Caninae

Life-forms

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Cladus: Synapsida
Cladus: Eupelycosauria
Cladus: Sphenacodontia
Cladus: Sphenacodontoidea
Cladus: Therapsida
Cladus: Theriodontia
Cladus: Cynodontia
Cladus: Eucynodontia
Cladus: Probainognathia
Cladus: Prozostrodontia
Cladus: Mammaliaformes
Classis: Mammalia
Cladus: Theriimorpha
Cladus: Theriiformes
Cladus: Trechnotheria
Cladus: Zatheria
Subclassis: Theria
Cladus: Eutheria
Infraclassis: Placentalia
Magnordo: Boreoeutheria
Superordo: Laurasiatheria
Cladus: Scrotifera
Grandordo: Ferungulata
Mirordo: Ferae
Cladus: Pancarnivora
Cladus: Carnivoramorpha
Cladus: Carnivoraformes
Ordo: Carnivora
Subordo: Caniformia

Familia: Canidae
Subfamilia: Caninae
Genera: Atelocynus - Canis - Cerdocyon - Chrysocyon - Cuon - Lupulella - Lycalopex - Lycaon - Nyctereutes - Otocyon - Speothos - Urocyon - Vulpes - †Cynotherium - †Dusicyon - †Eucyon - †Leptocyon - †Nurocyon - †Xenocyon

Name

Caninae Fischer, 1817
References

Hartstone-Rose, A.; Werdelin, L.; Ruiter, D.J., de; Berger, L.R.; Churchill, S.E. 2010: The Plio-Pleistocene ancestor of wild dogs, Lycaon sekowei n. sp. Journal of paleontology, 84: 299–308. DOI: 10.1666/09-124.1
Tedford, R.H.; Wang, X.; Taylor, B.E. 2009: Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae). Bulletin of the American Museum of Natural History, (325) DOI: 10.1206/574.1

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	Image	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]

References

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

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