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.     

Body size of the red fox Vulpes vulpes in Spain: the effect of agriculture

The body size of animals is affected by several factors, including ambient temperature and food availability. Ambient temperature is often negatively related to body size (Bergmann's rule) whereas an improved diet, especially during growth, has a positive effect. Animals commensal with man commonly exploit additional food sources (e.g. garbage dumps), thereby increasing their food supply. Using museum material, we studied morphological variation in skull size (and thus body size) among Spanish red foxes. Four measurements were taken of each skull and were related to the habitat from which the foxes were collected (agricultural and non-agricultural), and to latitude as a proxy for ambient temperature. The skull size of foxes collected in agricultural areas during the late 20th Century was significantly larger than that of those from non-agricultural areas, and was negatively related to latitude, thus contradicting Bergmann's rule. We suggest that increased food availability from animal husbandry is the cause for the observed increase in skull size (and thus body size).  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 729–734.     

Variation in the diet composition of a generalist predator, the red fox, in relation to season and density of main prey

Diet composition of a generalist predator, the red fox (Vulpes vulpes) in relation to season (winter or summer) and abundance of multi-annually cyclic voles was studied in western Finland from 1983 to 1995. The proportion of scats (PS; a total of 58 scats) including each food category was calculated for each prey group. Microtus voles (the field vole M. agrestis and the sibling vole M. rossiaemeridionalis) were the main prey group of foxes (PS = 0.55) and they frequently occurred in the scats both in the winter and summer (PSs 0.50 and 0.62, respectively). There was a positive correlation between the PSs of Microtus voles in the winter diet of foxes and the density indices of these voles in the previous autumn. Other microtine rodents (the bank vole Clethrionomys glareolus, the water vole Arvicola terrestris and the muskrat Ondatra zibethicus) were consumed more in winter than in summer. The unusually high small mustelid predation by red foxes (PS = approx. 0.10) in our study area gives qualitative support for the hypothesis on the limiting impact of mammalian predators on least weasel and stoat populations. None of the important prey groups was preyed upon more at low than at high densities of main prey (Microtus voles). This is consistent with the notion that red foxes are generalist predators that tend to opportunistically subsist on many prey groups. Among these prey groups, particularly hares and birds (including grouse), were frequently used as food by foxes.     

Foraging and diet of the red fox Vulpes vulpes in relation to variable food resources in Biatowieza National Park, Poland

Feeding ecology of red fox Vulpes vulpes was studied by scat analysis and snow-tracking m primeval temperate forest and adjacent meadows during four years (1985/86-1988/89) Winters varied from mild to unusually severe Main food resources for foxes were rodents of open meadows and river valleys (root vole Microtus oeconomus ). forest rodents (bank vole Clethrionomys glareolus and yellow-necked mouse Apodemus flavicollis ), hare Lepus europaetis and carcasses of wild boar Sus scrofa and red deer Cervus elaphus either killed by wolves and lynx or that had died from inanition Composition of fox diet m four cold seasons (autumn-winter) was compared to the abundance of main food resources Prolonged, sharp decline of Microtus was followed by only a twofold decrease of its share in fox diet Foxes continued to prey on declining Microlus The changes in the proportions of forest rodents and hare in fox diet clearly followed the fluctuations in numbers of these two prey Carcasses were alternative, buffer food to foxes and were taken considerably when Microlus and other prey were in low numbers or poorly accessible The depth of snow was the most important factor restricting foxes access to rodents Snow-tracking revealed that foxes dwelling in the forest widely used adjacent open areas In open meadows foxes mainly hunted for rodents, while in the forest the most significant foraging activity was scavenging Seasonal analysis of fox diet revealed that consumption of Microlus by foxes was stable throughout the year (37-47% of biomass consumed) Bank vole significantly contributed to fox diet in autumn, and hare in summer only Scavenging was most pronounced in winter and spring when carcasses made up 30% of biomass taken     

Reproduction of the bank vole, Clethrionomys glareolus, in northern and southern Sweden during several seasons and in different phases of the vole population cycle

The breeding biology of the bank vole was studied in northern and southern populations in Sweden in the years 1973–76. Trapped specimens supplied organ weights and histological data, which were utilized to describe the breeding season, number and size of litters, reproductive losses, spermatogenesis, and sexual maturation. In the north the populations are cyclic. During the increase phase of the cycle the breeding season was long, litter sizes and gonad weights were large, and there was maturation of yearlings in the summer. During the peak phase reproduction was impaired in all these respects. In southern Sweden there was no cyclicity, and the reproductive events varied less, but in 1976 the litters were larger than usual and the season was much extended.
In the south the voles born during the early part of the season regularly became sexually mature during their first summer. The differences in reproductive biology are related to population dynamics and discussed in terms of different hypotheses concerning vole population cyclicity.     

The effect of summer feeding on juvenile arctic fox survival - a field experiment

The arctic fox Alopex lagopus L population in Sweden is small and its numbers fluctuate widely with food availability, l e rodent populations This fluctuation is mediated through differences in recruitment rates between years The recruitment can be divided into three phases number of litters born, number of cubs per litter and cub survival rates The number of litters and their sizes have been shown to depend on food availability during winter and spring To examine cub survival during the summer and how it relates to food availability, we conducted a feeding experiment m northern Sweden during 1990, a year of low rodent density, involving six occupied arctic fox dens Feeding at dens lowered cub mortality rates However, condition and growth rates of juveniles were not influenced by supplementary feeding at dens, nor were they related to the probability of survival for an individual Thus arctic foxes seem to minimize risks rather than maximize growth The juvenile mortality from weaning and over the next 6 wk was 21%, mostly due to starvation Only 82% survived from weaning to the first breeding season Of the one-year-old foxes, 50% survived their second year Supplementary feeding of juveniles had no effect on the final survival rates over these two years However, the immediate, positive effect on cub survival could be used in a long-term, extensive management programme if combined with winter feeding     

The role of vole populations in prevalence of the parasite (Echinococcus multilocularis) in foxes

Effects of population fluctuation of the gray-sided vole(Clethrionomys rufocanus) on the prevalence (infection rates) of the parasiteEchinococcus multilocularis in red fox(Vulpes vulpes) populations was investigated from 1985 to 1992 in eastern Hokkaido (Abashiri, Nemuro, and Kushiro area), Japan. This parasite needs two hosts to complete its life cycle; the gray-sided vole as its intermediate host and the red fox as its final host. We found that: (1) Infection rates in foxes depended on the current-year abundance of voles in all three study areas, particularly in Abashiri. (2) In addition to this direct density-dependence, delayed density-dependence between the infection rate and the prior-year abundance of voles was detected in Nemuro and in Kushiro. (3) The regional differences in density-dependence pattern were related to regional differences in the winter food habits of red foxes: in Abashiri the proportion of voles in the fox’s diet greatly decreases in winter, while the proportion remains high in winter in Nemuro and in Kushiro, probably because of shallower snowpack. These results suggest that infection rates in foxes in Abashiri were less influenced by the prior-year prevalence, since the infection cycle might be interrupted in winter, when voles became less important in fox’s diet. In contrast, the state of the prevalence may carry over from year to year in Nemuro and in Kushiro, because red foxes continue to eat a considerable amount of voles throughout year. The regionally contrasted results for the relationship between infection rate in foxes and vole abundance were parallel to the regional difference in fluctuation pattern of vole populations, which are highly variable in Abashiri area, but less variable in Kushiro-Nemuro area. Drastic change in vole populations appears to affect the host-parasite system.     

Variation in the feeding pattern of red foxes in relation to changes in anthropogenic resource availability in a rural habitat of western Poland

The feeding pattern of red foxes Vulpes vulpes, i.e., the composition of their diet and dietary responses to fluctuations in the abundance of common voles Microtus arvalis were compared in the periods before and after the limitation of anthropogenic food availability in a rural area in western Poland. During the winter seasons of 1996/97–2014/15, the diet of the foxes was estimated by analysing the stomach contents of individuals shot from November to March and the index of vole abundance was obtained by counting their burrow entrances. The major component of the fox diet over the entire study period was anthropogenic food (mainly, the remains of farm animals) that showed both the highest average proportion in the stomach content volume (53.4%) and the highest average frequency of occurrence in the stomachs (62.7%), whereas small mammals were the second important component (27.5% and 54.2%, respectively). The frequency of anthropogenic food decreased and the frequencies of wild ungulate carrion and other wild vertebrates (mainly birds) increased after the limitation of anthropogenic resources. The observed fluctuations in the index of vole abundance resulted in positive changes in the small mammal frequency and negative changes in the anthropogenic food frequency, but the course of these relationships did not significantly differ between the two periods. The frequency of hares in fox stomachs did not change significantly between these periods, but recently the occurrence of hares in the diet of foxes increased with vole abundance. This study indicated that the reduction of anthropogenic food availability for carnivorous mammals may result in their switching to other prey species and potentially may have a negative influence on the demography of such prey species.     

Condition and growth and Red Foxes (Vulpes vulpes) in relation to food supply

Fat deposits of463 Red fox vixens shot during autumn-winter were monitored throughout a four-year cycle of vole abundance (1975-79) in south-central Sweden. The growth ofjuveniles during the same period was indexed by lower jaw length during winter. The reproduction of foxes in the study area was correlated with vole abundance. Fat deposits built up during autumn and were depleted during winter. Subadults had consistently smaller fat deposits than adults. There was no positive correlation between vole abundance and mean fat deposits in autumn or degree of depletion during winter. The mean amount of fat deposited during autumn was correlated with the frequency ofoccurrence offruit and berries in fox stomachs. The degree of depletion of fat deposits during winter was correlated with mean snow depth. It is hypothesized that the availability of fruit and berries (a major source of carbohydrates in the diet of foxes) determines the amount of fat reserves built up during autumn, whereas the energy requirement of moving in deep snow determines the degree of depletion during winter. Juvenile growth was correlated with vole abundance but growth later in life seemed to compensate for this variation.     

Harsh climate selects for small body size among Iceland's Arctic foxes

We studied the effect of the two environmental indices, the sub‐polar gyre (SPG), and winter and summer North Atlantic Oscillation (NAO), together with mean annual winter and summer temperatures and geographic location on mandible size and body mass of Arctic foxes in Iceland (6345 and 2732 specimens, respectively) during the year of their death. We predicted that when favorable conditions prevailed, large specimens would be selected for, and vice versa. Body size and body mass were significantly affected by the environmental parameters (i.e. SPG, NAO, ambient temperature and cloud cover) prevailing during the year of death. The effect of environmental conditions on body size was much stronger in the less productive region of eastern Iceland, apparently because in areas where food availability is meager, even a small difference in climate may tilt the balance from food sufficiency to food shortage. Western Iceland comprises only a quarter of the total surface area of the country, but its productive seashores are twice as long as those of all the rest of the country combined. It is interesting to note that the effect of the SPG, a marine phenomenon in the oceans surrounding Iceland, is reflected in the condition of the foxes more than the other climatic variables we used in this study, which are largely land‐related. Because Arctic foxes in Iceland feed largely on marine birds and invertebrates, the SPG seems to encompass more accurate information regarding the direct ocean forces that affect food availability to the foxes.     

Variation in the diet of red foxes on Scottish moorland in relation to prey abundance

The diet of red fox Vulpes vulpes was investigated through analysis of 340 scats collected during 1992-1996 from moorland in south-west Scotland. Rodents, game-birds, lagomorphs, carrion and insectivores were the most frequently occurring food types. Fox diet and prey abundance were compared between heather dominant and grass dominant habitat types within the study area. Rodents were the most frequently occurring food type in each habitat and occurred in 63% of seats overall. Rodents occurred more often in scats from grass dominant sites while gamebirds and lagomorphs were more frequent in scats from heather dominant sites. The occurrence of rodents in winter fox scats increased with rodent abundance. In contrast, the occurrence of gamebirds in winter fox scats was unrelated to gamebird abundance but negatively related to rodent abundance. It appeared that foxes switched to gamebirds in years, or habitats, where rodents were uncommon.     

Secondary types of food in the diet of a small-sized mammalian herbivore: from species ecology to habitat inferences

A macroscopic analysis of stomach contents is undertaken in the common vole, Microtus arvalis, obscurus chromosomal form in order to assess the patterns of non-herbivorous foraging activity of a herbivore distributed in different Eurasian biomes. The samples are collected in the Ural Mountains at midsummer (8 localities, 60° 38′ N to 51° 07′ N), and in winter (one locality, 56° 37′ N). Based on the frequency of occurrence and quantitative abundance in stomachs, photoassimilating parts of vascular plants are considered as the primary food category and inflorescences, roots, seeds, mycorrhizal fungi, arthropods, chitin fragments, colorless amorphous matter, and mosses are recognized as the secondary food categories. Although no alternatives to the primary food are found, some of the supplements substitute one another either geographically (fungi in the north versus mature seeds in the south) or by the vertical layers of habitats (seeds and arthropods in the herbaceous layer versus roots and fungi in the shallow layers of soil). Non-staple food consumption in different latitudinal zones at midsummer reflects spatiotemporal dynamics of food availability. From taiga to steppes of the Urals, the common vole is sporadically included in soil foraging but the consumption of mycorrhizal fungi (Glomerales) is only revealed within the zones of dark coniferous forests. The results suggest that the extents of inclusion of a herbivore in soil food webs might vary among ecosystems. In a paleoecological perspective, the analysis of stomach contents in a herbivorous micromammal might reveal fine-scale details of the habitats and provides insights into the stages of the growing season.     

Reproduction of the least weasel in captivity: basic observations and the influence of food availability

The least weaselMustela nivalis nivalis Linnaeus, 1766 is unique among carnivores because of its small body size and capacity for fast reproduction. It has been suggested to be the main agent for maintaining cyclic fluctuations in northern vole populations, largely based on its high reproductive potential and dependence on small rodents. This study describes basic reproductive data on the least weasel obtained during a captive breeding program. In the experiments, food availability was manipulated at the onset of breeding. Altogether, 65 litters were born during the 5-year study. The mean litter size was 5.1 (in 53 litters of known litter size), the most common litter size being six. The sex ratio of weaned young was not biased from 1∶1. The median date of birth was June 4. Food manipulations did not affect mating frequency suggesting that the observation of failure in breeding in years of low abundance of small mammals is due to mortality of embryos or young before weaning, but not due to avoidance of breeding at low food availability. In this respect, weasels differ from other main vole predators, owls and raptors, which often skip breeding if small mammals are scarce.     

Indirect Evidence that Flying Foxes Track Food Resources Among Islands in a Pacific Archipelago

Although flying foxes (fruit bats in the genus Pteropus ) in continental forests often fly between scattered resources, little is known about their ranging behavior among islands. The inhospitable water matrix that surrounds the food patches (islands) in archipelagos may prevent flying foxes from tracking resources as efficiently as their counterparts on larger landmasses do. Our aim in this study was to determine whether the abundance of foraging flying foxes ( Pteropus tonganus ) reflected food availability on islands in the Vava'u archipelago of Tonga, regardless of island size and isolation. Overall, food availability was the strongest determinant of flying fox abundance, and spatial aspects of the islands (land area within 10 km) had only a small influence. Food availability appears to regulate flying fox abundance only when food resources are low, but when food sources are plentiful, flying fox abundance may be high or low. These results provide indirect evidence that flying foxes are able to track food resources efficiently in an archipelago, and the water matrix that surrounds the food patches (islands) is not a strong deterrent for foraging animals.     

Bark consumption of voles in relation to snow cover, population density and grazing impact

The voles Microtus agrestis and Clethrionomys glareolus consume more bark in north than in south Sweden in spite of lower availability in the former area due to snow-pressed ground. Field experiments with genetically homogeneous aspen sticks demonstrated that this pattern reflects differences in animal behaviour and not differences in plant palatability. Within a region consumption in snow-pressed areas was usually less than in subnivean spaces or runways. The amount of bark consumption was generally correlated with population density and amount of grazing impact. Great differences in bark consumption between north Swedish areas with high peak vole populations and south-central Swedish areas with moderate peak numbers imply that nutrition is of different importance at population declines in different regions. The ability of voles to move into snow-pressed areas and locate woody sticks in the snow suggests that control of ground cover will not protect tree seedlings at reforestation.     

Spatial patterns of red fox (Vulpes vulpes) dens in a semi-arid landscape of North Africa

Different patterns of the use of space by red foxes ( Vulpes vulpes ) depend mainly on the distribution and availability of food patches. Nevertheless, other key resources such as suitable areas for dens can also influence the territory size and social organization of this predator. In fact, landscape modifications such as habitat patchiness made by human activities (agricultural practices, urbanization) should create ample unfavourable zones for den settlement. Several studies focused on the distribution and use of dens by the red fox in different habitat types but we do not have any data on the den settlement and habitat composition in the semi-arid regions of North Africa. This study was carried out at Djerba island (SE Tunisia), virtually deprived of surface water and with a semi-arid climate because of the bordering Sahara desert. We used line and random transects to find fox dens. The spatial pattern of dens varied mainly in different habitat types and geoclimatic regions. Highly fragmented areas were also occupied by foxes that built dens close together inside small suitable patches. Fox dens had fewer entrances in more arid regions and they mainly faced the south. Habitat selection was influenced by water availability and irrigated tree plantations that modify soil textures allowing digging of dens. Moreover, asphalt roads limit den settlements. We suggest that the choice of denning sites by foxes depends on persisting harsh conditions and human activities.     

Geographic variation of the winter diet of the Red Kite Milvus milvus in the Iberian Peninsula

This paper presents data on the winter diet of the Red Kite Milvus milvus in the Iberian Peninsula. The diet composition and its relationship with food abundance and habitat characteristics were studied in two areas (northern and southern plateaux) with different food availability and habitat composition. Red Kites showed a clear difference in diet between the two plateaux, with a higher consumption of carrion on the northern plateau and of game species on the southern plateau. Common Voles Microtus arvalis , despite being a very abundant prey on the northern plateau, did not appear in the diet in high numbers, but their frequency was related to the proportion of land occupied by nonirrigated (cereal) crops. High frequencies of "small" carrion (e.g. farmed Rabbits Oryctolagus cuniculus , poultry Gallus sp.) in the diet were correlated with a decrease in the trophic diversity which suggested an active choice of the food, at least in years with low vole densities. We suggest that the use of carrion is influenced not only by its abundance but by its availability to Red Kites. Refuse tips and muladares (dumps for dead livestock) and their management during the winter are important for the conservation of the species.     

Life history,population dynamics and production of Pontoporeia hoyi (Crustacea,Amphipoda) in relation to the trophic gradient of Lake George,New York

The life history characteristics, population dynamics and production of Pontoporeia hoyi in Lake George, New York, were studied from May 1981 through October 1982. P. hoyi, in terms of both density and standing crop, is the most prevalent member of the deep water macrobenthos of Lake George. It reproduces in the winter, with young being released in the late winter-early spring. At the southernmost study site, young released in the spring grew to 6–7 mm in length and bred during their first winter. At the remaining sites, P. hoyi required two years to complete its life cycle. This difference in life history characteristics can be related to food availability and temperature differences. The open waters of the south end of Lake George are not only more productive but are also more closely associated with the littoral zone, providing a wealth of bacteria-rich detritus for benthic deposit feeders. The greater food availability in the south basin of Lake George is reflected in significantly larger brood sizes and smaller size at maturity for P. hoyi populations from the south end of the lake.The southernmost study site has significantly greater P. hoyi density and standing crop than all other sites. The cohort of the year dominated density and standing crop at the southern site while the cohort of the previous year dominated standing crop at the other sites. Peak abundance ranged from 600 · m^–2 at the north site to 2 900 · m^–2 at the south site. Cohort production ranged from 2g · m^–2 at the north site to 15g · m^–2 at the south site.     

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.     

Contrasting phenotypic correlations in food provision of male Tengmalm's owls (Aegolius funereus) in a temporally heterogenous environment

Summary We examined the food provision rate of male Tengmalm's owls,Aegolius funereus, during one 3 year vole cycle consisting of consecutive low, increase and peake vole years. The data were collected in the midnestling period when males provisioned the whole family. In the low vole year, males with a low loading index (g/cm²) of flying area fed their offspring more often than did males with a high loading index, whereas in the peak vole year the opposite trend was evident. Similar relationships were found in the food mass provisioned to the nest. In the increase vole year, male body size had no effect on feeding efficiency. In the peak vole year, when large voles are abundant, heavy males preyed on larger voles than were generally available in their territories, indicating that largeness may increase strike power in hunting attempts. In the low vole year, when breeding is costly due to food scarcity and extensive hunting area, small males are more economical fliers and efficient hunters than large males. The contrasting trends in correlations between male size and feeding efficiency in years of vole abundance versus scarcity suggest that no fixed phenotype may most efficiently cope with variable food supply.