Ecology of wildlife rabies in Europe

• 1 The number of wildlife rabies cases has increased in Europe in recent years. We review the epizootiology of wildlife rabies in Europe, paying special attention to recent changes to the situation of two important vector species: the red fox and the raccoon dog. Red fox Vulpes vulpes has been the main vector of rabies since 1945, but the number and proportion of raccoon dog Nyctereutes procyonoides cases has rapidly increased during the past few years, particularly in north‐eastern Europe.
• 2 The transmission rate (average number of susceptible animals infected by each rabid animal) is critical for rabies spread and is partly determined by population density. Both raccoon dogs and foxes live in pairs. Foxes also live in family groups. Pairs and groups share their territories. Home range size usually correlates negatively with population density. Fox home ranges are 50–1500 ha, those of raccoon dogs 150–700 ha. The threshold value for rabies spread among foxes is estimated to be 0.63 individuals/km². Although fox density in eastern and northern Europe may be lower than this, the pooled density of foxes and raccoon dogs exceeds the threshold density.
• 3 Animal movements, especially dispersal of young, pose a risk for rabies spread. Although the likelihood of an epizootic is highest where fox and raccoon dog densities are highest, rabies may spread fastest where population densities are lower, because dispersal distances tend to correlate negatively with population density.
• 4 Oral vaccinations have been more effective in rabies control than culling foxes. Where two vector species exist, vaccination should be conducted twice a year, because most raccoon dogs disperse in autumn but some foxes do not disperse before mid‐ or late winter.
• 5 New rabies models, based on two vector species and their interaction, and which take into account the hibernation period of raccoon dogs, are needed for north‐eastern Europe.

     

Survival and Cause-Specific Mortality of Red Foxes in Agricultural and Urban Areas of Illinois

Abstract: Range expansion and population increase by coyotes (Canis latrans), reduced hunting and trapping, and intensified agricultural practices in the Midwest have altered red fox (Vulpes vulpes) mortality, although relative impacts of these factors are unknown. We examined mortality causes and survival of red foxes in urban and rural agricultural areas of Illinois, using radio telemetry data from 335 foxes (Nov 1996 to May 2002). We used Akaike's Information Criterion to evaluate six survival models for foxes reflecting 1) environmental effects, 2) intrinsic effects, 3) temporal effects, 4) behavioral effects, 5) social effects, and 6) a global model. Environmental and intrinsic models of survival were optimal for adult foxes. Adult foxes with low (0-20%) and high (80-100%) percentages of row crops in their home ranges had higher survival than adults with moderate percentages (40-70%). Heavier adults at capture also survived better. A global model (all covariates) was optimal for juvenile foxes. Higher juvenile survival associated with larger litters, lower body fat, and reduced dispersal time. Yearly survival ranged from 0.18 for rural male juveniles to 0.44 for rural female adults. Adult survival rates (0.35) were 11% higher than juvenile survival rates (0.24). Yearly survival varied for urban foxes due to cyclic outbreaks of sarcoptic mange (Sarcoptes scabei). Thus, summer survival (May-Sep) of urban juveniles ranged from 0.10 (mange present) to 0.83 (no mange recorded). Mange was the most common (45% of all fatalities) source of mortality for urban foxes, followed by road kill (31%). We recorded only 4 mange fatalities (2%) for rural foxes. Rural foxes experienced low hunting mortality (7%) and equivalent road kill and coyote predation fatalities (40% each). Sources of mortality for midwestern foxes have dramatically changed since the 1970s when hunting was the major cause of mortality. Coyote predation has effectively replaced hunting mortality, and cyclic patterns of mange outbreaks in urban fox populations might indicate a dynamic source or sink relationship to surrounding rural fox populations. Absent mange, urban areas might provide refugia for red foxes where coyote populations persist at high densities in rural areas. Managers of sympatric urban and rural wildlife populations must understand survival dynamics influencing the population at the landscape level.     

The impacts and management of foxes Vulpes vulpes in Australia

• 1 The successful introduction of the red fox Vulpes vulpes into Australia in the 1870s has had dramatic and deleterious impacts on both native fauna and agricultural production. Historical accounts detail how the arrival of foxes in many areas coincided with the local demise of native fauna. Recent analyses suggest that native fauna can be successfully reintroduced to their former ranges only if foxes have been controlled, and several replicated removal experiments have confirmed that foxes are the major agents of extirpation of native fauna. Predation is the primary cause of losses, but competition and transmission of disease may be important for some species.
• 2 In agricultural landscapes, fox predation on lambs can cause losses of 1–30%; variation is due to flock size, health and management, as well as differences in the timing and duration of lambing and the density of foxes.
• 3 Fox control measures include trapping, shooting, den fumigation and exclusion fencing; baiting using the toxin 1080 is the most commonly employed method. Depending on the baiting strategy, habitat and area covered, baiting can reduce fox activity by 50–97%. We review patterns of baiting in a large sheep‐grazing region in central New South Wales, and propose guidelines to increase landholder awareness of baiting strategies, to concentrate and coordinate bait use, and to maximize the cost‐effectiveness of baiting programs.
• 4 The variable reduction in fox density within the baited area, together with the ability of the fox to recolonize rapidly, suggest that current baiting practices in eastern Australia are often ineffective, and that reforms are required. These might include increasing landholder awareness and involvement in group control programs, and the use of more efficient broadscale techniques, such as aerial baiting.

     

Effects of Coyote Population Reduction on Swift Fox Demographics in Southeastern Colorado

ABSTRACT The distribution and abundance of swift foxes (Vulpes velox) has declined from historic levels. Causes for the decline include habitat loss and fragmentation, incidental poisoning, changing land use practices, trapping, and predation by other carnivores. Coyotes (Canis latrans) overlap the geographical distribution of swift foxes, compete for similar resources, and are a significant source of mortality amongst many swift fox populations. Current swift fox conservation and management plans to bolster declining or recovering fox populations may include coyote population reduction to decrease predation. However, the role of coyote predation in swift fox population dynamics is not well-understood. To better understand the interactions of swift foxes and coyotes, we compared swift fox population demographics (survival rates, dispersal rates, reproduction, density) between areas with and without coyote population reduction. On the Piñon Canyon Maneuver Site, Colorado, USA, we monitored 141 swift foxes for 65,226 radio-days from 15 December 1998 to 14 December 2000 with 18,035 total telemetry locations collected. Juvenile swift fox survival rate was increased and survival was temporarily prolonged in the coyote removal area. Adult fox survival patterns were also altered by coyote removal, but only following late-summer coyote removals and, again, only temporarily. Coyote predation remained the main cause of juvenile and adult fox mortality in both areas. The increase in juvenile fox survival in the coyote removal area resulted in a compensatory increase in the juvenile dispersal rate and an earlier pulse in dispersal movements. Adult fox dispersal rate was more consistent throughout the year in the coyote removal area. Coyote removal did not influence the reproductive parameters of the swift foxes. Even though juvenile survival increased, swift fox density remained similar between the areas due to the compensatory dispersal rate among juvenile foxes. We concluded that the swift fox population in the area was saturated. Although coyote predation appeared additive in the juvenile cohort, it was compensatory with dispersal.     

Can widespread generalist predators affect keystone prey? A case study with red foxes and European rabbits in their native range

Widespread generalist predators may affect declining keystone prey populations. However, this phenomenon is not well understood. In this paper, we assessed whether the abundance and population growth of European rabbits Oryctolagus cuniculus, a keystone prey species in Mediterranean Iberia, was related to the abundance and diet of red foxes, Vulpes vulpes, a widespread generalist predator. In a locality in central Spain, where rabbit population abundance declined, we estimated rabbit abundance during almost 3 years and determined fox abundance and diet during two concurrent years. We calculated a fox predation index (percentage of consumed rabbit biomass × fox abundance) to assess the importance of rabbits to foxes. We employed a multi–model approach to explain rabbit abundance and population growth. Foxes consumed between 60 and 99 % rabbit biomass in their diets, and this was independent of rabbit abundance. Periods of higher fox predation index coincided with lower rabbit density and vice versa. Two models best explained rabbit abundance and four rabbit population growth. They included the fox predation index and its interaction with rabbit abundance during the previous month. Altogether, fox predation, intraspecific density dependence, and their interaction partly explained rabbit population dynamics. We conclude that in order to propel the recovery of the rabbit in Iberia, it is essential to better understand the role of these factors in driving the abundance of the species.     

Toward reliable population density estimates of partially marked populations using spatially explicit mark–resight methods

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Lethal 1080 baiting continues to reduce European Red Fox (Vulpes vulpes) abundance after more than 25 years of continuous use in south‐west Western Australia

European Red Fox (Vulpes vulpes) baiting with 1080 poison (sodium fluoroacetate) is undertaken in many Australian sites to reduce fox abundance and to protect vulnerable native species from predation. The longest continuous use of fox baiting for fauna conservation commenced in south‐west Western Australia in the 1980s and includes baiting Dryandra Woodland and Tutanning Nature Reserve. The trap success of the Woylie (Bettongia penicillata) in these two reserves initially increased more than 20‐fold after the commencement of baiting and was maintained until 2000. Woylie captures then decreased rapidly, despite ongoing fox baiting, so the long‐term efficacy of 1080 baiting was questioned. Here, fox density and probabilities of detection, re‐detection and survival between replicated baited and unbaited sites were compared by modelling capture–recapture of individual foxes. These were identified from microsatellite DNA genotypes obtained non‐invasively from hair, scat and saliva samples. The frequency and duration of fox residencies were also quantified. Remote cameras were used to determine the fate of baits but uptake by foxes was low, whereas nontarget species' bait uptake was high. Nevertheless, foxes inhabiting baited reserves had significantly higher mortality, shorter residency times, and 80% lower density than foxes inhabiting unbaited reserves. Baiting continues to significantly reduce fox abundance after more than 25 years of continuous use. This has positive implications for fox control programmes throughout Australia but reduced fox abundance may facilitate increased predation by feral Cats (Felis catus).     

Landscape-Genetic Analysis of Population Structure in the Texas Gray Fox Oral Rabies Vaccination Zone

ABSTRACT In west-central Texas, USA, abatement efforts for the gray fox (Urocyon cinereoargenteus) rabies epizootic illustrate the difficulties inherent in large-scale management of wildlife disease. The rabies epizootic has been managed through a cooperative oral rabies vaccination program (ORV) since 1996. Millions of edible baits containing a rabies vaccine have been distributed annually in a 16-km to 24-km zone around the perimeter of the epizootic, which encompasses a geographic area >4 × 10⁵ km². The ORV program successfully halted expansion of the epizootic into metropolitan areas but has not achieved the ultimate goal of eradication. Rabies activity in gray fox continues to occur periodically outside the ORV zone, preventing ORV zone contraction and dissipation of the epizootic. We employed a landscape-genetic approach to assess gray fox population structure and dispersal in the affected area, with the aim of assisting rabies management efforts. No unique genetic clusters or population boundaries were detected. Instead, foxes were weakly structured over the entire region in an isolation by distance pattern. Local subpopulations appeared to be genetically non-independent over distances >30 km, implying that long-distance movements or dispersal may have been common in the region. We concluded that gray foxes in west-central Texas have a high potential for long-distance rabies virus trafficking. Thus, a 16-km to 24-km ORV zone may be too narrow to contain the fox rabies epizootic. Continued expansion of the ORV zone, although costly, may be critical to the long-term goal of eliminating the Texas fox rabies virus variant from the United States.     

Demonstrating frequency-dependent transmission of sarcoptic mange in red foxes

Understanding the relationship between disease transmission and host density is essential for predicting disease spread and control. Using long-term data on sarcoptic mange in a red fox Vulpes vulpes population, we tested long-held assumptions of density- and frequency-dependent direct disease transmission. We also assessed the role of indirect transmission. Contrary to assumptions typical of epidemiological models, mange dynamics are better explained by frequency-dependent disease transmission than by density-dependent transmission in this canid. We found no support for indirect transmission. We present the first estimates of R₀ and age-specific transmission coefficients for mange in foxes. These parameters are important for managing this poorly understood but highly contagious and economically damaging disease.     

Invading parasites cause a structural shift in red fox dynamics

The influence of parasites on host life histories and populations is pronounced. Among several diseases affecting animal populations throughout the world, sarcoptic mange has influenced many carnivore populations dramatically and during the latest epizootic in Fennoscandia reduced the abundance of red fox by over 70%. While the numerical responses of red fox populations, their prey and their competitors as well as clinical implications are well known, knowledge of how sarcoptic mange affects the structure of the dynamics of red fox populations is lacking. Integrating ecological theory and statistical modelling, we analysed the long-term dynamics (1955-1996) of 14 Danish red fox populations. As suggested by the model, invading sarcoptic mange significantly affected direct and delayed density dependence in red fox dynamics and concomitant shifts in fluctuation patterns were observed. Our statistical analyses also revealed that the spatial progressive spread of mange mites was mirrored in the autocovariate structures of red fox populations progressively exposed to sarcoptic mange.     

Not all predators are equal: a continent‐scale analysis of the effects of predator control on Australian mammals

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Does culling reduce fox (Vulpes vulpes) density in commercial forests in Wales, UK?

Forests within agricultural landscapes can act as safe harbourages for species that conflict with neighbouring landowners’ interests, including mammalian predators. The agency responsible for the management of forests in upland Wales, UK, has permitted the killing of foxes (Vulpes vulpes) on their land as a “good neighbour policy” with the aim of reducing fox numbers. The principal method used was the use of dogs to drive foxes to a line of waiting shooters; a small number of foxes were also killed by shooting at night with a rifle. However, it has been postulated that recent restrictions on the use of dogs to kill foxes in Britain could lead to an increase in fox numbers in plantations. The aim of this study was to determine whether over-winter culling (i.e. driving foxes to guns using dogs and rifle shooting) in these forests acted to reduce fox density. Fox faecal density counts were conducted in commercial forests in Wales in autumn 2003 and spring 2004. Data were analysed from 29 sites (21 individual forests and 8 forest blocks, the latter consisting of pooled data from 20 individual forests). The over-winter change in faecal density was negatively related to the proportion of felled land and the proportion of land more than 400 m altitude, these associations probably reflecting reduced food availability. Over-winter change was positively associated with culling pressure (i.e. more foxes were killed where more foxes were present, or vice versa), but this was not significant. The number of foxes killed was large relative to the estimated resident population, but losses appeared to be negated (most likely) by immigration. Pre-breeding (spring) faecal density counts were significantly positively related to culling pressure, i.e. more foxes were killed with increased fox density or vice versa. Overall, there was no evidence to suggest that culling reduced fox numbers. Consequently, restrictions on the use of dogs to control foxes are unlikely to result in an increase in fox numbers in commercial forests.     

Identifying and Controlling for Variation in Canid Harvest Data

An accurate understanding of harvest trends is required for effective wildlife management. Trapper harvest data represent valuable long-term data for evaluating patterns and trends for wildlife species at broad spatiotemporal scales. Inferring accurate trends from harvest data, however, first requires identifying and controlling for confounding factors that vary independent of abundance. We investigated trends in 43 years of trapper harvest data (1976–2018) from Illinois, USA, for red fox (Vulpes vulpes), gray fox (Urocyon cinereoargenteus), and coyote (Canis latrans) while controlling for factors that may affect trapper effort, including number of effective (i.e., successful) trappers, pelt price, gasoline price, winter unemployment, and winter weather conditions. Annual trapper harvest for red and gray foxes declined and was affected by gasoline price and winter unemployment, whereas annual trapper harvest for coyotes increased and was not strongly affected by other covariates. After adjusting for pelt price, harvest of red foxes was relatively stable, but harvest of gray foxes declined and harvest of coyotes increased. Effects of covariates on harvest per successful trapper varied by species; nevertheless, we detected an increasing trend for coyotes and decreasing trends for gray foxes and red foxes. Concordance across indices for gray foxes and coyotes was consistent with hypothesized declines for gray foxes and increases for coyotes in the midwestern United States. Trends for red foxes varied depending on how we accounted for potential confounding factors and it is unclear if these trends suggest population declines or distribution shifts to urban areas with reduced trapping susceptibility. Our results highlight the importance of understanding sources of variation in harvest data and that their effects can vary across species. © 2020 The Wildlife Society.     

Numbers and distribution of the Great Cormorant in Iceland: Limitation at the regional and metapopulation level

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The status of foxes Vulpes vulpes on the Isle of Man in 1999

1. Foxes Vulpes vulpes probably did not occur naturally on the Isle of Man, but were present in small numbers in the mid 19th century. They were introduced again in the 1980s, and in 1990 the population was estimated at 120–300 individuals (20–50 per 100 km²) on the basis of field signs. We used a nocturnal spotlight transect survey technique to assess the status of foxes on the Isle of Man in September 1999. This method had previously been used effectively to estimate fox densities in populations as sparse as 16 per 100 km². We surveyed a total of 852 km over a period of eight consecutive nights, during weather conditions which allowed excellent visibility. No foxes were seen. 2. By comparison with equivalent survey efforts in reference areas of mainland Britain, we conclude that post‐breeding fox density on the Isle of Man was certainly below 2.5 per 100 km², implying a maximum of 15 foxes on the entire island. We estimate a probability of only 15–25% that a fox population of 1 per 100 km² was present but not detected; this would be equivalent to only five or six individuals on the whole island. Foxes may even be entirely absent, although unsubstantiated sightings continue to be reported. 3. This finding is significant in understanding the ecology of the Isle of Man, and in planning the conservation of a number of ground‐nesting bird species there, as well as for farming interests, and for contingency planning against an accidental introduction of rabies.     

On food preference in the Red fox

• 1 Foxes treat different prey species in a variety of ways, eating, burying, or discarding them on the basis of preference.
• 2 Because foxes often utilize cached food and because they can remember which prey species is in a given cache the preference effect can be longstanding.
• 3 Evidence from the literature suggests that comparable effects of preference for different species of small mammals affects the diet of wild foxes and their behaviour in the same way as demonstrated in these experiments.
• 4 Certain distastes appear common to all foxes and most carnivores, for instance, insectivore and carnivore meat, in particular that of their own species.
• 5 An incidence of active cannibalism by a fox is reported.
• 6 The effect of food preference is shown to change during the course of one individual's life perhaps as a consequence of such factors as rearing cubs and competition for food.
• 7 One effect of the behavioural consequences of food preference is to defer the decision of what to eat.
• 8 That the fox will kill animals that it does not eat means that populations of animals that are not strictly fox prey are still at risk from fox predatory activity.

     

Europe‐wide biogeographical patterns in the diet of an ecologically and epidemiologically important mesopredator,the red fox Vulpes vulpes: a quantitative review

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Ineffectiveness of plastic mesh for protecting artificial freshwater turtle nests from red fox (Vulpes vulpes) predation

Invasive mammalian predators are efficient at driving native animal declines. The red fox (Vulpes vulpes) kills millions of endemic reptiles in Australia each year. In areas of south-eastern Australia, the eastern long-necked turtle (Chelodina longicollis) and Murray River turtle (Emydura macquarii) have declined by more than 50%. High rates of nest predation by foxes limit the recruitment of young turtles in these populations, but previous methods of fox control have been ineffective at protecting turtle nests. Here, we tested the effectiveness of plastic mesh for protecting artificial turtle nests from predation by foxes, in the mid-Murray catchment, Victoria. We also tested whether protecting a large number of artificial nests in a given area encourages foxes to give up foraging following predictions from giving-up density theory. We made a series of plots, each containing 32 artificial turtle nests. In each plot, we covered a percentage (0%, 25%, 50%, 81% or 100%) of the nests with either 1 or 2 sheets of plastic mesh. We used remote cameras to photograph and identify any predator that attacked nests in the plots. The cameras also allowed us to estimate the amount of time a fox was visible on each plot, as a metric of how much effort foxes expended on protected nests. Nest survival rate was not increased by either 1 or 2 sheets of mesh, and increasing the number of protected nests on a plot did not reveal a giving-up density (GUD) value for fox foraging behaviour. Our study demonstrates that plastic mesh is not effective for protecting artificial turtle nests from foxes in this region.     

Clinical Picture and Antibody Response to Experimental Sarcoptes scabiei var. vulpes Infection in Red Foxes (Vulpes vulpes)

Three red foxes (Vulpes vulpes) were experimentally infected with Sarcoptes scabiei isolated from a naturally infected wild red fox. A fourth red fox served as a control. The first signs of sarcoptic mange became evident on the 31st day post infection (dpi). The signs gradually increased thereafter and between dpi 49 and 77 characteristic lesions of hyperkeratosis developed. Two of the infected foxes developed severe sarcoptic mange, and one of these animals died on dpi 121. The third fox developed a chronic hyperkeratotic lesion on its back, at the site where the mites had been applied. On dpi 127 the surviving foxes were treated systemically with ivermectin, and within 4 weeks the skin lesions had healed except on the pinnae of one animal. Antibodies to S. scabiei var. vulpes were demonstrated in the infected foxes by an ELISA with which seroconversion was seen around 4 weeks post infection (wpi). Western blot analysis of sequential sera of the infected animals demonstrated antibody activity consistently after the 2nd wpi. The fourth, non-infected, fox did not show any skin lesions throughout the experimental period nor any specific antibodies to S. scabiei var. vulpes. kw|Keywords|k]sarcoptic mange; k]red fox; k]serodiagnosis; k]ELISA     

Evaluation of Camera Trapping for Estimating Red Fox Abundance

ABSTRACT The nature reserve Serra da Malcata, Portugal, was recently considered a site for Iberian lynx (Lynx pardinus) reintroduction. Because of potential disease risk posed by red foxes (Vulpes vulpes) in the area, a reliable estimate of fox abundance was critical for a dependable reintroduction program. We adapted camera-trapping techniques for estimating red fox abundance in the reserve. From July 2005 to August 2007, we conducted 7 camera-trapping sessions, allowing for individual identification of foxes by physical characteristics. We estimated abundance using the heterogeneity (M_h) model of the software program CAPTURE. Estimated density ranged from 0.91 ± 0.12 foxes/km²to 0.74 ± 0.02 foxes/km². By estimating red fox density, it is possible to define the number of foxes that must be sampled to assess the presence of potential fox-transmitted diseases that may affect lynx reintroduction.