Caninae

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.[5] 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](p122) some 7 million to 8 million years ago.[7]

Canines
Temporal range: Oligocene to Holocene (34 Ma-present)
Major canine clades, represented by a black-backed jackal (a true dog), a red fox (a vulpine) and a gray fox
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Subfamily: Caninae
Fischer de Waldheim, 1817
Genera[1][2][3][4]

Taxonomy and lineage
Canid subfamilies
 Canidae 
 

†Hesperocyoninae

 
 
 

†Borophaginae

 
 

Caninae

 
 
 

"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."[8]

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](p53) They first appeared in Sioux County, Nebraska in the Orellan era 34-32 million years ago, which was the beginning of the Oligocene.[5] 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](p53) Leptocyons various species branched 11.9 Mya into Vulpini (foxes) and Canini (canines).[5](pp174–175)

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](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:

Canina
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 (Lupulella mesomelas), side-striped jackal (Lupulella adusta), dhole (Cuon alpinus), and African wild dog (Lycaon pictus).[9]
Cerdocyonina
The South American canines include the bush dog (Speothos venaticus), hoary fox (Lycalopex uetulus), crab-eating fox (Cerdocyon thous), and maned wolf (Chrysocyon brachyurus).[9]
Vulpini
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).[9]
Urocyon
A monotypic taxon for the basal California island fox (Urocyon littoralis) and the gray fox (Urocyon cinereoargenteus).[9]

DNA analysis shows that the first three form monophyletic clades. The wolf-like canines and the South American canines together form the tribe Canini.[10] 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.[11]

The cladogram below is based on the phylogeny of Lindblad-Toh (2005)[11] modified to incorporate recent findings on Canis,[12] Vulpes,[13] Lycalopex species,[14] and Dusicyon.[15]

 Caninae 
 
 Canini 
Canina
 
 
 
 
 
 
 
 

Canis lupus familiaris (domestic dog)

 
 

Canis lupus (gray wolf)

 
 
 

Canis latrans (coyote)

 
 
 

Canis anthus (African golden wolf)

 
 
 

Canis simensis (Ethiopian wolf)

 
 
 

Canis aureus (golden jackal)

 
 
 

Cuon alpinus (dhole)

 
 
 

Lycaon pictus (African wild dog)

 
 
 
 

Lupulella adustus (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 procyonoides (raccoon dog)

 
 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
  1. McKenna, M.C.; Bell, S.K. (1997). Classification of Mammals above the Species Level. Columbia University Press. ISBN 978-0-231-11012-9.
  2. 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.
  3. 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.
  4. 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.
  5. 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.
  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.
  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.
  8. 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.
  9. 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.
  10. Jensen, Per (2007). The Behavioural Biology of Dogs. CABI. pp. 11–13. ISBN 978-1-84593-188-9.
  11. 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.
  12. 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.
  13. 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.
  14. 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.
  15. 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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