Belongs within: Caninae.
The genus Vulpes contains the standard foxes, found in Eurasia, Africa and North America. Foxes may be solitary or live in small groups of a male and up to five females (Macdonald 1984). The fennec fox V. zerda of northern Africa has sometimes been classified in its own genus Fennecus, but despite its distinctive large ears it is probably not much more divergent overall than other Vulpes species. The most widespread species is the red fox Vulpes vulpes, found across almost the entire Holarctic region (and introduced to Australia), though the North American population is sometimes treated as a separate species V. fulva.
Fantastic Mr Fox
Published 15 September 2008
Foxes are a widespread assemblage of canid predators, found through most of the Holarctic and drier Africa and also here in Australia, where the red fox Vulpes vulpes was introduced quite successfully. Too successfully, in fact—foxes are one of the most significant invasive species in Australia, and a dire threat to many native species. Of the slightly more than ten species in the fox genus Vulpes, the red fox is undoubtedly the most familiar, being both the most widespread species overall as well as the most abundant in developed countries. However, the familiarity of the red fox is a little misleading, as Vulpes vulpes is actually one of the more distinctive species in the genus, being considerably larger and arguably more dog-like than other foxes.
The morphological analysis of the Canidae by Tedford et al. (1995) supported a division of the living members of the family between two lineages, the Vulpini containing Vulpes, and the Canini including Canis (the genus including the domestic dog) and the South American canids. This early division is consistent with the early appearance of fossil species assigned to Vulpes, with V. stenognathus coming from the Late Miocene (Lyras & van der Geer 2003). The majority of Vulpini were included in Vulpes except for the bat-eared fox Otocyon megalotis and the two species of grey foxes Urocyon. Two species in the Vulpes clade, the fennec fox Vulpes zerda or Fennecus zerda and the Arctic fox Alopex lagopus have been regarded as separate genera, but it seems pretty well-established that doing so renders Vulpes paraphyletic (Zrzavý & Řičánková 2004). More recent analyses using both molecular and morphological data have continued to support the Vulpes-Canini division (Zrzavý & Řičánková 2004; Bardeleben et al. 2005), but results differ about the relationships of Vulpes to Urocyon and/or Otocyon, which may fall in a Vulpini clade or may be basal to the Vulpes + Canini split. Still, all recent authors seem to agree that, contrary to many older sources, the grey foxes should not be included in Vulpes.
Relationships within the Vulpes clade are fairly uncertain. Zrzavý & Řičánková (2004) tentatively suggested a division between two major groups that both may or may not be monophyletic, an ‘Afro-Asiatic’ clade and an ‘Holarctic’ clade. The ‘Afro-Asiatic’ group includes the fennec and pale fox V. pallida of northern Africa, the Cape fox V. chama of southern Africa, Blanford’s fox V. cana of central Asia and probably the Indian fox V. bengalensis, with the fennec and Blanford’s foxes forming a clade. Within the ‘Holarctic’ group, Rüppell’s fox V. ruppelli inhabits northern Africa, the red fox can be found across the entire Holarctic, and the corsac V. corsac and Tibetan foxes V. ferrilata are found in central Asia. The circumpolar Arctic fox forms a clade in the Holarctic group with the V. velox/V. macrotis complex, the swift and kit foxes, of North America. All fox species seem to inhabit temperate or dry climates—note particularly the wide geographical division between the Cape fox and all other species of the genus.
While foxes are generally characterised as solitary animals, and certainly do not form packs in the manner of Canis and closely related genera, individuals of at least some species may form small groups, usually a male and a number of vixens. Members of a group will still forage for food separately (Macdonald 1984). All foxes use a characteristic high pounce in capturing prey, springing upwards and landing on their quarry from directly above it.
Of course, foxes are also famed for being one of the few animals able to transform their appearance at will. They share this ability with the tanuki Nyctereutes procyonoides, a canid whose phylogenetic position relative to the Vulpes-Canini split remains uncertain. Nevertheless, the foxes would doubtless like to point out that they are not as prone to buffoonery as tanuki. Because of the uncertain position of the tanuki and a shocking shortage of studies of shape-changing abilities in fox species other than V. vulpes, we cannot presently comment whether the shape-changing ability is a plesiomorphy of crown canids that has been lost in the Canini, or has been acquired independently in foxes and tanuki.
Systematics of Vulpes
Characters (from Macdonald 1984): Muzzle pointed; ears triangular and erect; tail long and bushy; skull relatively flattened. Tail tip often differently coloured from remainder of body; black triangular face marks between eyes and nose. Frontal bones forming slightly indented or dished brows.
Vulpes Bowdich 1821 (nom. cons.) [incl. Fennecus Desmarest 1804]VG78
|--V. zerda (Zimmermann 1780)FS15, VG78 [=Canis zerdaVG78, Fennecus zerdaM84]
`--+--V. cana (Blandford 1877)FS15, VG78 [=Canis (Vulpes) canaVG78]
`--+--+--+--V. chama (Smith 1833)FS15, VG78 [=Canis chamaVG78]
| | `--V. pallida (Cretzschmar 1826)FS15, VG78 [=Canis pallidaVG78]
| `--+--V. velox (Say 1823)FS15, B75 [=Canis veloxVG78]
| | |--V. v. veloxM84
| | `--V. v. hebesM84
| `--+--V. lagopus (Linnaeus 1758)WT08 [=Canis lagopusVG78, Alopex lagopusFS15]
| `--V. macrotis Merriam 1888FS15, WT08
| |--V. m. macrotisMB86
| |--V. m. deviaBP87
| |--V. m. muticaBP87
| `--V. m. zinseri Benson 1938MB86
`--+--+--V. rueppelli (Schinz 1825)FS15, VG78 [=Canis rueppelliVG78]
| `--V. vulpes (Linnaeus 1758)FS15, M76 [=Canis vulpesVG78]
| |--V. v. vulpesB75
| |--V. v. fulvaB75
| `--V. v. regalis Merriam 1900B75
`--+--V. bengalensis (Shaw 1800)FS15, VG78 [=Canis bengalensisVG78]
`--+--V. corsac (Linnaeus 1768)FS15, VG78 [=Canis corsacVG78]
`--V. ferrilata (Hodgson 1842)FS15, VG78 [=Canis (Vulpes) ferrilataVG78]
Vulpes incertae sedis:
V. alopecoides (Major 1873)WT08
V. angustidens Thenius 1954WT08
V. beihaiensis Qiu & Tedford 1990WT08
V. chikushanensis Young 1930WT08
V. flavescensT66
V. galaticus Ginsburg 1998WT08
V. kernensisWT08
V. kiyomasai Kishida & Mori 1929TYM08
V. niloticusT66
V. pattisoniS78
V. praecorsac Kormos 1932WT08
V. praeglacialis Kormos 1932WT08 [=Alopex praeglacialisME05]
V. pulcherS78
V. riffautae Bonis et al. 2007WT08
V. stenognathus Savage 1942WT08
V. vetusWT08
V. vulgarisT66
Inorganic: Fennecus zerda minilorientalus Okamura 1987O87*Type species of generic name indicated
References
Bardeleben, C., R. L. Moore & R. K. Wayne. 2005. A molecular phylogeny of the Canidae based on six nuclear loci. Molecular Phylogenetics and Evolution 37 (3): 815–831.
[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.
[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.
[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.
Lyras, G. E., & A. A. E. van der Geer. 2003. External brain anatomy in relation to the phylogeny of Caninae (Carnivora: Canidae). Zoological Journal of the Linnean Society 138 (4): 505–522.
[M84] Macdonald, D. W. 1984. Foxes. In: Macdonald, D. (ed.) All the World’s Animals: Carnivores pp. 60–67. Torstar Books Inc.: New York.
[M76] Masui, M. 1976. Nihon no Doobutsu. Kogakukan: Tokyo.
[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.
[ME05] Moullé, P.-E., A. Echassoux, F. Lacombat, E. Desclaux & S. Bailon. 2005. L’environnement animal des premiers habitants de l’Europe méditerranéenne: les grands mammifères contemporains de l’homme du Vallonnet, données taxonomiques et biostratigraphiques pour la deuxième moitie du Pléistocène inférieur. BAR International Series 1364: 105-113.
[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.
[S78] Savage, R. J. G. 1978. Carnivora. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 249–267. Harvard University Press: Cambridge (Massachusetts).
Tedford, R. H., B. E. Taylor & X. Wang. 1995. Phylogeny of the Caninae (Carnivora: Canidae): the living taxa. American Museum Novitates 3146: 1–37.
[TYM08] Tennent, W. J., M. Yasuda & K. Morimoto. 2008. Lansania Journal of arachnology and zoology—a rare and obscure Japanese natural history journal. Archives of Natural History 35 (2): 252–280.
[T66] Tristram, H. B. 1866. Report on the mammals of Palestine. Proceedings of the Zoological Society of London 1866: 84–93.
[VG78] Van Gelder, R. G. 1978. A review of canid classification. American Museum Novitates 2646: 1–10.
[WT08] Wang, X., & R. H. Tedford. 2008. Dogs: Their fossil relatives and evolutionary history. Columbia University Press: New York.
Zrzavý, J., & V. Řičánková. 2004. Phylogeny of Recent Canidae (Mammalia, Carnivora): relative reliability and utility of morphological and molecular datasets. Zoologica Scripta 33 (4): 311–333.