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The Caninae, known as canines,[6](p182) 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 have been found in Lower Oligocene North America, and they did not spread to Asia until the end of the Miocene,[6](p122) some 7 million to 8 million years ago.[7] Many extinct species of Caninae were endemic to North America, living from 34 million to 11,000 years ago.[8]

Taxonomy and lineage

Canid subfamilies


Borophaginae +

Hesperocyoninae +

"Derived characteristics that distinguish the Caninae from other canids include small, simple, well-spaced premolars, a humerus without an entepicondylar foramen, and a metatarsal 1 which is reduced to a proximal rudiment."[9]

The genus Leptocyon (Greek: leptos slender + cyon dog) includes 11 species and was the first canine. They were small and weighed around 2 kg. They first appeared in North America around 34 million years ago in the Oligocene at the same time as the Borophaginae with whom they share features with, 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](p53)

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](pp143-144)

Phylogenetic relationships

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:

The wolf-like canines (genus Canis, Cuon, and Lycaon) include the domestic dog (Canis lupus familiaris), gray wolf (Canis lupus), red wolf (Canis rufus), eastern wolf (Canis lycaon), coyote (Canis latrans), Eurasian golden jackal (Canis aureus), African golden wolf (Canis anthus), Ethiopian wolf (Canis simensis), black-backed jackal (Canis mesomelas), side-striped jackal (Canis adustus), dhole (Cuon alpinus), and African wild dog (Lycaon pictus).[10]
The South American canines include the bush dog (Speothos venaticus), hoary fox (Lycalopex uetulus), crab-eating fox (Cerdocyon thous), and maned wolf (Chrysocyon brachyurus).[10]
The fox-like canines include the kit fox (Vulpes velox), red fox (Vulpes vulpes), Cape fox (Vulpes chama), Arctic fox (Vulpes lagopus), fennec fox (Vulpes zerda), the raccoon dog (Nyctereutes procyonoides), and the bat-eared fox (Otocyon megalotis).[10]
A monotypic taxon for the basal California island fox (Urocyon littoralis) and the gray fox (Urocyon cinereoargenteus).[10]

DNA analysis shows that the first three form monophyletic clades. The wolf-like canines and the South American canines together form the tribe Canini.[11] 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.[12]

The cladogram below is based on the phylogeny of Lindblad-Toh et al. (2005),[12] modified to incorporate recent findings on Canis,[13]Vulpes,[14]Lycalopex, species.[15] and Dusicyon[16]


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

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


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

Lycalopex sechurae (Sechuran fox or Peruvian desert fox)

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 procyonoides (raccoon dog) Dogs, jackals, wolves, and foxes (Plate XXXII).png


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



  1. ^ a b 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.
  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 2008.
  6. ^ a b c d Wang, Xiaoming; Tedford, Richard H. (2008). Dogs: Their Fossil Relatives and Evolutionary History. New York, NY: Columbia University Press. 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. ^ "Caninae Basic info". Paleobiology Database.[permanent dead link].
  9. ^ Munthe, Kathleen (1998). "Chapter 7, Canidae". In Janis, C.M.; Scott, K.M.; Jacobs, L.L. (eds.). Evolution of Tertiary mammals of North America. New York, NY: Cambridge University Press. pp. 124-143.
  10. ^ 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.
  11. ^ Jensen, Per (2007). The Behavioural Biology of Dogs. CABI. pp. 11-13. ISBN 978-1-84593-188-9.
  12. ^ 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.
  13. ^ 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.
  14. ^ 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.
  15. ^ 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.
  16. ^ Slater, G. J.; Thalmann, O.; Leonard, J. A.; Schweizer, R. M.; Koepfli, K.-P.; Pollinger, J. P.; et al. (2009). "Evolutionary history of the Falklands wolf". Current Biology. 19 (20): R937-R938. doi:10.1016/j.cub.2009.09.018. ISSN 0960-9822. PMID 19889366.

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