Life History Reproduction of red foxes is described by Storm et al. (1976) and Henry (1993). The gestation period and time of weaning are constant over different populations. Timing of mating and birth, by contrast, may vary geographically and is even subject to variation within a small area. Dispersal of juvenile foxes takes place every year, and because of variation in the time of birth, one might suspect that the time at which dispersal begins would also reflect strong geographic variation. In fact, the time of year at which dispersal is initiated seemed to be constant over all the populations studied. These and other aspects of the life of an urban fox over a single year are detailed in the following paragraphs. Courtship for foxes begins in mid to late December. Spermatogenesis begins in October (even in first-year males), and continues through March (Storm et al., 1976; Henry, 1993). When courtship starts, previously solitary foxes travel in pairs, or sometimes in groups of three. Mating usually takes place in January and February, although it may be earlier in more southern areas and later in northern areas. In the city of Minneapolis, I observed fox cubs that were about six weeks old as early as 7 May, and as late as 5 August. Therefore, there is at least a three-month window during which mating can take place and result in production of viable offspring. Given a gestation period of 52-53 days, which is consistent through the literature for all fox populations, these observations indicate that mating takes place in the Minneapolis area between late January and late April, and possibly earlier or later. Although Storm et al. (1976) claimed that breeding can take place as late as April, they noted that 94% of the rural foxes they studied bred between mid-December and mid-February, with a peak during the third week of January. Only 1% of their sample had bred after mid-February. Harris and Smith (1987) reported that for foxes in both London and Bristol the peak birth period was March, indicating a peak breeding period in January. They noted that in those cities matings can also take place in early December (about 10%), or after the first week of February (about 25%). No study has been conducted directly comparing the lengths of breeding seasons in urban areas as opposed to nearby rural areas, but it seems that the breeding season may be extended somewhat in urban areas. If this is true, the most likely explanation is the protection an urban environment provides from the extremes of weather. Most importantly, food is not scarce in urban areas in winter, as it often is in rural areas (Harris, 1977; Storm et al., 1976). The availability of food also affects the composition of fox family groups (von Schantz, 1984; Zabel and Taggart, 1989). A typical family group consists of the dog fox and vixen, plus up to four females that may or may not breed. Macdonald (1979) referred to these females as "helpers," and observed a definite dominance hierarchy in the family group, wherein the "helpers" were subordinate to the breeding female. Harris (1981a) also noted that some family groups in Bristol produced more than one litter of cubs in a given year, indicating that some of the "helpers" may also breed under certain conditions, presumably when resources are plentiful. Space is an important resource for foxes. Each family needs several safe den sites in which to raise their cubs in addition to safe resting sites outside of the den. Den space seems to be readily available for urban and rural foxes alike. Foxes maintain territories, and it is common for them to have several different dens within their territory. Prior to parturition, the mated pair selects a den, called the natal den, in which to bear the cubs. This den is usually larger than other available dens, and has several exits (Henry, 1993). The other dens can be as simple as a single entrance leading to a hole where the foxes can take shelter. In urban areas, a great variety of den sites is available to foxes, but places to rest during the day are scarcer. Denning, home ranges, and daytime resting sites are covered in a separate section. The natal den must be safe, because the fox family is most vulnerable to predators during the first two weeks after parturition. Fox cubs are blind and helpless for about ten days after birth. The vixen stays with them constantly for up to two weeks (longer in colder climates than in warmer climates), during which time she is dependent on the dog fox and any "helpers" for food. After this time, she may leave to hunt for herself, but will return regularly to nurse the cubs. The parents may first move the cubs to an alternate den when they are about two weeks old. The cubs usually are moved at least once, and may be moved several times during their first summer. For example, all the dens I located were abandoned in late spring. In two of the rural dens, human disturbance might have led to the abandonment, but humans frequented the immediate areas of the urban dens at least every other day, so it is unlikely that human disturbance was the only cause of abandonment of the urban dens. This suggests that urban foxes, like rural foxes, move frequently between dens. Movement may help protect the fox family from humans and other predators when the cubs are most vulnerable. When the cubs are about five weeks old, they begin to venture outside the den. Weaning takes place at this time, so the parents bring back prey for the cubs to eat. Later in the summer the cubs learn to hunt for themselves, and their parents will provide less food and spend less time at the den. The cubs range farther from the den as they grow older, hunting for themselves but remaining in the territory of their parents. October is the month during which dispersal of foxes is most likely to begin. Not all cubs disperse, and not all of those that disperse leave in their first year. In a study of dispersal, Harris (1988) found that only 3.5% of female cubs and 10.8% of male cubs dispersed before 1 October. By the end of December, Harris (1988) found that 32.5% of tagged females and 58.5% of tagged males had dispersed. Most of the animals that dispersed did so by the end of their second year, the final proportions being 37.8% for females and 75.8% for males. This corresponds closely with data gathered by Storm et al. (1976) on rural foxes, which indicated that most of them also disperse in October. The proportion of foxes dispersing in the rural population they studied was somewhat higher, reaching 58% for females and 96% for males. Population density is probably the most important factor affecting dispersal. In urban areas, for example, Harris (1988) found that male cubs from large litters were more likely to disperse than male cubs from small litters. He also found that animals from areas of low fox density (indicating poor fox habitat) dispersed farther than animals from areas of high or medium fox density. Another factor affecting dispersal is the food supply in the territory. Cubs might remain in their natal territory longer in urban areas, where food is plentiful, than in rural areas. However, the sample sizes and geographical areas of the studies by Harris (1988) and Storm et al. (1976) are too small to draw a definite conclusion. Once the cubs have dispersed, the adults and remaining cubs stay on the same territory, but continue to hunt alone. They may rest with other foxes in areas where resting habitats are limited, but do not associate in regular pairs until courtship begins again in December.