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Canines
Temporal range: Oligocene to Holocene (34Ma-present)
Caninae (Canini, Vulpini, Urocyon).jpg
Major canine clades, represented by a black-backed jackal (a true dog), 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
Genera
  • Leptocyon
  • Urocyon
  • Tribe Canini
    • Subtribe Canina
      • Canis
        • Xenocyon
      • Cuon
      • Lycaon
      • Cynotherium
      • Eucyon
    • Subtribe Cerdocyonina
      • Atelocynus
      • Cerdocyon
      • Chrysocyon
      • Lycalopex
      • Speothos
      • Dusicyon
      • Nurocyon
      • Protocyon
      • Theriodictis
  • Tribe Vulpini
    • Metalopex Tedford, Wang, & Taylor 2008
    • Nyctereutes
    • Otocyon
    • Prototocyon
    • Vulpes

The Caninae, known as canines, are one of three subfamilies found within the canid family. The other two canid subfamilies are the extinct Borophaginae and Hesperocyoninae. The Caninae includes all living canids and their most recent fossil relatives. 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, some 7 million to 8 million years ago.

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."

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

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.

Phylogenetic relationships

MSU V2P1a - Vulpes, Nyctereutes, Cuon & Canis skulls
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 (Canis mesomelas), side-striped jackal (Canis adustus), dhole (Cuon alpinus), and African wild dog (Lycaon pictus).
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).
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).
Urocyon
A monotypic taxon for the basal California island fox (Urocyon littoralis) and the gray fox (Urocyon cinereoargenteus).

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

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

 Caninae 

 Canini 
Canina








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 golden 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





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



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




 Cerdocyonina 


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 

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






 Vulpini 

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 


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 

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



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





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