Mammals and agri-environment schemes: hare haven or pest paradise?

1. Agri-environment schemes (AESs) are designed to create landscape-scale improvements in biodiversity. While the specific aims of AESs do not always include the enhancement of species of conservation concern, associated conservation strategies, such as the UK Biodiversity Action Plan, often rest on the assumption that AESs enhance environmental conditions and thereby improve the conservation status of target species. However, there is little evidence for the general efficacy of AESs in this respect. 2. To evaluate the effects of the Environmentally Sensitive Area (ESA) scheme, a widespread AES in Northern Ireland, a spotlight survey of the relative abundance of three mammal species, Irish hare Lepus timidus hibernicus, European rabbit Oryctolagus cuniculus and red fox Vulpes vulpes, was conducted. Of these, the Irish hare is a priority species for conservation action and the focus of a species action plan, while rabbit and fox are commonly considered agricultural pests. The effects of ESA designation and habitat on each species were assessed at 150 ESA and 50 non-ESA sites, matched for landscape characteristics. 3. The ESA scheme had no demonstrable effect on the abundance of Irish hares, and this agri-environment scheme did not target the landscape and habitat variables associated with hares. 4. In contrast, the abundance of rabbits and foxes was significantly greater within ESAs than the wider countryside. Agricultural factors such as reduced livestock stocking density, reduced overgrazing and field boundary enhancements may create more favourable conditions for both species. Aside from the implications for farm economics, the proliferation of rabbit populations within conservation areas may raise issues concerning the grazing of important plant communities, while increases in fox populations may adversely affect ground-nesting birds and other animal species of conservation concern. 5. Synthesis and applications: The abundance of rabbits and foxes corroborates recent work that suggests AESs may benefit common species but can not be relied upon to encourage rarer species. The Irish hare species action plan relies on agri-environment schemes to enhance the species’ status and realize the target of increasing the hare population by 2010 by promoting suitable habitat. However, the ESA scheme is unlikely to help in achieving these objectives. Targeted and evidence-based agri-environment prescriptions are clearly required in order to ensure the realization of species-specific conservation targets.

Habitat-species interactions in a carnivore community

Na maioria das area protegidas nacionais existe uma considerável falta de informação científica relativa aos mamíferos carnívoros. A Reserva Natural da Serra da Malcata, localizada no centro-este de Portugal, tem vindo a desenvolver, desde os últimos 20 anos, estudos de ecologia e esta tese pretende dar seguimento a esse esforço de monitorização, através do desenvolvimento de métodos simples e eficientes, de monitorização de carnívoros, que possam servir como percursores de trabalhos a longo-prazo em áreas relevantes para a conservação. Através do uso de armadilhagem fotográfica, foram estudadas relações espécies-habitats para 5 espécies: gato-bravo (Felis silvestris), fuinha (Martes foina), raposa (Vulpes vulpes), gineta (Genetta genetta), e sacarrabos (Herpestes ichneumon). Foram desenvolvidos métodos para se determinar a densidade absoluta de raposa e gineta e a população de gato-bravo foi estudada em detalhe. As principais conclusões do estudo foram: 1) a ocupação de raposas é uniforme e parece ser independente de variáveis ambientais; 2) a ocupação de fuinha encontra-se relacionada com variáveis de habitat, estrutura paisagística e presas; 3) a ocupação de gineta está relacionada com a cobertura de folhosas e distribuição de presas; 4) para o sacarrabos verificase que a ocupação é influenciada pelas extensões de habitat arbustivos, 5) a população de gato-bravo sofreu um forte declínio durante o trabalho e requer urgentes medidas de conservação. Metodologicamente foi demonstrada a importância da modelação das probabilidades de detecção para espécies para as quais este parâmetros apresenta valores baixos ou é muito variável. Esta tese também demonstrou a grande importância da Serra da Malcata para a conservação de carnívoros e a necessidade do desenvolvimento de técnicas de monitorização padronizadas para uma correcta gestão adaptativa das áreas protegidas.

Population and landscape genetics of the stone marten and red fox in Portugal : implications for conservation management of common carnivores

Tese de doutoramento, Biologia (Biologia da Conservação), Universidade de Lisboa, Faculdade de Ciências, 2015

Behavioral changes associated with a population density decline in the facultatively social red fox

Understanding the causal mechanisms promoting group formation in carnivores has been widely investigated, particularly how fitness components affect group formation. Population density may affect the relative benefits of natal philopatry versus dispersal. Density effects on individual behavioral strategies have previously been studied through comparisons of different populations, where differences could be confounded by between-site effects. We used a single population of red foxes (Vulpes vulpes) in the city of Bristol, UK, that underwent a natural perturbation in density to compare key changes in 1) group structure, 2) within-group relatedness, 3) mating system, 4) dispersal, and 5) dominance attainment. At high densities (19.6-27.6 adults km(-2)), group sex ratios were equal and included related and unrelated individuals. At low densities (4.0-5.5 adults km(-2)), groups became female biased and were structured around philopatric females. However, levels of within-group relatedness were unchanged. The genetic mating patterns changed with no instances of multiple-paternity litters and a decline in the frequency of extrapair litters of cubs from <= 77% to <= 38%. However, the number of genetically monogynous groups did not differ between periods. Dispersal was male biased at both high and low densities. At high density, most dominant males in the study groups appeared to have gained dominance after dispersing, but natal philopatry was an equally successful strategy at low density; conversely, most dominant females were philopatric individuals at both high and low densities. These results illustrate how density may alter behavioral strategies such as mating patterns and how this, in turn, alters group structure in a single population.

Environmental variation at the onset of independent foraging affects full-grown body mass in the red fox

The period following the withdrawal of parental care has been highlighted as a key developmental period for juveniles. One reason for this is that juveniles cannot forage as competently as adults, potentially placing them at greater risk from environmentally-induced changes in food availability. However, no study has examined this topic. Using a long-term dataset on red foxes (Vulpes vulpes), we examined (i) dietary changes that occurred in the one-month period following the attainment of nutritional independence, (ii) diet composition in relation to climatic variation, and (iii) the effect of climatic variation on subsequent full-grown mass. Diet at nutritional independence contained increased quantities of easy-to-catch food items (earthworms and insects) when compared with pre-independence. Interannual variation in the volume of rainfall at nutritional independence was positively correlated to the proportion of earthworms in cub diet. Pre-independence cub mass and rainfall immediately following nutritional independence explained a significant proportion of variance in full-grown mass, with environmental variation affecting full-grown mass of the entire cohorts. Thus, weather-mediated availability of easy-to-catch food items at a key developmental stage has lifelong implications for the development of juvenile foxes by affecting full-grown mass, which in turn appears to be an important component of individual reproductive potential.

Urban mammals: what does the future hold? An analysis of the factors affecting patterns of use of residential gardens in Great Britain

1Urban areas are predicted to grow significantly in the foreseeable future because of increasing human population growth. Predicting the impact of urban development and expansion on mammal populations is of considerable interest due to possible effects on biodiversity and human-wildlife conflict. 2The British government has recently announced a substantial housing programme to meet the demands of its growing population and changing socio-economic profile. This is likely to result in the construction of high-density, low-cost housing with small residential gardens. To assess the potential effects of this programme, we analysed the factors affecting the current pattern of use of residential gardens by a range of mammal species using a questionnaire distributed in wildlife and gardening magazines and via The Mammal Society. 3Twenty-two species/species groups were recorded. However, the pattern of garden use by individual species was limited, with only six species/species groups (bats, red fox Vulpes vulpes, grey squirrel Sciurus carolinensis, hedgehog Erinaceus europaeus, mice, voles) recorded as frequent visitors to > 20% of gardens in the survey. 4There was a high degree of association between the variables recorded in the study, such that it was difficult to quantify the effects of individual variables. However, all species/species groups appeared to be negatively affected by the increased fragmentation and reduced proximity of natural and semi-natural habitats, decreasing garden size and garden structure, but to differing degrees. Patterns of garden use were most clearly affected by house location (city, town, village, rural), with garden use declining with increasing urbanization for the majority of species/species groups, except red foxes and grey squirrels. Increasing urbanization is likely to be related to a wide range of interrelated factors, any or all of which may affect a range of mammal species. 5Overall, the probable effects of the planned housing development programme in Britain are not likely to be beneficial to mammal populations, although the pattern of use examined in this study may represent patterns of habitat selection by species rather than differences in distribution or abundance. Consequently, additional data are required on the factors affecting the density of species within urban environments.

Change in the diet of sooty owls (Tyto tenebricosa) since European settlement: from terrestrial to arboreal prey and increased overlap with powerful owls

The current diet of the sooty owl (Tyto tenebricosa) was determined by analysing freshly regurgitated pellets collected beneath their roosting sites in East Gippsland, Victoria. Comparisons were then made with: (i) prehistoric and historic diet from bone deposits found in cave roosts, and (ii) diet of a sympatric owl species, the powerful owl (Ninox strenua). Sooty owls consumed a large array of terrestrial mammal species before European settlement, but only three terrestrial species were detected in their current diet, a reduction of at least eight species since European settlement. To compensate, sooty owls have increased their consumption of arboreal prey from 55% to 81% of their diet. Arboreal species are also a major component of the powerful owl diet and this prey shift by sooty owls has increased dietary overlap between these two species. Predation by foxes (Vulpes vulpes) and other feral species is likely to have reduced the amount of terrestrial prey available to sooty owls since European settlement. Investigation of changes in the diet of sooty owls may offer a unique monitoring system for evaluating the ability of fox-control strategies to influence increases in numbers of critical-weight-range mammals.

Camera traps : the research paparazzi

The use of cameras to monitor wildlife is commonplace; however, little is known of the effectiveness of different camera technologies for the detection of mammals. We compared the detection success of three different camera systems, a passive infrared (IR) system, an active IR and a constant video camera, alongside a trapping grid of Elliott and cage traps to determine their effectiveness at detecting mammals at multiple locations in the Otways National Park, Victoria, Australia (n = 160 events; 40 ± 23 [SD] events per night). Species detected and detection rates differed between methods (χ2 = 57.95, df = 2, p < 0.0001). Only house mice (Mus musculus) were detected by camera and traditional trapping techniques. Camera systems alone detected foxes (Vulpes vulpes) and a koala (Phascolarctos cinereus), while traditional traps captured bush rats (Rattus fuscipes), agile antechinus (Antechinus agilis) and a brush-tailed possum (Trichosurus vulpecula) which were not detected by the camera systems. Assuming that the video camera detected all mammals at the camera trap, the passive IR system detected almost all mammals detected by the video and it detected significantly more species than the active IR system. The choice of method will ultimately depend on the species of interest, logistics and the study site, and may substantially influence the results of a study.

Does baiting influence the relative composition of the diet of foxes?

The changes in the diet of foxes (Vulpes vulpes) in the Jervis Bay Region was assessed following a long-term baiting program by analysing the composition of fox faecal excreta (scats). In all, 470 fox scats were collected between April and August 2003 from two baited sites, Booderee National Park (BNP) and Beecroft Peninsula, and from two unbaited sites in the southern and northern parts of Jervis Bay National Park (SJBNP and NJBNP respectively). Diet was compared between these sites and mammalian diet was also compared from scats collected before baiting in 1996 and after baiting in 2000 at Beecroft Peninsula and in 2001 at Booderee National Park. In 2003, the most common species consumed by foxes was the common ringtail possum (Pseudocheirus peregrinus), except at unbaited NJBNP, where the swamp wallaby (Wallabia bicolor) was the most frequent dietary item. Significant dietary differences were found between unbaited and baited sites, with the long-nosed bandicoot (Perameles nasuta) and P. peregrinus featuring more in the diet of foxes from the baited sites. Marked increases in the frequency of occurrence of P. peregrinus and P. nasuta in fox scats occurred from before baiting through to after baiting. Relative fox abundance, as indexed by the number of scats collected per kilometre, was lowest in Booderee, followed by Beecroft, then SJBNP, with NJBNP having the highest relative abundance of foxes. We suggest that baiting did affect the diet of foxes on both peninsulas and that the dietary changes across baiting histories were intrinsically related to an increase in abundance in some taxa as a result of relaxed predator pressure following sustained fox control. However, the lack of unbaited control sites over the whole study precludes a definitive conclusion.

Breeding behaviour and ecology of the sexually size-dimorphic brown songlark, Cinclorhamphus cruralis

The Australian endemic brown songlark, Cinclorhamphus cruralis, is one of the most sexually size-dimorphic of all birds, and yet its breeding ecology remains poorly documented. Here we redress this situation by describing the breeding activities of brown songlarks over three years (1998–2000) in the semi-arid grasslands of south-western New South Wales. Study populations of this nomadic species were selected in late August of each year on the basis of high adult abundance. Adult males at these sites were, on average, 2.3 times heavier than females. Over the three seasons, nesting activities started in early to late August and continued until early November or December. Males were highly polygynous and, on average, occupied territories of about 4.0 ha. Nests were well concealed at the base of small shrubs and grass tussocks or in thick herbage. Clutches ranged in size from 2 to 5 eggs (mean 3.2) and were incubated exclusively by the female for 11–13 days (mean 12.1). Nestlings received a range of invertebrate prey, mainly from the female, for 10–14 days (mean 11.5) before leaving the nest. Only 17% of nesting attempts were estimated to be successful, and each of these nests produced an average of 2.7 fledglings. Predators, including foxes, Vulpes vulpes, and brown snakes, Pseudonaja textilis, were the main cause of nest failure. Some females produced replacement clutches following nest failure, while others laid second clutches after the success of an earlier brood. We speculate that extreme size dimorphism has evolved in this species because (i) males compete physically for breeding territories, and (ii) habitat heterogeneity and excellent visibility of their surroundings allow some males to defend territories of sufficient size to support nesting by multiple females

Conditioned taste aversion reduces fox depredation on model eggs on beaches

Context:  Conditioned taste aversion (CTA) is induced by an association of a food item with a negative experience, such as illness, which causes animals to avoid subsequent consumption of that particular food item. Inducing CTA may help reduce depredation rates of threatened fauna where predator population control is undesirable, impractical or unsuccessful.

Aims:  We investigated whether CTA could be induced among foxes (Vulpes vulpes) to model eggs which mimicked those of the threatened hooded plover (Thinornis rubricollis).

Methods:   Model eggs treated with a potential CTA-inducing chemical (sodium carbonate) and control eggs free of the agent were exposed to fox depredation for 28 days to simulate a hooded plover incubation period. To investigate whether CTA would persist in wild foxes, we implemented a part-time agent treatment (an initial 14 day exposure period of model eggs with the CTA agent followed by a second 14 day period when model eggs were free of the agent).

Key results:  Similar intervals to the first depredation event were found for all model eggs regardless of treatment. After the first depredation event by foxes, the rate and likelihood of fox depredation was significantly lower in treated eggs than in control eggs. The likelihood or rate of depredation across the three treatments did not differ between the first and second periods.

Conclusions: Our results suggest that during an exposure period of at least 28 days, CTA can be induced in wild foxes to eggs on beaches. Our results also suggest that 14 days may be insufficient time for wild foxes to develop a lasting CTA to familiar food items such as eggs.

Implications:  Treatment of eggs with a CTA-inducing chemical may present a viable alternative to traditional predator control techniques for hooded plovers, as well as other ground-nesting birds, provided that an extended exposure to the CTAinducing agent occurs.

Top predators as biodiversity regulators : the dingo Canis lupus dingo as a case study

Top-order predators often have positive effects on biological diversity owing to their key functional roles in regulating trophic cascades and other ecological processes. Their loss has been identified as a major factor contributing to the decline of biodiversity in both aquatic and terrestrial systems. Consequently, restoring and maintaining the ecological function of top predators is a critical global imperative. Here we review studies of the ecological effects of the dingo Canis lupus dingo, Australia's largest land predator, using this as a case study to explore the influence of a top predator on biodiversity at a continental scale. The dingo was introduced to Australia by people at least 3500 years ago and has an ambiguous status owing to its brief history on the continent, its adverse impacts on livestock production and its role as an ecosystem architect. A large body of research now indicates that dingoes regulate ecological cascades, particularly in arid Australia, and that the removal of dingoes results in an increase in the abundances and impacts of herbivores and invasive mesopredators, most notably the red fox Vulpes vulpes. The loss of dingoes has been linked to widespread losses of small and medium-sized native mammals, the depletion of plant biomass due to the effects of irrupting herbivore populations and increased predation rates by red foxes. We outline a suite of conceptual models to describe the effects of dingoes on vertebrate populations across different Australian environments. Finally, we discuss key issues that require consideration or warrant research before the ecological effects of dingoes can be incorporated formally into biodiversity conservation programs.

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator–prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey.

2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs.

3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs.

4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3·5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators.

5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes.

6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.

Surviving on cached foods - the energetics of egg-caching by arctic foxes

Food-caching by arctic foxes (Vulpes lagopus (L., 1758)) is a behavioural adaptation thought to increase winter survival, especially in bird colonies where a large number of eggs can be cached during a short nesting season. In this paper, we measured the energy content of greater snow goose (Chen caerulescens atlantica Kennard, 1927) eggs and evaluated their perishability when cached in tundra soil for a whole summer. We estimated that eggs lost only ~8% of their dry mass over 60 days of storage in the ground. We used published estimates on digestibility of nutrients by arctic foxes to estimate that fresh and stored goose eggs contained 816 and 730 kJ of metabolizable energy, respectively, a difference of 11%. Using information on arctic fox energetics, we evaluated that 145 stored eggs were required to sustain the growth of one pup from the age of 1 to 3 months (nutritional independence). Moreover, 23 stored eggs were energetically equivalent to the average fat deposit of an arctic fox during winter. Finally, we calculated that an adult arctic fox would need to recover 160-220 stored eggs to survive 6 months in resting conditions during cold winter temperatures. This value increased to 480 when considering activity cost. Based on egg acquisition and caching rates observed in many goose colonies, we conclude that cached eggs represent an important source of energy relative to the needs of an arctic fox during winter, and have thus a high fitness value.

Effect of Direct and Indirect Cues of Predation Risk on the Foraging Behavior of the White-Footed Mouse (Peromyscus leucopus)

Understanding predator-prey dynamics requires an understanding of how prey assess predation risk. This study tested the effect of microhabitat, moon stages, and mammalian predator urines (Vulpes vulpes [Red Fox], Mustela vison [Mink], and Procyon lotor [Raccoon]) on the degree of predation risk perceived by Peromyscus leucopus (White-footed Mouse). Giving-up densities from artificial food patches were used to quantify perceived predation risk. White-footed Mice exhibited a strong preference for cover microhabitat and for the new moon stage. However, the mice did not significantly alter their foraging behavior in response to the predator urines compared to a water control. Additionally, mice foraged less on colder nights. The results suggest that mammalian predator urines may not provide reliable information on actual predation risk for the White-footed Mice and that the mice extensively use indirect cues to assess predation risk.