Space use and resting site selection of red foxes (<Emphasis Type="Italic">Vulpes vulpes</Emphasis>) living near villages and small towns in Southern Germany

In 2005–2008, we radio-tracked 17 foxes in rural areas of Southern Germany. The mean home range size was 76.6 ha (95% MCP) or 138.9 ha (95% fixed kernel), and the built-up area formed an integral part of the home range. Home ranges of juvenile foxes were significantly smaller than home ranges of adult foxes. Gender-specific differences among adult foxes were not established. A minimum population density of 2.7 foxes per km² and summer densities of up to 13.4 foxes per km² were calculated. Therefore, the fox density was three to eight times higher than that of strictly rural foxes. Daytime resting sites of foxes were mostly found in forests (62.2%) and reedbed areas (20.6%). Of the resting sites, 14.8% were situated inside settlements, in fallow gardens or gardens of residents. During the day, foxes exhibited habitat preferences for forests and reedbed areas. A habitat structure that offers plenty of cover or dense vegetation is essential for its selection as a safe resting site. If this basic requirement is fulfilled, foxes also choose resting sites within settlements, and are not disturbed by human presence.     

Complex housing environment for farmed blue foxes (Vulpes lagopus): use of various resources

The present study was designed to measure the use of various, simultaneously available resources in a complex housing environment in juvenile blue foxes. Twelve blue fox sibling (male–female) pairs were housed in two-section experimental cages from the age of 8 weeks until the age of 7 months (from June to December). Each experimental cage was furnished with two platforms, a nest box, a sand box and a wooden block. This housing set-up provided the foxes with social contact, and an opportunity for oral manipulation, scratching and nesting, as well as the choice of staying on a solid floor material or on an elevated location. The foxes’ behaviour was recorded at three time points during autumn (September, November and December). The foxes used all available resources. The most utilised resource was the nest box, possibly because it could be utilised in several ways (as a shelter, an elevated location, an object for scratching and for oral manipulation). The foxes also stayed more in the cage section containing the nest box than in the cage section containing a sand box. The foxes rested much on the cage floor, but they also used the interior of the nest box and elevated locations for resting. Social contact often occurred during resting. Thus, the nest box and elevated location, in conjunction with social contact seem to be valuable while resting. While active, the foxes utilised the cage floor and roof of the nest box instead of the platforms. Scratching, digging and an interaction with the wooden block were seldom observed. Activity occurred mainly on the ‘empty’ cage area. In conclusion, all studied resources provided blue foxes with a distinct value, as they all were used in the complex housing environment. The nest box is used most and for most variable behaviours.     

Den attendance patterns in swift foxes during pup rearing: varying degrees of parental investment within the breeding pair

Parental investment varies in mammalian species, with male care of young being more common in social and monogamous species. Monogamy is commonly observed in canid species, with both males and females, and often “helper” individuals, providing some degree of care for the young. Social units of the swift fox (Vulpes velox), a small North American canid species, usually consist of a male–female pair and occasionally helpers. The role of parental investment and behavior in swift fox society is currently poorly understood. We observed swift fox dens during the pup-rearing season in each of 2 years to evaluate attendance and frequency of visits to natal dens by adult males and females. Female foxes remained at dens longer and visited them more frequently than did male foxes. Female attendance and visitation decreased throughout the pup-rearing season as pups became older and more independent. Environmental factors, including climate and its effect on prey, appeared to contribute to differences in fox behavior between the 2 years. We observed only one fox outside of the breeding pair attending a den in each of the 2 years, both of which were males. We concluded that each of these two foxes were living within the social unit of the male–female pair as a trio, but not serving as a helper and contributing to the care of the pups. Our results increased knowledge of the ecology and behavior of the swift fox, a species of conservation concern in the Great Plains of North America.     

The city-fox phenomenon: genetic consequences of a recent colonization of urban habitat

The red fox (Vulpes vulpes) is one of the best-documented examples of a species that has successfully occupied cities and their suburbs during the last century. The city of Zurich (Switzerland) was colonized by red foxes 15 years ago and the number of recorded individuals has increased steadily since then. Here, we assessed the hypothesis that the fox population within the city of Zurich is isolated from adjacent rural fox populations against the alternative hypothesis that urban habitat acts as a constant sink for rural dispersers. We examined 11 microsatellite loci in 128 foxes from two urban areas, separated by the main river crossing the city, and three adjacent rural areas from the region of Zurich. Mean observed heterozygosity across individuals and the number of detected alleles were lower for foxes collected within the city as compared with their rural conspecifics. Genetic differentiation was significantly lower between rural than between rural and urban populations, and highest value of pairwise FST was recorded between the two urban areas. Our results indicate that the two urban areas were independently founded by a small number of individuals from adjacent rural areas resulting in genetic drift and genetic differentiation between rural and urban fox populations. Population admixture and immigration analysis revealed that urban-rural gene flow was higher than expected from FST statistics. In the five to seven generations since colonization, fox density has dramatically increased. Currently observed levels of migration between urban and rural populations will probably erode genetic differentiation over time.     

From breeding pairs to fox towns: the social organisation of arctic fox populations with stable and fluctuating availability of food

Food availability can impact group formation in Carnivora. Specifically, it has been suggested that temporal variation in food availability may allow a breeding pair to tolerate additional adults in their territory at times when food abundance is high. We investigate group occurrence and intraspecific tolerance during breeding in a socially flexible canid, the arctic fox (Vulpes lagopus). We compare Iceland and Sweden where resource conditions differ considerably. A breeding pair was the most common social unit in both populations, but as predicted, groups were more frequent where food abundance varied substantially between years (Sweden: 6 %) than where food availability was stable (Iceland: ≤2 %). Within Sweden, supplemental feeding increased group occurrence from 6 to 21 %, but there was no effect of natural variation in lemming (Lemmus lemmus) availability since group formation was rare also at lemming highs. Thus, additional factors appeared to influence the trade-off between intraspecific territoriality and tolerance. We report two cases where related females showed enduring social relationships with good-neighbour strategies. Related females also engaged in alloparental behaviour in a ‘fox town’ with 31 foxes (4 adults, 3 litters). In contrast, when unrelated foxes bred close to each other, they moved or split their litters during summer, presumably because of territorial conflict. We suggest that fluctuating food availability is linked to group formation in this Arctic carnivore, but also when food availability increases, additional factors such as relatedness, alloparental benefits, competition and predator defence appear necessary to explain group formation.     

Structure of arctic fox (Alopex lagopus beringensis) colonies in the northern extremity of Bering Island

Data on the spatial structure of an Arctic fox (Alopex lagopus beringensis) colony were obtained in July-August 1995, using walk counts and observations near living dens around the Northern rookery of the northern fur seals located on Bering Island (Commander Islands). The home ranges of 31 Arctic fox families (61 adults and 145 pups inhabiting 66 dens) were found over 27 km of the coastline. Sixty individuals (3 adults and 57 pups) were marked by color ear-tags. Among adult foxes, 24 (39.3%) were recognized as females and 12 (19.7%) as males; the sex of 25 (41.0%) foxes was not recognized. Among 57 marked cubs, 26 (45.6%) were females and 31 (54.4%) were males. The best studied families (13) had 3–11 pups (6.7 ± 0.7, on average). The survival of cubs at an age younger than 2.0–2.5 months was 82.5%; 30.8% of the families consisted of more than two adults. The distribution of the Arctic fox dens and home ranges along the coastline has been studied; specific features of the location of dens have been described. In the studied area, Arctic foxes have been foraging on birds (67.6% of dens with food remains), northern fur seals (40.5), other marine mammals (13.5), Pacific salmon (29.7), and reindeer (2.7%), as well as on amphipods and voles. Rich constant food sources (rookeries, marine bird colonies, and spawning places of the blueback salmon) were found in 7 home ranges of the Arctic fox; 6 home ranges included temporary food sources (spawning streams of the humpback salmon); and 18 home ranges were poor in food resources. Arctic foxes whose home ranges lie within 6–7 km around a “food patch” used the concentrated food resources together. Food resources are supposed to become important only after the raised pups turn to self-feeding. Differences in the use of space, foraging and breeding of the two Arctic fox subspecies (A. l. beringensis and A. l. semenovi), and arrangement of colonies around the northern fur seal rookeries are discussed.     

Food limitation and social regulation in a red fox population

This study evaluates a conceptual model on functional and numerical response to short-term fluctuating vole populations of a red fox ( Vulpes vulpes L.) population in south-central Sweden. The model assumes that this particular population is located in between socially regulated stable populations to the south and direct food-limited populations to the north. The model predicts: (1) food availability as the primary factor for limiting fox numbers, causing reduced rates of reproduction and survival during years of low vole densities, and (2) density-dependent regulation during years of increasing and high vole densities resulting in increased group sizes within territories of fixed dimensions. During 1973–1980 data were obtained from 1216 fox scats, 874 fox carcasses, 63 tagged foxes, nine radio-collared females and from yearly den counts in an area of 130 km², Eight predictions of the model were tested. These concerned the occurrence of small rodents in fox diet, fluctuations in the density of foxes, variations in the number of fox litters, the effect on reproduction of providing supplemental food during January–May, the proportion of vixens bearing a litter different years, dispersal of young males relative to that of young females throughout the vole cycle, and variations in mortality rates of young males and females. All tests were in favour of the conceptual model, and contradictory to alternative models.     

Activity patterns and interactions of red foxes (Vulpes vulpes) in Oxford city

The activity patterns of red foxes ( Vulpes vulpes L.) living and breeding in urban areas of Oxford, and the interactions between individuals, were analysed from detailed radio-tracking data on 17 adults. Foxes were nocturnal and active during the night for a mean of 6h 52min min, irrespective of the time of year. The number of active/resting periods increased in autumn and winter, and in winter and spring the night was characteristically divided into several short cycles of activity of 2–2 1/2 h each, interspersed with similar periods of rest. The foxes occupied mutually exclusive group ranges and group members remained within 50m of each other for up to one-third of the time spent active at night. The implications of fox spatial relationships for rabies transmission are discussed.     

Inter- and intraspecific competition between the fox species Alopex lagopus and Vulpes vulpes: an evaluation trial under penned conditions

This study compared competition capacity and dominance relations between arctic foxes (Alopex lagopus) and red foxes (Vulpes vulpes). Experiments were carried out in semi-natural earthen floor enclosures using farm-bred colour types of both species (blue fox and silver fox) as subjects. Results of the dominance scoring and open field behaviour after weaning in August-September showed that blue foxes dominated over silver foxes. Thereafter, the situation gradually became reversed and silver foxes were dominant during the breeding and whelping seasons. Housing both species together from weaning produced more curious animals as compared to when these species were placed in common quarters after the autumn equinox. In the case of blue foxes, the male dominated highly over all females. In silver foxes, the difference in dominance between the sexes was, however, less pronounced. The most dominant individuals in the study groups were typically among the heaviest. Breedings and whelpings succeeded better in silver than in blue foxes. However, none of litters born survived more than one week. The present results support the conclusion that when both fox species are housed together, Vulpus vulpus tends to dominate over Alopex lagopus. Received: 22 March 1996/Accepted: 30 June 1996     

Food habits of the Indian fox (Vulpes bengalensis) in Kutch, Gujarat, India

We characterized the diet of the Indian fox (Vulpes bengalensis) during the breeding season in a semi-desert region of Western India. Diet was estimated using scat analysis. We used Index of Relative Importance (IRI) to determine the contribution of prey items in the diet of the Indian fox.Indian foxes were observed to feed on a wide variety of prey items. Arthropods were the most frequently occurring prey in their diet. IRI scores were highest for the group Coleoptera and Orthoptera followed by rodents, termites, Ziziphus fruits and spiny tailed lizards (Uromastyx hardwickii). IRI scores for rodents were higher for pups, differing significantly from proportions present in adult diet, thus indicating that they are crucial food items for the young ones. Prey proportions in the fox diet differed between the two habitats in the study area (grassland and scrubland).Our data suggest that the Indian fox is essentially an omnivore showing similar diet (in terms of high incidence of arthropods) to foxes inhabiting arid and semi-arid regions. The opportunistic and generalist strategy has probably helped the species to survive in varied habitats across the Indian subcontinent.     

Non-invasive faecal steroid monitoring of ovarian and adrenal activity in farmed blue fox (Alopex lagopus) females during late pregnancy, parturition and lactation onset

In the semi-domesticated blue fox, handling stress may influence reproductive performance and increase perinatal pup loss. Ovarian and adrenal steroids were analysed in faecal samples collected from mid-gestation through the first week of lactation in 40 female blue foxes to characterize hormone patterns during this important reproductive period. Daily faecal samples were collected from 40 foxes during 30 pregnancies, one late abortion and nine bred-matched non-pregnancies. Mean concentrations of faecal progestagens over the 10 days before birth were significantly higher in pregnant compared to non-pregnant females (51+/-1.50 microg/g versus 36+/-3.72 microg/g, respectively; P < 0.01). From 10 to 3 days before whelping, total faecal oestrogen concentrations also were higher (P < 0.01) in pregnant (1082+/-41.69 ng/g) than non-pregnant (628+/-72.43 ng/g) foxes, before declining to non-pregnant values (402+/-24.88 ng/g) after parturition. Overall mean faecal corticoid concentrations from 3 to 20 days before whelping differed between pregnant and non-pregnant foxes (128+/-3.11 ng/g versus 103+/-5.86 ng/g, respectively; P < 0.01). Furthermore, in pregnant foxes, corticoid excretion increased further from 2 days before to 3 days after whelping (216+/-13.71 ng/g; P < 0.01). Thereafter, corticoid concentrations were similar between pregnant and non-pregnant females (P > 0.05). In sum, the faecal steroid hormone patterns for oestrogens and progestagens were similar to those previously obtained by analyses of fox serum hormones, with both steroids being higher in pregnant than non-pregnant foxes at the end of gestation. The elevation in corticoid concentrations in pregnant females suggests that adrenal activation is involved in the initiation of parturition in the blue fox. Thus, faecal steroid analyses can be used to monitor ovarian activity during pregnancy and pseudopregnancy in farmed blue fox females.     

Fox‐baiting in agricultural landscapes in south‐eastern Australia: a case‐study appraisal and suggestions for improvement

Summary In south‐eastern Australia, the introduced Red Fox (Vulpes vulpes) is a major predator of native wildlife and livestock. Fox control in agricultural landscapes is heavily reliant on the laying of poisoned baits by private landholders, yet there have been few assessments of the application or success of landholder‐baiting practices. We evaluated a community‐based fox‐baiting campaign, typical of programs employed throughout the agricultural regions of south‐eastern Australia to control foxes. We recorded the spatial coverage of 1080 baits deployed by landholders, assessed baiting procedures, monitored the survival of six radio‐collared foxes during and after baiting, and compared the spatial coverage and likely effectiveness of the baiting program with two alternative (theoretical) baiting strategies. Relative to other baiting programs, coordination among neighbours was reasonably high, with 37.5% of baited properties (n = 40) adjoining ≥3 neighbouring properties that also contained baits. Nevertheless, the maximum distance from the centre of a baited property to the nearest edge of an unbaited property was <750 m (mean = 380 m ± 147 m SD). On average, 33% (±17% SD) of each fox’s home range overlapped with baited properties, but only two foxes died during the baiting program. The remaining four foxes were still alive 10 weeks after baiting ceased. Modelling of simulated fox home ranges showed that 13.5% contained no bait stations based on the community baiting program, whereas alternative roadside‐ and grid‐baiting strategies (theoretically) delivered baits to all simulated home ranges. Some landholders employed practices that could reduce the effectiveness of baiting programs such as not removing decayed baits before deploying new ones or placing bait stations too close together. Our research illustrates the difficulties of managing a coordinated baiting program on private land that effectively controls foxes. Alternative baiting strategies such as roadside baiting need to be considered to improve fox control in agricultural landscapes.     

Exclusion by interference competition? The relationship between red and arctic foxes

The distribution of many predators may be limited by interactions with larger predator species. The arctic fox in mainland Europe is endangered, while the red fox is increasing its range in the north. It has been suggested that the southern distribution limit of the arctic fox is determined by interspecific competition with the red fox. This has been criticised, on the basis that the species co-exist on a regional scale. However, if the larger red fox is superior and interspecific competition important, the arctic fox should avoid close contact, especially during the breeding season. Consequently, the distribution of breeding dens for the two species would be segregated on a much smaller spatial and temporal scale, in areas where they are sympatric. We tested this hypothesis by analysing den use of reproducing arctic and red foxes over 9 years in Sweden. High quality dens were inhabited by reproducing arctic foxes more often when no red foxes bred in the vicinity. Furthermore, in two out of three cases when arctic foxes did reproduce near red foxes, juveniles were killed by red foxes. We also found that breeding arctic foxes occupied dens at higher altitudes than red foxes did. In a large-scale field experiment, red foxes were removed, but the results were not conclusive. However, we conclude that on the scale of individual territories, arctic foxes avoid areas with red foxes. Through interspecific interference competition, the red fox might thus be excluding the arctic fox from breeding in low altitude habitat, which is most important in years when food abundance is limited and competition is most fierce. With high altitude refuges being less suitable, even small-scale behavioural effects could scale up to significant effects at the population level.     

Habitat use and ranging behaviour of the red fox (Vulpes vulpes) in a Mediterranean rural area: is shelter availability a key factor?

Factors affecting activity, habitat use, and home-range size of the red fox were analysed in a highly heterogeneous rural environment. Individual differences in behaviour were used to test our hypotheses. Food habits tended to depend on food availability, which, in turn, was mainly influenced by temperature. Diet was highly heterogeneous. Insects, e.g. grasshoppers and beetles, and cultivated fruits were the staple of the diet, but no diet component stood out clearly from all others. For an opportunistic species such as the fox, habitat heterogeneity may be the main factor underlying a wide trophic niche. All foxes selected the vineyard as part of their home ranges, whereas they preferred for activity the abandoned olive-yard among the habitats of their home ranges. Human intolerance of foxes affects their pattern of activity, habitat selection, and ranging behaviour. Foxes were strongly nocturnal. Cover-rich habitats were preferred for resting and for movements in daylight. Areas under human management were mainly used at night. Selectivity was higher for resting than for activity sites. Variation in home-range size and shape can be influenced not only by the dispersion of the main food patches, but also by the location of shelters     

Innovative parental care in a myrmecophageous mammal

Male bat-eared foxes, Otocyon megalotis, are known to contribute extensively to parental care. Yet, the exact roles that males and females play in raising offspring remain relatively unexplored. Here, we describe interactions between adult foxes and their presumed offspring based on a pilot study on three family groups of a wild population in South Africa. We report the first recorded instance of dung provisioning observed in canids. A male bat-eared fox provided dung to his offspring during a foraging trip, presumably to give them access to the ensconced insects. Further, this male provisioned the young foxes with large, live insects. Similar to other researchers, we never observed provisioning by females, but the females in this population did interact socially with their young in addition to suckling. We emphasize the importance of anecdotal reports of novel behavioural responses in wild canids, as an accumulation of such evidence may reveal patterns of innovative behaviour presently unrecognized in this family.     

The flea epifauna of a suburban fox (Vulpes vulpes) population

Five hundred and thirteen fleas, of eight different species, were collected from a sample of 252 foxes killed in suburban London. 25–8% of foxes carried fleas, with a mean of 204 fleas per fox. Levels of infestation of male and female hosts did not differ significantly. Possible sources of the fleas infesting foxes are discussed with respect to their seasonal occurrence and fox prey composition. No evidence was found to support the suggestion that foxes obtain the majority of their fleas from prey items, although occasional heavy infestations of some flea species were probably derived from recent meals. Although Pulex irritans, Paraceras m. melis and Ctenocephalides canis , which contributed 35 % of the flea epifauna, could be considered parasitic on the fox, it seems probable that foxes pick up the majority of their fleas from the habitat through which they move. Thus, two particularly heavily infested categories of foxes were found: (1) juveniles during July-September, their fleas probably being accumulated during exploratory and play activities, and (2) all animals during the period October-December.     

On the origin of a domesticated species: Identifying the parent population of Russian silver foxes (Vulpes vulpes)

The foxes at Novosibirsk, Russia, are the only population of domesticated foxes in the world. These domesticated foxes originated from farm-bred silver foxes (Vulpes vulpes), whose genetic source is unknown. In this study we examined the origin of the domesticated strain of foxes and two other farm-bred fox populations (aggressive and unselected) maintained in Novosibirsk. To identify the phylogenetic origin of these populations we sequenced two regions of mtDNA, cytochrome b and D-loop, from 24 Novosibirsk foxes (8 foxes from each population) and compared them with corresponding sequences of native red foxes from Europe, Asia, Alaska and Western Canada, Eastern Canada, and the Western Mountains of the USA. We identified seven cytochrome b - D-loop haplotypes in Novosibirsk populations, four of which were previously observed in Eastern North America. The three remaining haplotypes differed by one or two base change from the most common haplotype in Eastern Canada. Φ(ST) analysis showed significant differentiation between Novosibirsk populations and red fox populations from all geographic regions except Eastern Canada. No haplotypes of Eurasian origin were identified in the Novosibirsk populations. These results are consistent with historical records indicating that the original breeding stock of farm-bred foxes originated from Prince Edward Island, Canada. Mitochondrial DNA data together with historical records indicate two stages in the selection of domesticated foxes: the first includes captive breeding for ~50 years with unconscious selection for behaviour; the second corresponds to over 50 further years of intensive selection for tame behaviour.     

Genetic polymorphism of plasma α1B-glycoprotein and transferrin in arctic and silver foxes

Plasma samples of 235 foxes from 38 complete families (14 of arctic foxes, 21 of silver foxes and 3 with arctic x silver fox hybrid offspring) were analysed by one-dimensional horizontal polyacrylamide gel electrophoresis (PAGE) pH 9.0 followed by general-protein staining of gels. A major postalbumin of fox plasma was identified as alpha 1B-glycoprotein (alpha 1B) by using immunoblotting with antiser m specific to human or pig plasma alpha 1B. Four codominant, autosomal alleles of alpha 1B were found in arctic foxes. Two transferrin (TF) alleles (TfF, TfS) were observed in arctic foxes and two (TfD, Tff) in silver foxes; the TF F type of both of the fox species showed identical electrophoretic mobilities. The arctic foxes showed a high degree of polymorphism for both TF and alpha 1B. The silver foxes showed a scarce polymorphism of TF and were monomorphic for alpha 1B. The arctic fox, silver fox and their hybrids could be clearly differentiated from one another by their plasma protein patterns obtained by the PAGE method.     

Prevalence of Toxoplasma gondii infection diagnosed by PCR in farmed red foxes, arctic foxes and raccoon dogs

The aim of this study was to compare Toxoplasma gondii infection in three canid species: red fox Vulpes vulpes, arctic fox Vulpes lagopus and raccoon dog Nyctereutesprocyonoides kept at the same farm. Anal swabs were taken from 24 adult and 10 juvenile red foxes, 12 adult arctic foxes, three adult and seven juvenile raccoon dogs. Additionally, muscle samples were taken from 10 juvenile red foxes. PCR was used to detect T. gondii DNA. T. gondii infection was not detected in any of the arctic foxes; 60% ofraccoon dogs were infected; the prevalence of the parasite in material from red fox swabs was intermediate between the prevalence observed in arctic foxes and raccoon dogs. It is possible that susceptibility and immune response to the parasite differ between the three investigated canid species. T. gondii DNA was detected in muscle tissue of five young foxes. The results of this study suggest that T. gondii infection is not rare in farmed canids.     

Estimating number of families for an urban fox population by using two public data sets

The number of families in the urban fox population of Sapporo, Japan, was estimated from two sets of data reported by the public to government: records of road-killed foxes (information-A) and records of complaints about foxes (information-B). We assumed that fox populations consist of families that have exclusive home ranges, i.e., territories, during the period between gestation and dispersal. The urban area was then divided into hexagons that correspond to the territories. The locations from the two sets of records during the territorial period were plotted on the map. The number of fox families for which information-A and/or B was reported was estimated by counting the number of hexagons that include the record. The total number of families was estimated by using a double-observation method. We adopted Chapman’s unbiased estimator which is based on the hypergeometric distribution that corresponds to the conditional likelihood. We demonstrated the possibility of estimating the abundance of animals from government data such as road kill and complaints if the animals have territories. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.