Foxes – Ecology and sociality

The red fox Vulpes vulpes is found over much of the planet, and with a degree of ecological flexibility which is closely linked to its dietary breadth. Exploration of fox ecology, made possible by advances in technology beginning in the 1970s, has revealed how the sociology of foxes is considerably more complex than was previously thought, and how that complexity is related to their food resource ecology. Foxes live in territories with marked boundaries, most often in pairs where resources are scarce and widely dispersed. But where resources are both abundant and aggregated, family groups occur. In rural Oxfordshire, these family groups consist of a dominant breeding pair with several non-breeding vixens. The sub-dominant vixens assist with feeding and guarding of cubs. Urban foxes have a similar social system, but less stable territories, probably because the urban environment is hazardous and less predictable. Territories are configured to include resource-rich habitats, and even adjacent territories can be comprised of very different habitat types. Patchier territories tend to be smaller. Scat analysis shows variation in territory structure is reflected in very varied diets, and that diet tracks the seasonal abundance of different food sources. At small spatial scales foxes can optimize their use of food patches, and also time their activity to coincide with maximum availability. The extraordinary plasticity of fox ecology complicates attempts to manage them, either to protect stock or for disease control.

David W. Macdonald
Patrick Doncaster
Malcolm Newdick
Heribert Hofer
Fiona Mathews
Paul J. Johnson


Albone ES, Gosden PE, Ware GC, Macdonald DW, Hough NG. 1977. Bacterial action and chemical signalling in the red fox (Vulpes vulpes) and other mammals. Pages 78-91. Proceedings of the American Chemical Society Symposium.

Bacon PJ, Macdonald DW. 1980. To control rabies: vaccinate foxes. New Scientist 87:640-645.

Bacon PJ, Macdonald DW. 1981. Habitat and the spread of Rabies. Nature 289:634-635.

Bacon PJ, Macdonald DW. 1981. Habitat classification, fox populations and rabies spread. Ecology IoT, Grange-over-Sands.

Carbone C, Mace GM, Roberts SC, Macdonald DW. 1999. Energetic constraints on the diet of terrestrial carnivores (Addendum). Nature 402:442-442.

Carr GM, Macdonald DW. 1986. The sociality of solitary foragers: a model based on resource dispersion. Animal Behaviour 34:1540-1549.

Creel SR, Macdonald DW. 1995. Sociality, group size, and reproductive suppression among carnivores. Advances in the Study of Behaviour 24:203-257.

Doncaster CP, Dickman CR, Macdonald DW. 1990. Feeding ecology of red foxes (Vulpes vulpes) in the city of Oxford, England. Journal of Mammalogy 71:188-194.

Doncaster CP, Macdonald DW. 1991. Drifting territoriality in the red fox Vulpes vulpes. Journal of Animal Ecology 60:423-439.

Doncaster CP, Macdonald DW. 1991. Ecology and ranging behaviour of red foxes in the city of Oxford. Atti Io Simp. Carnivori (Univ. Pavia), Hystrix 3:11-20.

Doncaster CP, Macdonald DW. 1992. Optimum group size for defending heterogenous distributions of resources: a model applied to red foxes, Vulpes vulpes, in Oxford City. Journal of Theoretical Biology 159:189-198.

Doncaster CP, Macdonald DW. 1996. Intraspecific variation in the movement behaviour of foxes (Vulpes vulpes): a reply to White, Saunders and Harris. Journal of Animal Ecology 65:126-127.

Doncaster CP, Macdonald DW. 1997. Activity patterns and interactions of red foxes (Vulpes vulpes) in Oxford city. Journal of Zoology 241:73-87.

Fenn MGP, Macdonald DW. 1995. Use of middens by red foxes: risk reverses rhythms of rats. Journal of Mammalogy 76:130-136.

Geffen E, Gompper ME, Gittleman JL, Luh HK, Macdonald DW, Wayne RK. 1996. Size, life history traits, and social organization in the Canidae: a re-evaluation. American Naturalist 147:140-160.

Hofer H, Macdonald DW. 1986. Patterns of resource distribution and exploitation by the red fox (Vulpes vulpes) and the Eurasian badger (Meles meles) : a comparative study. Pages 309, [375] leaves, [100] leaves of plates. University of Oxford, 1986.

Johnson D, Macdonald DW, Kays R, Blackwell PG. 2003. Response to Revilla, and Buckley and Ruxton: the resource dispersion hypothesis. Trends in Ecology and Evolution 18:381-382.

Johnson DDP, Kays R, Blackwell PG, Macdonald DW. 2002. Does the resource dispersion hypothesis explain group living? Trends in Ecology & Evolution 17:563-570.

Johnson DDP, Macdonald DW. 2003. Sentenced without trial: reviling and revamping the Resource Dispersion Hypothesis. Oikos 101:433-440.

Macdonald DW. 1976. Food caching by red foxes and some other carnivores. Zeitschrift für Tierpsychologie 42:170-185.

Macdonald DW. 1977. The behavioural ecology of the red fox. Pages 70-90 in Kaplan C, editor. Rabies, the facts. Oxford University Press, Oxford.

Macdonald DW. 1977. The behavioural ecology of the red fox, vulpes vulpes : a study of social organisation and resource exploitation. Page 2 v. University of Oxford, 1977.

Macdonald DW. 1977. On food preference in the red fox. Mammal Review 7:7-23.

Macdonald DW. 1979. “Helpers” in fox society. Nature 282:69-71.

Macdonald DW. 1979. Some observations and field experiments on the urine marking behaviour of the red fox, Vulpes vulpes L. Zeitschrift für Tierpsychologie 51:1-22.

Macdonald DW. 1980. Patterns of scent marking with urine and faeces amongst carnivore communities. Pages 107-139. Proceedings of the Symposia of the Zoological Society of London.

Macdonald DW 1980. Rabies and wildlife: a biologist’s perspective. Oxford University Press, New York.

Macdonald DW. 1980. The red fox, Vulpes vulpes, as a predator upon earthworms, Lumbricus terrestris. Zeitschrift für Tierpsychologie 52:171-200.

Macdonald DW. 1980. Social factors affecting reproduction behaviour among red foxes (Vulpes vulpes L., 1758). Pages 131-183 in Zimen E, editor. The red fox: behaviour and ecology. Dr. W. Junk bv Publishers, The Hague.

Macdonald DW. 1981. Resource dispersion and the social organisation of the red fox (Vulpes vulpes). Pages 918-949 in Chapman JA, and Pursley D, editors. The First International Worldwide Furbearer Conference, Frostburg, Maryland.

Macdonald DW. 1983. The ecology of carnivore social behaviour. Nature 301:379-384.

Macdonald DW. 1984. Carnivore social behaviour – does it need patches? A reply to von Schantz. Nature 307:390-390.

Macdonald DW 1987. Running with the Fox. Unwin Hyman, London.

Macdonald DW 1992. The Velvet Claw. BBC Books, London.

Macdonald DW. 1995. Wildlife rabies: the implications for Britain. Unresolved questions for the control of wildlife rabies: social perturbation and interspecific interactions. Pages 33-48. Rabies in a changing world. Proceedings of the British Small Animal Veterinary Association, Cheltenham.

Macdonald DW. 2012. Red Fox. Pages 70-74 in Kingdon JD, editor. Mammals of Africa. A & C Black.

Macdonald DW. 2013. Genus Suricata. Pages 346-352 in Kingdon JS, and Hoffmann M, editors. Mammals of Africa Vol V. Carnivores, Pangolins, Equids and Rhinoceroses. Bloomsbury, London.

Macdonald DW, Bacon PJ. 1982. Fox Society, Contact Rate and Rabies Epizootiology. Comparative Immunology Microbiology and Infectious Diseases 5:247-256.

Macdonald DW, Barrett P 1995. Collins Field Guide to Mammals of Britain & Europe. HarperCollins, London.

Macdonald DW, Brown LJ, Yerli S, Canbolat A-F. 1994. The behaviour of red foxes, Vulpes vulpes, caching the eggs of loggerhead turtles, Carreta carreta. Journal of Mammalogy 75:985-988.

Macdonald DW, Brown RE, editors. 1985. Social Odours in Mammals. Oxford University Press, Oxford.

Macdonald DW, Buesching C, D., Stopka P, Henderson J, Ellwood SA, Baker SE. 2004. Encounters between two sympatric carnivores: red foxes (Vulpes vulpes) and European badgers (Meles meles). Journal of Zoology 263:385-392.

Macdonald DW, Bunce RGH, Bacon PJ. 1981. Fox Populations, Habitat Characterization and Rabies Control. Journal of Biogeography 8:145-151.

Macdonald DW, Carr G. 1979. A report on the trends in the trade in red fox pelts of British origin.

Macdonald DW, Carr GM. 1989. Food security and the rewards of tolerance. Pages 75-99 in Standen V, and Folley RA, editors. Comparative socioecology: the behavioural ecology of humans and other mammals. Blackwell Scientific Publications, Oxford.

Macdonald DW, Courtenay O, Forbes S, Mathews F. 1999. The red fox (Vulpes vulpes) in Saudi Arabia: loose-knit groupings in the absence of territoriality. Journal of Zoology 249:383-391.

Macdonald DW, Creel S, Mills MG. 2004. Canid society. Pages 85-106 in Macdonald DW, and Sillero-Zubiri C, editors. The biology and conservation of wild canids. Oxford University Press, Oxford, UK.

Macdonald DW, Doncaster P, Royal Society for the Prevention of Cruelty to Animals. 1985. Foxes in your neighbourhood? RSPCA, Horsham.

Macdonald DW, Halliwell EC. 1994. The rapid spread of red foxes, Vulpes vulpes, on the Isle of Man. Global Ecology and Biogeography Letters 4:9-16.

Macdonald DW, Johnson PJ. 1996. The impact of sport hunting: a case study. Pages 160-207 in Dunstone N, and Taylor VL, editors. The exploitation of mammal populations: 1994 Symposium of the Zoological Society of London. Chapman & Hall, London.

Macdonald DW, Newdick MT. 1982. The distribution and ecology of foxes, Vulpes vulpes (L.), in urban areas. Pages 123-135 in Bornkamm R, Lee JA, and Seaward MRD, editors. Urban ecology. Blackwell Scientific Publications, Oxford.

Macdonald DW, Reynolds J. 2004. Red fox (Vulpes vulpes). Pages 129-135 in Sillero-Zubiri C, Hoffmann M, and Macdonald DW, editors. Canids: foxes, wolves, jackals and dogs. Status survey and conservation action plan. Gland, Switzerland, and Cambridge, UK, Gland, Switzerland, and Cambridge, UK, IUCN.

Macdonald DW, Reynolds JC, Carbone C, Mathews F, Johnson PJ. 2003. The bioeconomics of fox control. Pages 220-236 in Tattersall FH, and Manley WJ, editors. Conservation and Conflict: Mammals and Farming in Britain. Linnean Society Occasional Publication, Westbury Publishing, Yorkshire.

Macdonald DW, Riordan P, Mathews F. 2006. Biological hurdles to the control of of TB in cattle: A test of two hypothesis concerning wildlife to explain the failure of control. Biological Conservation 131:268-286.

Macdonald DW, Voigt DR. 1985. The biological basis of rabies models. Pages 71-108 in Bacon PJ, editor. Population dynamics of rabies in wildlife. Academic Press, London.

Newdick MT, Macdonald DW, University of Oxford. Dept. of Zoology., University of Oxford. Life and Environmental Sciences Division. 1983. The behavioural ecology of urban foxes, vulpes vulpes, in Oxford.

Smith GC, Thulke H-H, Fooks AR, Artois M, Macdonald DW, Eisinger D, Selhorst T. 2008. Towards the Elimination of Rabies in Eurasia: What is the future of rabies control in Europe? Pages 283-289 in Dodet B FA, Miller T, Tordo N, editor. Developments in Biologicals.

Voigt DR, Macdonald DW. 1984. Variation in the spatial and social behaviour of the red fox, Vulpes vulpes. Acta Zoologica Fennica 171:261-265.