Understanding the intricacy of canid social systems: Structure and temporal stability of red fox (Vulpes vulpes) groups.

Red foxes have a highly flexible social system. Despite numerous studies worldwide, our understanding of the pattern and stability of fox social relationships remains limited. We applied social network analysis to camera trap data collected at high-quality foraging patches to examine the social structure of a population of urban red foxes. Foxes encountered a conspecific on 13% of patch visits, and had significant preferred and avoided companionships in all seasons. They also associated in communities that matched territorial space use, confirming that territories can be analysed separately to increase power without excluding too many social partners. Foxes maintained stable, long-term relationships with other territory residents, but the average longevity of relationships varied seasonally, suggesting that social connectivity, particularly between foxes from different social groups, is influenced by their annual life cycle. The probability of re-association after a given time lag was highest in spring and summer, during cub birth and rearing, and lowest in the winter mating season, when mean relationship duration was shorter. 33% of fox relationships lasted for four consecutive seasons and were probably between territory residents. 14% lasted for around 20 days and were probably between residents and visitors from adjacent territories. The majority (53%) lasted less than one day, particularly during dispersal and mating, and were probably between foxes from non-adjacent social groups. Social structure varied between groups; in one group the death of the dominant male caused significant social disruption for two seasons. This is the first application of social network analysis to multiple red fox social groups. However, our analyses were based on interactions at high quality food patches; social connections may differ when foxes are resting, travelling and foraging elsewhere in their territory. Our results will inform management practices, particularly for disease spread and population control.