Caninae

Canines; Temporal range: Oligocene to Holocene (34 Ma-present) 34–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	Major canid clades, represented by a black-backed jackal (a wolf-like canine), a red fox (a vulpine) and a gray fox
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Family:	Canidae
Subfamily:	Caninae; Fischer de Waldheim, 1817
Genera
	†Leptocyon; Tribe Urocyonini †Metalopex Tedford, Wang, & Taylor 2008; Urocyon; ; Tribe Canini Subtribe Canina Canis †Xenocyon; ; Cuon; Lupulella; Lycaon; †Cynotherium; †Aenocyon; †Eucyon; †Mececyon; †Megacyon; ; Subtribe Cerdocyonina Atelocynus; Cerdocyon; Chrysocyon; Lycalopex; Speothos; †Dusicyon; †Nurocyon; †Protocyon; †Theriodictis; ; ; Tribe Vulpini Nyctereutes; Otocyon; Vulpes; †Prototocyon; ;

Caninae (whose members are known as canines (/keɪnaɪnz/)^[6]^: 182 is the only living subfamily within Canidae, alongside the extinct Borophaginae and Hesperocyoninae.^[7]^[1] They first appeared in North America, during the Oligocene around 35 million years ago, subsequently spreading to Asia and elsewhere in the Old World at the end of the Miocene,^[6]^: 122 some 7 million to 8 million years ago.^[7]

Taxonomy and lineage

Canid subfamilies

Canidae

†Hesperocyoninae

	†Borophaginae

	Caninae

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

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

Divisions	Description	Genus	Species
Subtribe Canina^[8]	The wolf and wolf-like canines	Canis Linnaeus, 1758	Canis aureus Canis familiaris Canis latrans Canis lupaster Canis lupus Canis rufus Canis lycaon Canis simensis † Canis antonii † Canis armbrusteri † Canis chihliensis † Canis edwardii † Canis etruscus † Canis falconeri † Canis mosbachensis † Canis palmidens † Canis variabilis
		Cuon Hodgson, 1838	Cuon alpinus
		Lycaon Brookes, 1827	Lycaon pictus †Lycaon sekowei
		Lupulella Hilzheimer, 1906	Lupulella adusta Lupulella mesomelas
Subtribe Cerdocyonina^[8]	The South American canines	Speothos Lund, 1839	Speothos venaticus †Speothos pacivorus
		Lycalopex Burmeister 1854	Lycalopex culpaeus Lycalopex fulvipes Lycalopex griseus Lycalopex gymnocercus Lycalopex sechurae Lycalopex vetulus
		Cerdocyon C. E. H. Smith, 1839	Cerdocyon thous
		Chrysocyon Smith, 1839	Chrysocyon brachyurus
		Atelocynus Cabrera, 1940	Atelocynus microtis
Tribe Vulpini^[8]	The fox-like canines	Nyctereutes Temminck, 1838	N. procyonoides N. viverrinus †Nyctereutes abdeslami †Nyctereutes donnezani †Nyctereutes megamastoides †Nyctereutes sinensis †Nyctereutes tingi †Nyctereutes vinetorum
		Otocyon S. Müller, 1835	Otocyon megalotis
		Vulpes Garsault, 1764	Vulpes bengalensis Vulpes cana Vulpes chama Vulpes corsac Vulpes ferrilata Vulpes lagopus Vulpes macrotis Vulpes pallida Vulpes rueppellii Vulpes velox Vulpes vulpes Vulpes zerda
Genus Urocyon^[8]	Gray foxes	Urocyon Baird, 1857	U. cinereoargenteus U. littoralis † U. citrinus † U. galushai † U. minicephalus † U. progressus † U. webbi

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]

Caninae

Canini

Canina

Canis latrans (coyote)

	Canis rufus (red wolf)

	Canis lycaon (algonquin wolf)

	Canis lupus (gray wolf)

	Canis familiaris (domestic dog)

Canis anthus (African wolf)

Canis simensis (Ethiopian wolf)

Canis aureus (golden jackal)

Cuon alpinus (dhole)

Lycaon pictus (African wild dog)

	Lupulella adusta (side-striped jackal)

	Lupulella mesomelas (black-backed jackal)

Cerdocyonina

Speothos venaticus (bush dog)

	Chrysocyon brachyurus (maned wolf)

	†Dusicyon australis (Falkland Islands wolf)

Lycalopex

Lycalopex vetulus (hoary fox)

Lycalopex sechurae (Sechuran fox or Peruvian desert fox)

	Lycalopex fulvipes (Darwin's fox)

	Lycalopex gymnocercus (pampas fox)

	Lycalopex griseus (South American gray fox or chilla)

	Lycalopex culpaeus (culpeo or Andean fox)

Cerdocyon thous (crab-eating fox)

Atelocynus microtis (short-eared dog)

Vulpini

Otocyon megalotis (bat-eared fox)

Nyctereutes (raccoon dogs)

Vulpes

	Vulpes zerda (fennec fox)

	Vulpes cana (Blanford's fox)

Vulpes chama (Cape fox)

	Vulpes vulpes (red fox)

	Vulpes rueppellii (Ruppell's fox)

	Vulpes corsac (corsac fox)

	Vulpes ferrilata (Tibetan sand fox)

	Vulpes macrotis (kit fox)

	Vulpes lagopus (Arctic fox)

Urocyon

	Urocyon littoralis (island fox)

	Urocyon cinereoargenteus (gray fox)

References

^ ^a ^b ^c ^d Tedford, Richard; Wang, Xiaoming; Taylor, Beryl E. (2009). "Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae)". Bulletin of the American Museum of Natural History. 325: 1–218. doi:10.1206/574.1. hdl:2246/5999. S2CID 83594819.
^ McKenna, M.C.; Bell, S.K. (1997). Classification of Mammals above the Species Level. Columbia University Press. ISBN 978-0-231-11012-9.
^ 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. S2CID 84448363.
^ 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.
^ 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.
^ ^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.
^ ^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.
^ ^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.
^ Jensen, Per (2007). The Behavioural Biology of Dogs. CABI. pp. 11–13. ISBN 978-1-84593-188-9.
^ ^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.
^ 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.
^ 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.
^ 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.
^ 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. hdl:10261/58562. ISSN 0960-9822. PMID 19889366. S2CID 36185744.

[Tedford2009-1] Tedford, Richard; Wang, Xiaoming; Taylor, Beryl E. (2009). "Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae)". Bulletin of the American Museum of Natural History. 325: 1–218. doi:10.1206/574.1. hdl:2246/5999. S2CID 83594819.

[McKennaBell1997-2] McKenna, M.C.; Bell, S.K. (1997). Classification of Mammals above the Species Level. Columbia University Press. ISBN 978-0-231-11012-9.

[Lyras-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. S2CID 84448363.

[msw3-4] 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.

[Sotnikova2006-5] 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.

[wang2008-6] 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.

[miklosi2015-7] Miklosi, Adam (2015). Dog Behaviour, Evolution, and Cognition. Oxford Biology (2 ed.). Oxford University Press. pp. 103–107. ISBN 978-0199545667 – via Google Books.

[wayne1993-8] 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.

[LindbladToh2005-10] 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.

[Koepfli2015-11] 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.

[ZhaoZhang2016-12] 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.

[14] 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. hdl:10261/58562. ISSN 0960-9822. PMID 19889366. S2CID 36185744.

[6]