Dens and Home Ranges The most distinctive feature of an urban environment is the abundance of man-made structures that offer shelter for many non- human animals. Foxes take advantage of man-made structures, sometimes even setting up their natal dens in occupied houses (Harris, 1981a; Harris, 1980). Harris (1977) and Storm et al. (1976) determined that the most important factor governing the distribution of breeding dens for both urban and rural foxes was human disturbance. Harris (1977) located 97 dens in London, 46 of which were under residential buildings (36 of these under garden sheds). Of the remaining 51, more than half (29) were in banks of earth, such as garden banks or railway embankments. Some foxes raised their cubs under the floorboards of occupied houses, often gaining entrance through a pet door. In one case, a domestic cat and dog, both of which tolerated the foxes, were present in the household where a fox den was located (Harris, 1980). Usually, foxes avoid dogs; the distribution of foxes in Bristol showed a significant negative correlation to the distribution of feral dogs (Harris, 1981a). Fox distribution in urban areas is also strongly influenced by the local habitat. The number and type of habitats available to foxes in an urban area are highly variable. Harris (1977) listed some of the habitats found in London, and noted that most of the foxes he studied were recovered from residential habitats, industrial habitats, and vacant land with no public access. Surprisingly, despite reports that open public parks were important for daytime shelter of foxes, Harris recovered only 8.73% of his foxes from parks, although parks comprised 10.53% of suburban land. By contrast, nearly 60% of his foxes were recovered from residential habitats, which comprised just over 40% of suburban land. Harris and Rayner (1986) found a significant positive correlation between areas of high fox density and residential areas where the housing was owner- occupied. Harris (1977) speculated that space for daytime shelter is an important limiting resource in London, and that the availability of suitable daytime resting habitat has a strong influence on the distribution of foxes. He found that between autumn and spring, several foxes will often share a daytime retreat. This clumping is not unknown in rural fox populations, but it is more common and takes place for a longer period of time in urban and suburban fox populations (Harris, 1977). Clumping is to be expected in urban and suburban populations not only because daytime resting sites are rare, but also because fox densities are generally higher in urban areas than in rural areas. Rural foxes are known to prefer regions of diverse habitat, such as edge habitats (Henry, 1993). The abundance of such diverse regions in an urban area may account for the unusually high density of foxes found in cities -- Harris (1977) recorded up to five family groups per km2 in several of his studies. By contrast, two families of foxes studied by Storm et al. (1976) in Minnesota stayed within separate well-defined areas of 9.6 km2 each. The average home range of an urban fox is significantly smaller than the average home range of a rural fox (Harris, 1980; Coman et al., 1991; Doncaster and MacDonald, 1991). In a study of foxes in Australia, Coman et al. (1991) found that two family groups of foxes living on farmland had home ranges of 5.0 and 7.2 km2, while three family groups of foxes on the fringe of an urban area had home ranges of 0.6, 1.3, and 0.8 km2. Doncaster and Macdonald (1991) found a similar difference between foxes living in central Oxford and foxes living in suburban and rural areas around Oxford, and also found that the home ranges of urban foxes changed over time. "City ranges were not spatially stable over months or even weeks," Doncaster and MacDonald (1991) wrote. "They moved in step-wise extensions to encompass new areas whilst at the same time contracting other parts of the range to expel old areas." This movement of territories is unique among foxes. Doncaster and Macdonald speculated that movement of home ranges is a behavioral adaptation developed since foxes invaded urban areas in England in the 1930's. The average amount of food available over the area of a city is higher than in a similarly sized rural area, but the variance in food availability is also much greater. For foxes to make the best use of the resources in urban environments, they must constantly explore new areas and re- explore old ones. This activity would require far too much energy to be viable in a large rural home range, but the high density of different patches in a city permit foxes following this strategy to survive and prosper.