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Temporal range: Oligocene to Holocene (34 Ma-present) 34–0 Ma
Caninae (Canini, Vulpini, Urocyon).jpg
Major canine clades, represented by a black-backed jackal (a wolf-like canine), a red fox (a vulpine) and a gray fox
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Subfamily: Caninae
Fischer de Waldheim, 1817

The Caninae, known as canines,[6]: 182  are one of three subfamilies found within the canid family. The other two canid subfamilies are the extinct Borophaginae and Hesperocyoninae.[7] The Caninae includes all living canids and their most recent fossil relatives.[1] Their fossils were first found in North America and dated to the Oligocene era, then spreading to Asia at the end of the Miocene era,[6]: 122  some 7 million to 8 million years ago.[7]

Taxonomy and lineage[edit]

Canid subfamilies

Hesperocyoninae Hesperocyon (white background).png

Borophaginae Aelurodon illustration.png

Caninae Dogs jackals wolves and foxes Plate X.png

The genus Leptocyon (Greek: leptos slender + cyon dog) includes 11 species and was the first primitive canine. They were small and weighed around 2 kg.[6]: 53  They first appeared in Sioux County, Nebraska in the Orellan era 34-32 million years ago, which was the beginning of the Oligocene.[1] This was the same time as the appearance of the Borophaginae with whom they share features, indicating that these were two sister groups. Borophaginae skull and dentition were designed for a powerful killing bite compared with the Leptocyon which were designed for snatching small, fast-moving prey. The species L. delicatus is the smallest canid to have existed. At the close of their genus 9 million years ago one Leptocyon lineage resembled the modern fox.[6]: 53  The various species of Leptocyon branched 11.9 Mya into Vulpini (foxes) and Canini (canines).[1]: 174–175 

The canines spent two-thirds of their history in North America, before dispersing 7 million years ago into Asia, Europe, and Africa. One of the characteristics that distinguished them from the Borophaginae and Hesperocyoninae was their possession of less weight in their limbs and more length in their legs, which may have aided their dispersion. The first canine to arrive in Eurasia was the coyote-sized Canis cipio, whose scant fossils were found in Spain. However, the assignment of C. cipio within the canines to the genus Canis or genus Eucyon is not clear.[6]: 143–144 

Phylogenetic relationships[edit]

Skulls of various canine genera; Vulpes (corsac fox), Nyctereutes (raccoon dog), Cuon (dhole), and Canis (Eurasian golden jackal)

The results of allozyme and chromosome analyses have previously suggested several phylogenetic divisions:

Divisions Description Image Genus Species
Canina[8] The wolf-like canines Wolf on alert.jpg Canis Linnaeus, 1758
Dhole (Asiatic wild dog) cropped.jpg Cuon Hodgson, 1838
African wild dog (Lycaon pictus pictus).jpg Lycaon Brookes, 1827
Lupulella mesomelas schmidti Kenya 2020 2 (cropped).jpg Lupulella Hilzheimer, 1906
Cerdocyonina[8] The South American canines Chien des buissons Safari de Peaugres 07082017 2.jpg Speothos Lund, 1839
South american grey fox (cropped).jpg Lycalopex Burmeister 1854
Crab-eating Fox (cropped).JPG Cerdocyon C. E. H. Smith, 1839
Lobo Guará andando.jpg Chrysocyon Smith, 1839
Atelocynus microtis en amazonie péruvienne.jpg Atelocynus Cabrera, 1940
Vulpini[8] The fox-like canines タヌキ - panoramio - Masashi Taomoto (cropped).jpg Nyctereutes Temminck, 1838
  • N. procyonoides
  • N. viverrinus
  • Nyctereutes abdeslami
  • Nyctereutes donnezani
  • Nyctereutes megamastoides
  • Nyctereutes sinensis
  • Nyctereutes tingi
  • Nyctereutes vinetorum
Bat eared fox Kenya crop (cropped).jpg Otocyon S. Müller, 1835
Vulpes vulpes ssp fulvus.jpg Vulpes Garsault, 1764
Urocyon[8] Gray foxes Urocyon cinereoargenteus False Cape State Park.jpg Urocyon Baird, 1857

DNA analysis shows that the first three form monophyletic clades. The wolf-like canines and the South American canines together form the tribe Canini.[9] Molecular data imply a North American origin of living Canidae some 10 Mya and an African origin of wolf-like canines (Canis, Cuon, and Lycaon), with the jackals being the most basal of this group.

The South American clade is rooted by the maned wolf and bush dog, and the fox-like canines by the fennec fox and Blanford's fox. The gray fox and island fox are basal to the other clades; however, this topological difference is not strongly supported.[10]

The cladogram below is based on the phylogeny of Lindblad-Toh (2005)[10] modified to incorporate recent findings on Canis,[11] Vulpes,[12] Lycalopex species,[13] and Dusicyon.[14]


Canis lupus familiaris (domestic dog) Tibetan mastiff (transparent background).png

Canis lupus (gray wolf) Dogs, jackals, wolves, and foxes (Plate I).png

Canis latrans (coyote) Dogs, jackals, wolves, and foxes (Plate IX).png

Canis anthus (African wolf) Dogs, jackals, wolves, and foxes (Plate XI).png

Canis simensis (Ethiopian wolf) Dogs, jackals, wolves, and foxes (Plate VI).png

Canis aureus (golden jackal) Dogs, jackals, wolves, and foxes (Plate X).png

Cuon alpinus (dhole) Dogs, jackals, wolves, and foxes (Plate XLI).png

Lycaon pictus (African wild dog) Dogs, jackals, wolves, and foxes (Plate XLIV).png

Lupulella adusta (side-striped jackal) Dogs, jackals, wolves, and foxes (Plate XIII).png

Lupulella mesomelas (black-backed jackal) Dogs, jackals, wolves, and foxes (Plate XII).png


Speothos venaticus (bush dog) Dogs, jackals, wolves, and foxes (Plate XLIII).png

Chrysocyon brachyurus (maned wolf) Dogs, jackals, wolves, and foxes (Plate VII).png

Dusicyon australis (Falkland Islands wolf)


Lycalopex vetulus (hoary fox) Dogs, jackals, wolves, and foxes (Plate XXXI).png

Lycalopex sechurae (Sechuran fox or Peruvian desert fox)

Lycalopex fulvipes (Darwin's fox)

Lycalopex gymnocercus (pampas fox) Dogs, jackals, wolves, and foxes (Plate XVII).png

Lycalopex griseus (South American gray fox or chilla)

Lycalopex culpaeus (culpeo or Andean fox) Dogs, jackals, wolves, and foxes (Plate XIV).png

Cerdocyon thous (crab-eating fox) Dogs, jackals, wolves, and foxes (Plate XV).png

Atelocynus microtis (short-eared dog) Dogs, jackals, wolves, and foxes (Plate XVI).png


Otocyon megalotis (bat-eared fox) Dogs, jackals, wolves, and foxes BHL19827472 white background.png

Nyctereutes (raccoon dogs) Dogs, jackals, wolves, and foxes (Plate XXXII).png


Vulpes zerda (fennec fox) Dogs, jackals, wolves, and foxes (Plate XXXVI).png

Vulpes cana (Blanford's fox) Blandford's fox.png

Vulpes chama (Cape fox) Dogs, jackals, wolves, and foxes (Plate XXXIII).png

Vulpes vulpes (red fox) Dogs, jackals, wolves, and foxes (Plate XXII).png

Vulpes rueppellii (Ruppell's fox) Dogs, jackals, wolves, and foxes (Plate XXXV).png

Vulpes corsac (corsac fox) Dogs, jackals, wolves, and foxes (Plate XXVII).png

Vulpes ferrilata (Tibetan sand fox) Tibetan sand fox illustration, transparent background.png

Vulpes macrotis (kit fox) Dogs, jackals, wolves, and foxes (Plate XXV).png

Vulpes lagopus (Arctic fox) Dogs, jackals, wolves, and foxes (Plate XXVI).png


Urocyon littoralis (island fox) Vulpes littoralis transparent background.png

Urocyon cinereoargenteus (gray fox) Dogs, jackals, wolves, and foxes (Plate XX).png


  1. ^ a b c d Tedford, Richard; Wang, Xiaoming; Taylor, Beryl E. (2009). "Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae)" (PDF). Bulletin of the American Museum of Natural History. 325: 1–218. doi:10.1206/574.1. hdl:2246/5999. S2CID 83594819.
  2. ^ McKenna, M.C.; Bell, S.K. (1997). Classification of Mammals above the Species Level. Columbia University Press. ISBN 978-0-231-11012-9.
  3. ^ Lyras, G.A.; van der Geer, A.E.; Dermitzakis, M.; de Vos, J. (2006). "Cynotherium sardous, an insular canid (Mammalia: Carnivora) from the Pleistocene of Sardinia (Italy), and its origin". Journal of Vertebrate Paleontology. 26 (3): 735–745. doi:10.1671/0272-4634(2006)26[735:CSAICM]2.0.CO;2.
  4. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
  5. ^ Sotnikova, M. (2006). "A new canid Nurocyon chonokhariensis gen. et sp. nov.(Canini, Canidae, Mammalia) from the Pliocene of Mongolia" (PDF). Courier Forschungsinstitut Senckenberg. 256: 11. Retrieved 4 May 2008.
  6. ^ a b c d e Wang, Xiaoming; Tedford, Richard H. (2008). Dogs: Their Fossil Relatives and Evolutionary History. Columbia University Press, New York. pp. 1–232. ISBN 978-0-231-13529-0.
  7. ^ a b Miklosi, Adam (2015). Dog Behaviour, Evolution, and Cognition. Oxford Biology (2 ed.). Oxford University Press. pp. 103–107. ISBN 978-0199545667 – via Google Books.
  8. ^ a b c d Wayne, Robert K. (June 1993). "Molecular evolution of the dog family". Trends in Genetics. 9 (6): 218–224. doi:10.1016/0168-9525(93)90122-x. PMID 8337763.
  9. ^ Jensen, Per (2007). The Behavioural Biology of Dogs. CABI. pp. 11–13. ISBN 978-1-84593-188-9.
  10. ^ a b Lindblad-Toh, Kerstin; Wade, Claire M.; Mikkelsen, Tarjei S.; Karlsson, Elinor K.; Jaffe, David B.; Kamal, Michael; et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature. 438 (7069): 803–819. Bibcode:2005Natur.438..803L. doi:10.1038/nature04338. PMID 16341006.
  11. ^ Koepfli, Klaus-Peter; Pollinger, John; Godinho, Raquel; Robinson, Jacqueline; Lea, Amanda; Hendricks, Sarah; et al. (2015). "Genome-wide evidence reveals that African and Eurasian Golden Jackals are distinct species". Current Biology. 25 (16): 2158–2165. doi:10.1016/j.cub.2015.06.060. PMID 26234211.
  12. ^ Zhao, Chao; Zhang, Honghai; Liu, Guangshuai; Yang, Xiufeng; Zhang, Jin (2016). "The complete mitochondrial genome of the Tibetan fox (Vulpes ferrilata) and implications for the phylogeny of Canidae". Comptes Rendus Biologies. 339 (2): 68–77. doi:10.1016/j.crvi.2015.11.005. ISSN 1631-0691. PMID 26868757.
  13. ^ Tchaicka, Ligia; de Freitas, Thales Renato Ochotorena; Bager, Alex; Vidal, Stela Luengos; Lucherini, Mauro; Iriarte, Agustín; et al. (2016). "Molecular assessment of the phylogeny and biogeography of a recently diversified endemic group of South American canids (Mammalia: Carnivora: Canidae)" (PDF). Genetics and Molecular Biology. 39 (3): 442–451. doi:10.1590/1678-4685-GMB-2015-0189. PMC 5004827. PMID 27560989.
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