed successfully for a range of natural and modified ecosystems, a critical gap that needs addressing is better prediction of resource pulses. A recent model used change in summer temperature from one year to the next (ΔT) for predicting masts for forest and grassland plants in New Zealand. We extend this climate-based method in the framework of a model for consumer-resource dynamics to predict invasive house mouse (Mus musculus) outbreaks in forest ecosystems. Compared with previous mast models based on absolute temperature, the ΔT method for predicting masts resulted in an improved model for mouse population dynamics. There was also a threshold effect of ΔT on the likelihood of an outbreak occurring. The improved climate-based method for predicting resource pulses and consumer responses provides a straightforward rule of thumb for determining, with one year's advance warning, whether management intervention might be required in invaded ecosystems. The approach could be applied to consumer-resource systems worldwide where climatic variables are used to model the size and duration of resource pulses, and may have particular relevance for ecosystems where global change scenarios predict increased variability in climatic events.

Modelling landscape-level numerical responses of predators to prey: the case of cats and rabbits.

Predator-prey systems can extend over large geographical areas but empirical modelling of predator-prey dynamics has been largely limited to localised scales. This is due partly to difficulties in estimating predator and prey abundances over large areas. Collection of data at suitably large scales has been a major problem in previous studies of European rabbits (Oryctolagus cuniculus) and their predators. This applies in Western Europe, where conserving rabbits and predators such as Iberian lynx (Lynx pardinus) is important, and in other parts of the world where rabbits are an invasive species supporting populations of introduced, and sometimes native, predators. In pastoral regions of New Zealand, rabbits are the primary prey of feral cats (Felis catus) that threaten native fauna. We estimate the seasonal numerical response of cats to fluctuations in rabbit numbers in grassland-shrubland habitat across the Otago and Mackenzie regions of the South Island of New Zealand. We use spotlight counts over 1645 km of transects to estimate rabbit and cat abundances with a novel modelling approach that accounts simultaneously for environmental stochasticity, density dependence and varying detection probability. Our model suggests that cat abundance is related consistently to rabbit abundance in spring and summer, possibly through increased rabbit numbers improving the fecundity and juvenile survival of cats. Maintaining rabbits at low abundance should therefore suppress cat numbers, relieving predation pressure on native prey. Our approach provided estimates of the abundance of cats and rabbits over a large geographical area. This was made possible by repeated sampling within each season, which allows estimation of detection probabilities. A similar approach could be applied to predator-prey systems elsewhere, and could be adapted to any method of direct observation in which there is no double-counting of individuals. Reliable estimates of numerical responses are essential for managing both invasive and threatened predators and prey.

Predicted responses of invasive mammal communities to climate-related changes in mast frequency in forest ecosystems.

Predicting the dynamics and impacts of multiple invasive species can be complex because ecological relationships, which occur among several trophic levels, are often incompletely understood. Further, the complexity of these trophic relationships exacerbates our inability to predict climate change effects on invaded ecosystems. We explore the hypothesis that interactions between two global change drivers, invasive vertebrates and climate change, will potentially make matters worse for native biodiversity. In New Zealand beech (Nothofagus spp.) forests, a highly irruptive invasive mammal community is driven by multi-annual resource pulses of beech seed (masting). Because mast frequency is predicted to increase with climate change, we use this as a model system to explore the extent to which such effects may influence invasive vertebrate communities, and the implications of such interactions for native biodiversity and its management. We build on an established model of trophic interactions in the system, combining it with a logistic probability mast function, the parameters of which were altered to simulate either contemporary conditions or conditions of more or less frequent masting. The model predicts that increased mast frequency will lead to populations of a top predator (the stoat) and a mesopredator (the ship rat) becoming less irruptive and being maintained at appreciably higher average abundances in this forest type. In addition, the ability of both current and in-development management approaches to suppress invasive mammals is predicted to be compromised. Because invasive mammals are key drivers of native fauna extinction in New Zealand, with the additional loss of associated functions such as pollination and seed dispersal, these predictions imply potentially serious adverse impacts of climate change for the conservation of biodiversity and ecosystem function. Our study also highlights the importance of long-term monitoring data for assessing and managing future impacts of global change drivers.

Thresholds in plant-herbivore interactions: predicting plant mortality due to herbivore browse damage.

Patterns of herbivore browse at small scales, such as the rate of leaf consumption or plant preferences, drive the impact of herbivores on whole-plant processes, such as growth or survival, and subsequent changes in plant population structure. However, herbivore impacts are often non-linear, highly variable and context-dependent. Understanding the effect of herbivores on plant populations therefore requires a detailed understanding of the relationships that drive small-scale processes, and how these interact to generate dynamics at larger scales. We derive a mathematical model to predict annual rates of browse-induced tree mortality. We model individual plant mortality as a result of rates of foliage production, turnover and herbivore intake, and extend the model to the population scale by allowing for between-tree variation in levels of herbivore browse. The model is configurable for any broadleaved tree species subject to vertebrate or invertebrate browse, and is designed to be parameterized from field data typically collected as part of browse damage assessments. We parameterized and tested the model using data on foliage cover and browse damage recorded on kamahi trees (Weinmannia racemosa) browsed by possums (Trichosurus vulpecula) in New Zealand forests. The model replicated observed patterns of tree mortality at 12 independent validation sites with a wide range of herbivore densities and browse damage. The model reveals two key thresholds; in plant foliar cover, indicating when individual trees may be at high risk from browse-induced mortality, and in herbivore intake, leading to high rates of mortality across the whole population.

Unexpected consequences of control: competitive vs. predator release in a four-species assemblage of invasive mammals.

Invasive species are frequently the target of eradication or control programmes to mitigate their impacts. However, manipulating single species in isolation can lead to unexpected consequences for other species, with outcomes such as mesopredator release demonstrated both theoretically and empirically in vertebrate assemblages with at least two trophic levels. Less is known about the consequences of species removal in more complex assemblages where a greater number of interacting invaders increases the potential for selective species removal to result in unexpected changes in community structure. Using a replicated Before-After Control-Impact field experiment with a four-species assemblage of invasive mammals we show that species interactions in the community are dominated by competition rather than predation. There was no measurable response of two mesopredators (rats and mice) following control of the top predator (stoats), but there was competitive release of rats following removal of a herbivore (possums), and competitive release of mice following removal of rats.

Populations in variable environments: the effect of variability in a species' primary resource.

Mechanistic models for herbivore populations responding to rainfall-driven pasture are used to explore the effect of temporal variability in a primary resource on the abundance and distribution of a species. If the numerical response of the herbivore to pasture is a convex function, then gains made over time intervals with above average rainfall do not compensate for losses incurred when rainfall is below average. Populations therefore fare worse when rainfall is variable compared with when rainfall is reliable. It is demonstrated that this result is independent of the distribution of rainfall. Sensitivity of a species to variability, and hence the limit to its distribution in variable environments, is directly proportional to the difference between population growth rate under ideal conditions and the estimated rate of decline as the species' resource tends to zero. When density dependence is included in the numerical response, the average abundance of a species declines with increasing variability in its primary resource. However, a model for the dynamics of pasture and rabbits (Oryctolagus cuniculus) and red foxes (Vulpes vulpes) in southern Australia, is used to illustrate that trophic interactions can reverse the effect of variability: in the absence of foxes, the mean abundance of rabbits declines with variability as expected, but in the full model the mean abundance of rabbits increases.