Encounters between Red Foxes (Vulpes vulpes): Implications for Territory Maintenance, Social Cohesion and Dispersal

“Although the proximity loggers cannot achieve absolute spatial precision, they provide much better spatial resolution than radiotracking studies [5,23] and far more complete and less labourintensive data collection than direct observation methods (e.”

“They concluded, however, that neither the social subordination hypothesis nor the social cohesion hypothesis alone explained the dispersal behaviour of foxes (White & Harris 1994).”

“Avoidance of roads was quantified by comparing the observed number of roads crossed during radiotracking sessions with the predicted number crossed for randomly generated patterns of movement (White & Harris 1994).”

“The expected number of roads crossed per night was estimated from randomly generated patterns of movement (hereafter, ‘‘simulated’’ movement) using the random walk model developed by White and Harris (1994).”

“Although these factors are expected to contribute to variation in detection distances, this is trivial compared to positional error of traditional radiotelemetry locations (White and Harris 1994, Gehrt and Fritzell 1998, White et al. 2000, Ramsey et al. 2002, Atwood and Weeks 2003).”

“Past researchers have 61 quantified contact rates by observing contacts visually (Totton et al. 2002) or using 62 telemetry to infer how often animals come in close proximity (White and Harris 1994, 63 Caley et al. 1998, Ramsey et al. 2002, White et al. 2003, Ji et al. 2005).”

“Similarly, direct transmission between adults within social groups, and from adult to cubs, is also likely to be common (see White & Harris, 1994; Baker et al., 1998; Baker & Harris, 2000).”

“Radiotelemetry has been used to obtain detailed information on animal ecology (Saunders et al. 1993, White and Harris 1994), but the advent of the GPS collar has enabled researchers to assess these behaviors remotely on far ranging species moving rapidly across inaccessible terrain (Weimerskirch et…”

“Dispersal in carnivores has been studied mainly in social species (e.g., Harris and White, 1992; Harrison, 1992; White and Harris, 1994), but few studies have focused on solitary felids, and most of them were on species of large size (Beier, 1995; Maehr et al.”

“The highest incidence of bite wounds occurred in winter and spring, coincident with the mating and dispersal seasons (White & Harris 1994).”

“Foxes, like most territorial animals, mark their territories and contact their neighbours regularly (White and Harris, 1994; Goszczynski, 2002).”

“White & Harris (1994) found that direct encounters, although almost always aggressive, were uncommon in red foxes (Vulpes vulpes), and therefore a relatively unimportant means of territory defense.”

“We analyzed each movement singly, with the assessment corridor set to 25-m width, corresponding with the accuracy of the fixes (White and Harris 1994).”

“Survival analysis Capture–mark–recapture of foxes in Bristol was carried out as part of an ongoing study into the social and spatial organisation of red foxes (White and Harris 1994; Baker et al. 1998).”

“Traditionally, studies have relied on radio-tracking data to infer dyadic encounters between individuals, with an encounter (typically referred to as a ‘contact’) being defined as broadly as two individuals logging spatial locations within 100 m of each other (White and Harris 1994; Ramsey et al. 2002; Schauber et al.…”

“…rabies and on social fox communities in which intragroup contact rates are assumed high enough to spread an infection throughout the whole group within at least two months (for estimates of group contacts see, for example, MacDonald et al. (1981), White & Harris (1994) and White et al. (1995)).”

“In the wild, antagonistic encounters occurring between unfamiliar individuals is common (White and Harris 1994).”

“However, our understanding of red fox sociality in Australia is limited (Newsome 1995) and males may be more likely to be involved in aggressive encounters with conspecifics than females (White and Harris 1994).”

“Here, we study a modified version of the territorial random walk model where animals are random walkers with an attraction towards a central place, such as a den or nest site where the animals return occasionally [9], or a core area where animals tend to spend most of their time [10].”

“The detection distance of the proximity collars was set to approximately 5 m, which we estimated to be a reasonable distance from which two animals could detect one another (White and Harris 1994) or for infectious disease to be transmitted via aerosol (Gorham and Brandly 1953; Xie et al. 2007).”

“Encounters between unfamiliar male territory holders are often aggressive, especially during the mating season (White and Harris, 1994).”

“We used the lowest observed seasonal estimate (spring, 0.05) as our low contact rate, the highest observed seasonal estimate (winter, 0.76) as our high contact rate, and the overall mean of 0.28 as our midrange contact rate (White & Harris 1994; White et al. 1995) (Supporting Information).”

“Brief meetings between foxes sharing a territory are quite common, but contacts between foxes of adjacent territories are generally avoided (Harris, 1986; Macdonald, 1987; White and Harris, 1994).”

“…terms, relating to the territorial nature of foxes (Catling and Philips 1991; Coman et al. 1991; Phillips and Catling 1991; Adkins and Stott 1998) and their use of scent markings to delineate territorial boundaries (White and Harris 1994) and to mark the locations of food sources (Henry 1977).”

“Previous research showed that young foxes can be highly aggressive towards each other (White and Harris 1994), and aggression among young canids can result in the emigration of lower ranking individuals (Gese et al. 1996).”

“While this is questionable, some modified versions, as the one proposed by [58], use correlated random walks as null models and simulated independent trajectories under these models to replace DE by a more realistic reference value.”

“…ones (Gaines & McClenaghan, 1980; Greenwood, 1980; Baird & Birney, 1982; Adler et al., 1984; Mech, 1987; Anderson, 1989; Hanski et al., 1991; White & Harris, 1994; Gese et al., 1996; Dobson et al., 1998; Altwegg et al., 2000; Clobert et al., 2001; Serrano et al., 2001), and when dispersing,…”

“Such risk of physical aggression during a neighbour encounter also applies to canids [53, 54].”

“…frequency and duration of contact between individuals include visual observation (Grenier et al. 1999, Totton et al. 2002), radiotelemetry (White and Harris 1994, Caley et al. 1998, Ramsey et al. Received: 22 December 2010; Accepted: 7 August 2011; Published: 28 October 2011 1E-mail:…”

“In this sense, the barking sequence may function in spacing individuals for mutual avoidance (e.g. Harrington & Mech 1983) or exchanging information without close contact, which may otherwise be likely to lead to an aggressive encounter (e.g. White & Harris 1994).”

“…females, trespass into another group’s territory, perhaps searching for extra-group mating opportunities (see also Pal et al. (1998a) for dogs; Bekoff and Wells (1986) and Gese (2001) for coyotes; and Voigt and Macdonald (1984), Saunders et al. (1993) and White and Harris (1994) for red foxes).”

“The pressure of neighboring groups may have negative consequences because of agonistic encounters [9, 10, 25], which may affect the maintenance of territories [26].”

“…daily travel are used to compensate for altered food availability, periods with different resources should lead to varied patterns of between-group scramble (BGS) and between-group contest competition (BGC) (e.g., for BGC: red foxes, White and Harris 1994; African elephants, Wittemyer et al. 2007).”

“Social organisation has also been shown to vary temporally in relation to changes in activity connected with breeding and dispersal (White and Harris 1994; Ji et al. 2005; Böhm et al. 2008).”

“…(ii) group size estimates are smallest, and most realistic, when based on social connections because foxes avoid encounters with nongroup members (White & Harris, 1994); (iii) groups are largest in autumn prior to the onset of dispersal (Baker et al., 2007; Soulsbury et al., 2008); (iv)…”

“In some instances this can lead to increased territoriality and home range stability (White and Harris 1994; Soulsbury et al. 2010), but in most cases it leads to a breakdown of territoriality and an increase in home range overlap (Hadidian et al. 2010; Harris et al. 2010).”

“In fact. the mating season is riskier for small males (White and Harris, 1994), which face high er risks from incursions by larger males in their home ranges.”

“As in other canids, including red foxes (Doncaster and Macdonald 1997; Macdonald 1977, 1979; Meia and Weber 1996; White and Harris 1994), interactions between foxes in the same social group were almost always affiliative, often involving allogrooming, play, resting, or foraging together.”

“In the wild, antagonistic encounters between unfamiliar individuals occur (White & Harris, 1994) and the level of risk increases with increasing density, with associated competition for space, mates, and F I G U R E 4 The effect of competing predators on the waiting distance of collared leopards…”

“In many mammals, natal dispersal is triggered by aggression received from same-sexed adults (White and Harris 1994; Smale et al. 1997; Feder et al. 2019), although evidence for this dispersal mechanism in male blue monkeys is mixed (Rudran 1978; Ekernas and Cords 2008).”

“Intraspecific,intergroup aggressiveinteractions are frequent in many social animals (e.g., insects, 1 birds, 2 and mammals 3–6 ).”

“Among Vulpes species in North America and Europe, contact rates were shown to peak during the mating season in winter (White and Harris 1994; Murdoch et al. 2008; Ralls et al. 2013), suggesting peaks in contact rates typically occur during the winter or dry season in canids worldwide.”

“, 2004), although aggressive displays can still escalate into fights (White and Harris, 1994).”

“This pattern of aggression is also seen in free-living foxes, where the sub-adult fighting injuries increased at the start of the dispersal period (White and Harris, 1994).”