Evolutionary implications of morphological variation in the lower carnassial of red fox<Emphasis Type="Italic">Vulpes vulpes</Emphasis>

Research on the morphological variability of the occlusal surface of M₁ talonid in the red foxVulpes vulpes (Linnaeus, 1758) in the Holarctic has been carried out on 2271 specimens originating from 42 populations. The Nearctic was represented by 666 specimens belonging to 13 populations, whereas Palearctic was represented by 1605 specimens from 29 populations. Analyses of the developmental level and formation of cristids between the hypoconid and entoconid allowed the differentiation of 34 shape variants of the occlusal surface of the talonid in the red fox. Because of the complicated variation of cristids, 34 variants were assigned to 5 morphotypes of group P. In the Palearctic and Nearctic a significant geographic variation occurred of P morphotypes and their variants. Primitive variants of the talonid structure on M₁ are predominant in populations from the south of the Asian range of the red fox, while more progressive characters of the occlusal surface of the lower carnassial are typical of the northern and centrally located red fox populations in the Palearctic and Nearctic. The geographic differentiation is probably connected with different Pleistocene histories of particular populations.     

Geography of sexual dimorphism in the tooth size of the red fox Vulpes vulpes (Mammalia, Carnivora)

Geographic variation in sexual dimorphism of tooth size was assessed for the red fox Vulpes vulpes (Linnaeus, 1758) across the whole northern range of the species. Twenty-one measurements of tooth size and skull length were taken from 2849 specimens (1577 males and 1272 females) originating from 12 Nearctic and 25 Palearctic localities. The index of sexual dimorphism was calculated as a quotient of the mean measure of certain characters in males by the respective mean in females ( M _m/ M _f). In the whole range, the males were larger than females and mean dimorphism index of tooth size ranged from 1.01 to 1.06. On average, the tooth measurements in males were 3.6% larger than in females. The highest dimorphism was observed in the canines. Dimorphism of tooth size was higher in the Palearctic than Nearctic. Statistically significant differences between regions were found for lengths of C¹, C₁ and M₁. In the Palearctic, higher values of the dimorphism indices were observed particularly in the southern parts of the Eurasian range of the red fox and in Great Britain. For a few metrical traits, sexual dimorphism indices presented significant relations to some geo-climatic variables. The geographic pattern of size dimorphism in the red fox seems to be shaped by sexual selection, intraspecific and interspecific competition and population density.     

Microsatellite differentiation between Phillip Island and mainland Australian populations of the red fox Vulpes vulpes

Predation by the red fox Vulpes vulpes is believed to be threatening the little penguin Eudyptula minor on Phillip Island in Victoria. Polymorphism at seven microsatellite loci was examined to estimate the extent of differentiation between Phillip Island and mainland populations of V. vulpes. Loss of alleles has occurred on Phillip Island where foxes first appeared = 88 years ago compared with mainland populations. Genetic differentiation between the Phillip Island and mainland populations was high. The relatively high differentiation found between the two populations could be due to either low migration rates, the effect of the composition of founder animals or both effects. Further ecological and historical information about the populations is needed to explore the likely significance of these effects.     

Geography of dental polymorphism in the red fox Vulpes vulpes and its evolutionary implications

Frequency distributions of dental morphotype characters (groups of morphotypes from A to S) in 43 red fox Vulpes vulpes (Linnaeus, 1758) populations from the Holarctic region were analysed. Definite but different geographical gradients in morphotype dental patterns were found both in the Palearctic and Nearctic. In the Nearctic, a great number of characters present geographical gradient related to longitude whereas, in the Palearctic, gradient relates to latitude. Mean annual temperature and mean annual sum of precipitations play a significant role in shaping the geographical variation of morphotype characters. The shape of the teeth in the centre of the morphogenetic field is more heavily geo-climatically conditioned than the morphotype characters in the teeth at the ends of the cheek teeth field. There was an attempt to reconstruct the evolutionary changes in the morphotype dental pattern in the V. vulpes line. The morphotype pattern in the red fox dentition partially follows the phylogeographical evolutionary trends in Canidae in the northern hemisphere. However, the picture of morphotype variation is influenced by both earlier episodes of human interference (reintroductions of foxes in North America) and the recent phenomenon of heavy opportunism of the species.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 61–84.     

The arctic fox Alopex lagopus in Fennoscandia: a victim of human-induced changes in interspecific competition and predation?

After a marked decline at the beginning of the 1900s, the arctic fox Alopex lagopus population in Fennoscandia has remained at a very low level. We suggest that the main cause for the population crash was winter starvation caused by (1) over-hunting of reindeer Rangifer tarandus populations, and thus reduced carcass availability in the mountains, and (2) increased interspecific competition for these carcasses because of increased invasion of red foxes Vulpes vulpes from lower altitudes. The failure of arctic fox populations to recover, despite increasing reindeer populations in the mid 1900s, can be explained by a concurrent strong increase in red fox numbers. Analyses of countywide hunting statistics from Norway 1891–1920 suggest that there actually was an increase in red fox numbers in the period of arctic fox decline, and that the increase in reindeer populations from the 1920s to the 1950s was accompanied by a new increase in red fox numbers. We conclude that restoring arctic fox populations most likely will require a substantial and lasting reduction of red fox populations.     

Survival and Cause-Specific Mortality of Corsac and Red Foxes in Mongolia

ABSTRACT The corsac fox (Vulpes corsac) and red fox (Vulpes vulpes) range widely across northern and central Asia and may be declining in many regions due to overhunting and other causes. However, details of the fundamental causes of survival and mortality of both species remain largely unquantified, but may be crucial for understanding interspecific relationships and developing effective conservation actions. We studied a radiomarked population of sympatric corsac and red foxes in central Mongolia to quantify survival and cause-specific mortality rates from April 2005 to April 2007. Survival probability was 0.34 for corsacs (n = 18) and 0.46 for red foxes (n = 17) and did not vary by year within or between each species. Among both foxes, mortality occurred mainly from hunting by humans, but also from predation by larger canids and unknown causes. Our results suggest that illegal human hunting represents the principal source of mortality for both species and that a recently initiated ranger patrol program in the study area did not affect fox survival. As such, more stringent protective measures will likely be necessary to halt declines of both foxes. Our results also suggest that interference competition occurs between species as red foxes killed but did not consume corsacs. Our results will be useful for developing science-based management strategies to protect foxes in Mongolia, and in understanding the competitive relationships between them.     

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.     

Range‐wide multilocus phylogeography of the red fox reveals ancient continental divergence,minimal genomic exchange and distinct demographic histories

Widely distributed taxa provide an opportunity to compare biogeographic responses to climatic fluctuations on multiple continents and to investigate speciation. We conducted the most geographically and genomically comprehensive study to date of the red fox (Vulpes vulpes), the world's most widely distributed wild terrestrial carnivore. Analyses of 697 bp of mitochondrial sequence in ~1000 individuals suggested an ancient Middle Eastern origin for all extant red foxes and a 400 kya (SD = 139 kya) origin of the primary North American (Nearctic) clade. Demographic analyses indicated a major expansion in Eurasia during the last glaciation (~50 kya), coinciding with a previously described secondary transfer of a single matriline (Holarctic) to North America. In contrast, North American matrilines (including the transferred portion of Holarctic clade) exhibited no signatures of expansion until the end of the Pleistocene (~12 kya). Analyses of 11 autosomal loci from a subset of foxes supported the colonization time frame suggested by mtDNA (and the fossil record) but, in contrast, reflected no detectable secondary transfer, resulting in the most fundamental genomic division of red foxes at the Bering Strait. Endemic continental Y‐chromosome clades further supported this pattern. Thus, intercontinental genomic exchange was overall very limited, consistent with long‐term reproductive isolation since the initial colonization of North America. Based on continental divergence times in other carnivoran species pairs, our findings support a model of peripatric speciation and are consistent with the previous classification of the North American red fox as a distinct species, V. fulva.     

<Emphasis Type="Italic">Echinococcus multilocularis</Emphasis> in the red fox (<Emphasis Type="Italic">Vulpes vulpes</Emphasis>) in Slovenia

Using the parasitological washing out method, we examined the intestines of 428 red foxes (Vulpes vulpes) for the presence of Echinococcus multilocularis (Leuckart 1863) and found that the overall prevalence was 2.6% (confidence interval 95% 1.3–4.5%). This is the first extended research reporting on the presence of E. multilocularis in the Slovenian fox population.     

Predation by red foxes limits recruitment in populations of eastern grey kangaroos

We investigated the impact of red fox (Vulpes vulpes Linnaeus 1758) predation on juvenile eastern grey kangaroos (Macropus giganteus Shaw 1790) using a replicated predator removal experiment. In two sites in Namadgi National Park, south‐eastern Australia, a persistent 1080 poisoning campaign over 18 months reduced fox density by more than 85%, and to less than 10% of the fox density in two other sites with no fox baiting. Changes in the mother : young ratios and densities of kangaroo populations were monitored twice monthly along 2‐km transects in each site from July 1993 to February 1995. Compared to nonremoval sites, where foxes were controlled, 25–40% more females retained juveniles over the period when these young became emergent from the pouch. This higher survival of emergent pouch young resulted in a significantly higher proportion of juveniles in kangaroo populations at fox control sites, which resulted in a significantly higher annual growth rate. We conclude that predation upon juveniles is an important limiting factor for kangaroo populations in Namadgi NP.     

Natal den selection by sympatric arctic and red foxes on Herschel Island,Yukon, Canada

In the twentieth century, red fox (Vulpes vulpes) expanded into the Canadian Arctic, where it competes with arctic fox (Vulpes lagopus) for food and shelter. Red fox dominates in physical interactions with the smaller arctic fox, but little is known about competition between them on the tundra. On Hershel Island, north Yukon, where these foxes are sympatric, we focused on natal den choice, a critical aspect of habitat selection. We tested the hypothesis that red fox displaces arctic fox from dens in prey-rich habitats. We applied an approach based on model comparisons to analyse a 10-year data set and identify factors important to den selection. Red fox selected dens in habitats that were more prey-rich in spring. When red foxes reproduced, arctic fox selected dens with good springtime access, notably many burrows unblocked by ice and snow. These provided the best refuge early in the reproductive season. In the absence of red foxes, arctic foxes selected dens offering good shelter (i.e. large isolated dens). Proximity to prey-rich habitats was consistently less important than the physical aspects of dens for arctic fox. Our study shows for the first time that red foxes in the tundra select dens associated primarily with prey-rich areas, while sympatric arctic foxes do not. These results fit a model of red fox competitively interfering with arctic fox, the first detailed study of such competition in a true arctic setting.     

Mercury in wild terrestrial carnivorous mammals from north-western Poland and unusual fish diet of red fox

Total mercury concentrations were determined in the kidney (K), liver (L), and pectoral muscle (M) of 19 individuals representing wild carnivorous mammals from NW Poland: 10 red foxes Vulpes vulpes (Linnaeus, 1758), 3 raccoon dogs Nyctereutes procyonoides Gray, 1834, 2 badgers Meles meles Linnaeus, 1758, 3 pine martens Martes martes Linnaeus, 1758, and 1 polecat Mustela putorius Linnaeus, 1758. The sample of red fox included 3 immature specimens found on Mielin Island; the island supports a black cormorant colony, and the foxes found there had fed mostly on cormorant nestlings as well as on fish and their remains. In addition to the Mielin Island foxes, the group of foxes included 3 other immature and 4 adult individuals. The highest mean of mercury concentrations were revealed in the Mielin red fox juveniles: 5.11, 4.52, and 1.56 mg/kg d.w. being recorded in K, L, and M. No significant differences in mercury concentrations in the respective tissues were found between the remaining immature and adult red foxes; their mercury concentrations were several times lower than those of the Mielin individuals. In all the animals except the Mielin foxes, mercury concentrations in K, L, and M did not exceed 1.3, 1.0 and 0.5 mg/kg d.w., respectively, the highest values being in badgers (which feed mostly on soil invertebrates), followed by pine martens and then the canids (red fox and raccoon dog). Studies on common and widely distributed terrestrial animals, particularly red fox and badger, may provide numerous valuable comparative data on mercury contamination of different areas of the northern hemisphere.     

The effects of sex,age, season and habitat on diet of the red fox <Emphasis Type="Italic">Vulpes vulpes</Emphasis> in northeastern Poland

The diet of the red fox Vulpes vulpes was investigated in five regions of northeastern Poland by stomach content analysis of 224 foxes collected from hunters. The red fox is expected to show the opportunistic feeding habits. Our study showed that foxes preyed mainly on wild prey, with strong domination of Microtus rodents, regardless of sex, age, month and habitat. Voles Microtus spp. were found in 73% of stomachs and constituted 47% of food volume consumed. Other food items were ungulate carrion (27% of volume), other mammals (11%), birds (9%), and plant material (4%). Sex- and age-specific differences in dietary diversity were found. Adult males and juvenile foxes had larger food niche breadths than adult females and their diets highly overlapped. Proportion of Microtus voles increased from autumn to late winter. Significant habitat differences between studied regions were found. There was a tendency among foxes to decrease consumption of voles with increasing percentage of forest cover. Based on our findings, red foxes in northeastern Poland can be recognized as a generalist predators, consuming easily accessible and abundant prey. However, high percentage of voles consumed regardless of age, sex, month, or habitats may indicate red fox specialization in preying on Microtus rodents.     

Feeding responses of the red fox (<Emphasis Type="Italic">Vulpes vulpes</Emphasis>) to different wild rabbit (<Emphasis Type="Italic">Oryctolagus cuniculus</Emphasis>) densities: a regional approach

We investigate the feeding responses of the red fox (Vulpes vulpes) at a regional scale to different densities of European wild rabbit (Oryctolagus cuniculus) in central–southern Spain. Rabbit abundance indices were obtained in 86 localities during summer 2002. The diet of the fox was studied by analysis of 114 scats collected in 47 of these localities. The feeding response of the fox was examined by a representation of the dry weight percent of rabbit in the diet as a function of the abundance of rabbits; this used data only from those localities where at least 3 scats were collected (70 fox scats from 18 localities). We evaluated the relationship between rabbit abundance and the diversity of the diet of the fox. The feeding patterns of red foxes approximated to Holling’s type III functional response, typical of opportunistic predators. There was a negative relationship between the diversity of the fox’s diet and the abundance of rabbits. Therefore, the fox apparently behaves as a facultative predator, feeding on rabbits when they are abundant and shifting to other prey (and hence a more diverse diet) when rabbits are scarce. These findings are the first step towards understanding the potential role of red foxes in regulating rabbit populations in central–southern Spain.     

Population structure of two rabies hosts relative to the known distribution of rabies virus variants in Alaska

For pathogens that infect multiple species, the distinction between reservoir hosts and spillover hosts is often difficult. In Alaska, three variants of the arctic rabies virus exist with distinct spatial distributions. We tested the hypothesis that rabies virus variant distribution corresponds to the population structure of the primary rabies hosts in Alaska, arctic foxes (Vulpes lagopus) and red foxes (Vulpes vulpes) to possibly distinguish reservoir and spillover hosts. We used mitochondrial DNA (mtDNA) sequence and nine microsatellites to assess population structure in those two species. mtDNA structure did not correspond to rabies virus variant structure in either species. Microsatellite analyses gave varying results. Bayesian clustering found two groups of arctic foxes in the coastal tundra region, but for red foxes it identified tundra and boreal types. Spatial Bayesian clustering and spatial principal components analysis identified 3 and 4 groups of arctic foxes, respectively, closely matching the distribution of rabies virus variants in the state. Red foxes, conversely, showed eight clusters comprising two regions (boreal and tundra) with much admixture. These results run contrary to previous beliefs that arctic fox show no fine‐scale spatial population structure. While we cannot rule out that the red fox is part of the maintenance host community for rabies in Alaska, the distribution of virus variants appears to be driven primarily by the arctic fox. Therefore, we show that host population genetics can be utilized to distinguish between maintenance and spillover hosts when used in conjunction with other approaches.     

High genetic diversity and low population structure in red foxes (Vulpes vulpes) from Croatia

The red fox (Vulpes vulpes) is a highly adaptable omnivorous mammal distributed across all continents on the northern hemisphere. Although the red fox is present throughout Europe, where it plays an important ecological and socio-economic role not only as a game species but also as a rabies reservoir, few studies have examined its population-level mitochondrial DNA variability. In this study, 27 mitochondrial DNA control region haplotypes were identified in 229 red fox samples taken from four regions in Croatia. Haplotype diversity of Croatian red foxes (0.901) was found to be among the highest of all European red fox populations studied to date. Genetic differentiation among regions was quite low, and statistically significant estimates of differentiation were obtained only when comparing the population from the peninsular region of Istria with the three continental populations. It seems that landscape barriers like rivers and small mountains do not restrict gene flow among foxes in the continental part of Croatia, while the combination of a narrow land bridge and altitudes exceeding 1000 m limit fox migration between Istria and the rest of the continent. Better understanding of small-scale population structure will require analysis of highly variable nuclear markers like microsatellites.     

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     

Arctic fox versus red fox in the warming Arctic: four decades of den surveys in north Yukon

During the last century, the red fox (Vulpes vulpes) has expanded its distribution into the Arctic, where it competes with the arctic fox (Vulpes lagopus), an ecologically similar tundra predator. The red fox expansion correlates with climate warming, and the ultimate determinant of the outcome of the competition between the two species is hypothesized to be climate. We conducted aerial and ground fox den surveys in the northern Yukon (Herschel Island and the coastal mainland) to investigate the relative abundance of red and arctic foxes over the last four decades. This region has undergone the most intense warming observed in North America, and we hypothesized that this climate change led to increasing dominance of red fox over arctic fox. Results of recent surveys fall within the range of previous ones, indicating little change in the relative abundance of the two species. North Yukon fox dens are mostly occupied by arctic fox, with active red fox dens occurring sympatrically. While vegetation changes have been reported, there is no indication that secondary productivity and food abundance for foxes have increased. Our study shows that in the western Arctic of North America, where climate warming was intense, the competitive balance between red and arctic foxes changed little in 40?years. Our results challenge the hypotheses linking climate to red fox expansion, and we discuss how climate warming’s negative effects on predators may be overriding positive effects of milder temperatures and longer growing seasons.