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.     

内蒙古达赉湖地区赤狐生境选择及生境景观特征分析

近年来,干旱、鼠害及人为对草原的过度开发利用等对达赉湖地区赤狐的数量及分布范围造成了很大影响。借助于主成分分析、Bailey’s判别分析、遥感和地理信息系统对达赉湖自然保护区赤狐的生境选择及生境的景观特征进行了研究。在赤狐家域内采用样线法和样方法,共设置10 m×10 m样方245个(实验样方101个,对照样方144个),并测定样方内的7个生境因子:植被类型、隐蔽级、食物丰富度、雪深、距水源距离、距围栏距离、距居民点距离。对样方内数据进行主成分分析得出,食物和隐蔽性是赤狐生境选择的主要因子,距居民点距离和距围栏距离是赤狐生境选择的次要因子,雪深和距水源距离是赤狐在生境选择中未表现出选择和利用的因子。利用Bailey’s方法和基于赤狐的生境分布图的景观统计得出:赤狐偏好选择柳灌丛和芦苇塘两种生境,这两种生境的总面积约为1093.47 km2,占研究区总面积的14.05%;赤狐随机利用河道、典型草原和草甸草原3种生境类型,这3种生境的总面积约为4721.79 km2,占研究区总面积的60.67%。;回避的生境为冰面和沙化草地,总面积约为1968.09 km2,占研究区总面积的25.29%。基于地理信息系统和Fragstats的景观特征分析得出,赤狐最适宜生境的面积最少,斑块数量最多,平均斑块面积最小,平均形状指数最小,平均斑块距离最大;其次是较适宜生境和不适宜生境;一般适宜生境的面积最多,斑块数量最少,平均斑块面积和平均形状指数最大,平均斑块距离最小。赤狐多分布于斑块较大的适宜其生存的生境或分布于由这些斑块形成的生境斑块镶嵌体中。     

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.     

Den distribution and den use by the red foxVulpes vulpes japonica in Kyushu

The distribution of dens and den use by the red fox was studied in Kumamoto Prefecture between 1968 and 1982. A total of 80 dens were classified into natal, residential, and temporarily-visited dens. Natal dens comprised 12.4% of all dens. Relative proportions were quite stable for a long period. This indicates a constant number of reproducing females in the area for a considerable length of time. Seasonally the utilization of den showed three peaks in February, May and October. The peaks coincided well with the peaks of reproductive activity of female foxes. The results of multiple regression analysis of distribution of dens with respect to selected environmental parameters around densites indicated that the fox selectively utilize open land, rather than densely vegetated areas. The persistence of dens, the selective utilization by foxes of open land and the sizes of dens were all related to the breeding activities of female foxes inhabiting the area. Reproductive females usually used plural dens (“den group”) in one reproductive season. The numbers of those females occuring in the area were maintained at a steady level for fairy long periods. These indicate that the existence of a stable social relationship between females mediated through occupancy of “den groups.”     

Arctic Fox Responses to Tourism Activity

As the interest for nature-based tourism activities increases, it is important to provide evidence-based guidelines for wildlife-human interactions to minimize the disturbance caused to wildlife. In Fennoscandia, the endangered arctic fox (Vulpes lagopus) is subject to increasing tourism interest and some regions recommend a minimum approach distance of 300 m, but the guidelines have not been scientifically validated. We conducted experimental human approaches towards arctic fox den sites to study activity and behavioral changes in response to the approaching observer. The first arctic foxes hid when approached within 300 m, but many had increased their vigilance already at the start distance of 500 m. At approximately 200 m, the hiding probability increased rapidly at dens disturbed and undisturbed by tourism activities. Arctic foxes at disturbed dens allowed the observer to approach more closely before they increased their vigilance and before they hid compared to foxes at undisturbed dens. We confirm that a minimum distance of 300 m might be sufficient for most arctic foxes to refrain from hiding, but a longer distance would be required to avoid causing any disturbance. We recommend a minimum approach distance of ≥300 m to be implemented in all Fennoscandian regions inhabited by the arctic fox. © 2020 The Wildlife Society.     

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.     

Red fox takeover of arctic fox breeding den: an observation from Yamal Peninsula, Russia

Here, we report from the first direct observation of a red fox (Vulpes vulpes) intrusion on an arctic fox (Vulpes lagopus) breeding den from the southern Arctic tundra of Yamal Peninsula, Russia in 2007. At the same time, as a current range retraction of the original inhabitant of the circumpolar tundra zone the arctic fox is going on, the red fox is expanding their range from the south into arctic habitats. Thus, within large parts of the northern tundra areas the two species are sympatric which gives opportunities for direct interactions including interference competition. However, direct first-hand observations of such interactions are rare, especially in the Russian Arctic. In the present study, we observed one red fox taking over an arctic fox breeding den which resulted in den abandonment by the arctic fox. On July 19, eight arctic fox pups were observed on the den before the red fox was observed on the same den July 22. The pups were never seen at the den or elsewhere after the red fox was observed on the den for as long as we stayed in the area (until August 10). Our observation supports the view that direct interference with red fox on breeding dens may contribute to the range retraction of arctic foxes from the southern limits of the Arctic tundra in Russia.     

Landscape structure, predation of red foxes on grey partridges, and their spatial relations

The aim of this study was to estimate the effect of agricultural landscape structure on the predation rate of red foxes Vulpes vulpes on grey partridges Perdix perdix during the breeding season and on their spatial relations. The number of partridge remains found around fox breeding dens (N=165) was used as an index of the predation rate in 10 study areas. Moreover, the distribution of both species and the searching intensity of partridge nesting habitat (permanent semi-natural vegetation) by foxes in relation to the landscape structure were studied using scat, track and call counts. The predation index (range 0.06–0.46 partridges/den) increased with spring partridge density and decreased with the occurrence of crop boundaries. The distribution of foxes and partridges in large fields was positively correlated with the occurrence of permanent vegetation, but no such effect was observed in small fields. The searching intensity in permanent vegetation by foxes decreased with the occurrence of these structures among small fields, but not among large fields. The study showed that in a differentiated landscape foxes and partridges had various structural elements at their disposal, which led to partial separation of the predator and its prey in the space.     

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     

Population persistence in a landscape context: the case of endangered arctic fox populations in Fennoscandia

Anthropogenic fragmentation of habitat and populations is recognized as one of the most important factors influencing loss of biodiversity. Since it is difficult to quantify demographic parameters in small populations, we need alternative methods to elucidate important factors affecting the viability of local populations. The Fennoscandian arctic fox inhabits a naturally fragmented alpine tundra environment, but historic anthropogenic impacts have further fragmented its distribution. After almost 80 yr of protection, the population remains critically endangered. Both intrinsic factors (related to the isolation and size of sub‐populations) and extrinsic factors (related to environmental conditions influencing patch quality and interspecific competition) have been proposed as explanations for the lack of population growth. To distinguish between these hypotheses, we conducted a spatially explicit analysis that compares areas where the species has persisted with areas where it has become locally extinct. We used characteristics of the fragments of alpine tundra habitat and individual arctic fox breeding dens (including both currently active dens and historically active dens) within the fragments to evaluate the importance of habitat characteristics and connectivity in explaining variation in persistence within a fragment. The number of reproductive events in a fragment was related to the size of the fragment, but not more than expected following a 1:1 relationship, suggesting little effect of fragment size on the relative number of reproductions. The likelihood of a den being used for breeding was positively associated with factors minimising interspecific competition as well as increasing within‐fragment connectivity. These results support the idea that the failure of Fennoscandian arctic fox to recover is caused by demographic factors that can be related to fine‐scale Allee or Allee‐like effects, as well as environmental influences related to increased competition and exclusion by red foxes.     

Red Fox Use of Landscapes with Nesting Shorebirds

Predation of nests and young is one of the limiting factors in the conservation of birds; understanding environmental covariates of predator distribution can assist with decisions regarding the best management strategies to reduce predation risk. The habitat of beach-nesting birds is often reshaped by storms in ways that may affect nest predation, such as by flattening vegetated dunes where mammals hunt, but human management of beaches tries to prevent the effects of storms on the landscape with unknown implications for predator distributions. Moreover, human development may affect predator distributions by subsidizing food and shelter. To determine the relationship between predator occupancy and landscape features in beach-nesting bird habitat, we repeated mammalian predator track surveys 8 times/year at 90 plots in southern New Jersey, USA, from 2015–2017. We used dynamic occupancy models to estimate the probability of use by red foxes (Vulpes vulpes) and to document changes in habitat use as related to landcover types over the avian breeding season within years. We had 373 red fox detections with years pooled. Detection probability for red foxes varied by year, and probability of use decreased as the distance to the nearest primary dune increased. We found no evidence that red fox habitat use depended on distance to human development. Our results suggest that conserving nesting habitat that includes open areas (i.e., storm overwash [whereby vegetation is scoured by tidal flooding]) may reduce predation risk because beach-nesting birds would not be forced into nesting close to dunes, which are typically used for hunting by red foxes. © 2020 The Wildlife Society.     

Effects of Roads on Endangered San Joaquin Kit Foxes

ABSTRACT San Joaquin kit foxes (Vulpes macrotis mutica) occur in central California, USA, and are endangered due to habitat loss and degradation. As the human population of California grows, more roads are being constructed in remaining kit fox habitat. We examined effects of 2-lane roads on demographic and ecological patterns of kit foxes on the Lokern Natural Area (LNA) from August 2001 to June 2004. Of 60 radiocollared kit foxes, only one was struck by a vehicle. Foxes were assigned to 1 of 3 risk categories (high, medium, or low) based on proportion of time spent in road-effect zones, which were defined by the probability of a fox encountering a road during nocturnal movements. Fox survival probabilities, reproductive success, litter size, nocturnal movements, and den placement all were similar among risk categories. Nocturnal locations of foxes were closer to roads than were den locations, and den fidelity was lowest for medium-risk foxes and highest for low-risk foxes but intermediate for high-risk foxes. Food availability and use were not affected by proximity to roads. We were unable to detect any significant detrimental effects from 2-lane roads on kit fox demography and ecology. Our results suggest that standard mitigation strategies, such as crossing structures and exclusionary fencing, would not benefit kit foxes on the LNA.     

The feeding choice of the Red Fox (Vulpes vulpes) in a semi‐arid fragmented landscape of North Africa in relation to water and energy contents of prey

In semi‐arid zones, the lack of surface water for drinking forced predators to obtain it from the body fluids of prey and this requires more different prey than that needed to provide energy alone. We examined Red fox (Vulpes vulpes) scats to ascertain if selective feeding compensates the shortage of free water by the ingestion of more prey rich in water such as insects. In addition, we discussed our results in light of increasing habitat fragmentation that, in turn, influences the prey availability. At different spatial levels, recombination of prey assemblages in the diet reaches a solution for variable shortage of water. This was true especially at the landscape level (different degree of aridity) and indirectly for habitat types. Nevertheless, the effects of landscape characteristics probably influenced foxes through the influence of suitable habitats, which partitioned prey species. In fact, increasing values of patchiness had a negative effect on measured energetic gain indices but foxes try to exploiting different patches including small and isolated ones because these surveys become important to obtain water moistures from alternative prey (Orthopters).     

Distinct patterns in alpine vegetation around dens of the Arctic fox

The arctic fox Alopex lagopus excavates its dens in gravely ridges and hillocks, and creates a local environment quite distinct from the surrounding tundra or heath landscape. In northern Sweden, the vegetation of 18 dens of the arctic fox was investigated, as well as reference areas off the dens but in geologically and topographically similar locations. The species composition showed considerable differences between den and reference areas, with grasses and forbs occurring more abundantly on the dens, and evergreen dwarf-shrubs occurring more in reference areas. The effect of the foxes' activities is thought to be either through mechanical soil disturbance, or through nutrient enrichment via scats, urine, and carcasses. This was expected to result in differences in plant traits with key functional roles in resource acquisition and regeneration, when comparing dens with reference areas. We hypothesised that the community mean of specific leaf area (SLA) would differ if nutrient enrichment was the more important effect, and that seed weight, inversely proportional to seed number per ramet and hence dispersal ability, would differ if soil disturbance was the more important effect. Specific leaf area showed a significant difference, indicating nutrient enrichment to be the most important effect of the arctic fox on the vegetation on its dens. Arctic foxes act as ecosystems engineers on a small scale, maintaining niches for relatively short-lived nutrient demanding species on their dens in spite of the dominance of long-lived ericaceous dwarf-shrubs in the landscape matrix. Thus, foxes contribute to the maintenance of species richness on the landscape level.     

Foraging ecology and spatial behaviour of the red fox (Vulpes vulpes) in a wet grassland ecosystem

We investigated diet composition, habitat selection and spatial behaviour of the red fox (Vulpes vulpes) in relation to the availability of wader nests in a coastal polder area in southwest Denmark. The predatory role of the red fox in wet grassland ecosystems has profound implications for conservation status of declining populations of grassland breeding waders. However, few studies have focussed on the foraging ecology and behaviour of the red fox in these landscapes. Faecal analyses revealed that fox diet consisted of birds (43 % of prey remains?/?32 % of biomass), rodents (39 %?/?21 %), sheep (mainly as carrion, 14 %?/?41 %) and lagomorphs (4 %?/?7 %). Charadriiformes (including waders) comprised 3–12 % of prey remains throughout the year. Telemetry data and spotlight counts indicated that foxes did not select areas with high densities of breeding waders, suggesting that foxes did not target wader nests while foraging. Foxes maintained stable home ranges throughout their lives, indicating that the area sustained a permanent fox population all year round. The population densities, estimated from spotlight surveys, were 0.74 visible foxes km^?2 (95 % CI; 0.34–1.61) on the preferred breeding habitat for waders and 1.21 km^?2 in other open habitats such as cultivated fields. Our results indicate that red fox predation on wader nests is incidental, consistent with the notion that red foxes are generalist predators that opportunistically subsist on many prey groups.     

Annual dynamics of den use by red foxesVulpes vulpes and badgersMeles meles in central Poland

This study on den use by red foxesVulpes vulpes (Linnaeus, 1758) and badgersMeles meles (Linnaeus, 1758) was conducted in central Poland (51°48’N, 19°53’E) over two years. We found a strong negative correlation between monthly den utilisation rate by foxes and air temperature (r=−0.83,p=0.0001). Correlation between temperature and den use by badgers was positive but less distinct (r=0.40,p=0.036). During the whole study period, a negative correlation between the monthly intensity of den use by badgers and by foxes was found (r=−0.51,p=0.008). The seasonal pattern of den utilisation differed between the species. Foxes used the dens mainly in winter and spring whereas in summer and autumn the intensity of den utilisation sharply decreased to a minimum in August-September. A peak in den use by foxes was observed in March and coincided with rut and selection of future breeding dens. In badgers, two peaks in den use were observed: the first in April and the second in August-September. An individual badger used more dens and used them more evenly throughout the year than an individual fox. The same dens were visited by foxes and badgers mostly from January to April, with a maximum in March.     

Habitat selection by two predators in an urban area: The stone marten and red fox in Wrocław (SW Poland)

We analyzed the habitat use of stone martens and red foxes based on incidental observations within the urbanized zone of Wrocław, SW Poland. We compared proportional habitat use at observation sites with randomly selected points and evaluated differences in distance to the water sources and to urban boundaries. Habitat use by both species was different from what we had expected from random points. Stone martens used high-density housing more frequently than red foxes and that expected from random points and avoided open and industrial areas, whereas red foxes used housing estates significantly more often than expected and avoided high-density housing. Both species used the other habitats according to their availability. Stone martens often selected habitats located closer to the city centre, whereas red foxes often selected habitats closer to urban borders than expected. The distribution of red foxes and stone martens is influenced by several factors including the availability of shelter and food, as well as the opportunity to move around undetected. Interspecific competition may also play an important role in habitat selection. Stone martens seem to be better adapted to urbanized areas than red foxes.     

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.     

Spatial use and interaction of the invasive raccoon dog and the native red fox in Central Europe: competition or coexistence?

The main objective was to discover extent of interference and/or exploitative competition between the native red fox (Vulpes vulpes) and the introduced, invasive raccoon dog (Nyctereues proconoides) in the intensively used, agricultural landscape of northeast Germany (Mecklenburg-Western Pomerania) using very high frequency (VHF) radio telemetry. We recorded location data for 12 foxes and 16 raccoon dogs between July 2004 and December 2006. Species had similar average home range sizes estimated in each season (K95). Home ranges of adjacent raccoon dogs and foxes overlapped from 0.5 to 74.5 % with a mean of 26.4 %. We found a significantly different home range overlap index between the species showing that raccoon dog ranges shifted between seasons to a greater extent than red fox ranges. The raccoon dog differed significantly from the red fox in its use of habitat types, preferring dense vegetation cover and avoiding open areas. The red fox displayed less preference for or avoidance of specific habitat types. Moreover, an almost neutral inter-specific interaction index ranging from ?0.12 to 0.12 indicates that raccoon dogs and red foxes ignored each other. It is concluded that widespread and available resources and differences in spatial use patterns prevent competition between red foxes and raccoon dogs in the agricultural landscape of northeast Germany.     

Resource partitioning among cape foxes,bat-eared foxes,and black-backed jackals in South Africa

Cape foxes (Vulpes chama) and bat-eared foxes (Otocyon megalotis) are sympatric with black-backed jackals (Canis mesomelas) over much of southern Africa, although competition with and/or predation by jackals may suppress local populations of both fox species. From 2005 to 2008, we captured, radio-collared, and monitored 11 cape foxes, 22 bat-eared foxes, and 15 black-backed jackals on a game ranch in South Africa to investigate their spatial, habitat, temporal, and dietary resource overlap. Mean annual home-range sizes were 27.7 km² for cape foxes, 5.0 km² for bat-eared foxes, and 17.8 km² for jackal family groups. Home ranges overlapped completely between species, although core areas overlapped less (<45%), with cape foxes and jackals overlapping the least (12%). When active, cape foxes, but not bat-eared foxes, used core areas of jackal groups less than expected. Additionally, both fox species used jackal core areas less than expected for their den sites, suggesting areas outside jackal core areas were used as refuges by foxes. Strong levels of habitat partitioning were not apparent at the study site or home-range levels, although habitat selection for den sites differed between jackals and cape foxes. Jackals were the most diurnal across seasons, whereas cape foxes were the most nocturnal. Diets overlapped little (R₀ = 0.20–0.34) among the canid species, with bat-eared foxes overlapping the least with the others. Jackals killed at least 5 collared bat-eared foxes and 1 collared cape fox, indicating potential interference competition, probably for exclusive use of territorial space rather than over shared resources. We conclude that bat-eared foxes coexisted with jackals primarily by their dietary specialization and group living. Cape foxes coexisted with jackals by exhibiting high levels of spatial, habitat, temporal, and dietary partitioning. Surprisingly, the fox species exhibited positive associations with each other. Our results show the mechanisms that may allow jackals to suppress fox populations, yet also show how foxes, in turn, use different mechanisms to coexist with a dominant canid. © 2012 The Wildlife Society.