Viverridae

This article is about the extant family of carnivorans. For an extinct family of carnivorous mammals, see Viverravidae.

Viverridae is a family of small to medium-sized feliform mammals, comprising 14 genera with 33 species. This family was named and first described by John Edward Gray in 1821.^[3] Viverrids occur all over Africa, in southern Europe, and in South and Southeast Asia on both sides of the Wallace Line.^[4]

Viverridae[1] Temporal range: 34–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Eocene to Recent[2]
Viverrids (clockwise from top left): species of Paradoxurus, Genetta, Arctictis and Paguma
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Infraorder:	Aeluroidea
Parvorder:	Viverroidea
Family:	Viverridae Gray, 1821
Type genus
Viverra Linnaeus, 1758
Genera
Subfamily Genettinae Genetta Poiana †Semigenetta Subfamily Hemigalinae Hemigalus Chrotogale Cynogale Diplogale Macrogalidia Subfamily Paradoxurinae Paradoxurus Paguma Arctictis Arctogalidia †Kanuites †Kichechia †Siamictis †Tugenictis Subfamily Viverrinae Civettictis Viverra Viverricula
Distribution of living viverrid species

The species of the subfamily Genettinae are known as genets and oyans. The viverrids of the subfamily Viverrinae are commonly called civets; the Paradoxurinae and most Hemigalinae species are called palm civets.

Characteristics

Binturong (Arctictis binturong) on display at the Museum of Osteology

Viverrids have four or five toes on each foot and half-retractile claws. They have six incisors in each jaw and molars with two tubercular grinders behind in the upper jaw, and one in the lower jaw. The tongue is rough with sharp prickles. A pouch or gland occurs beneath the anus, but there is no cecum.^[3] The male's urethral opening is directed backward.^[5]

Viverrids are the most primitive of all the families of feliform Carnivora and clearly less specialized than the Felidae. In external characteristics, they are distinguished from the Felidae by the longer muzzle and tuft of facial vibrissae between the lower jaw bones, and by the shorter limbs and the five-toed hind foot with the first digit present. The skull differs by the position of the postpalatine foramina on the maxilla, almost always well in advance of the maxillopalatine suture, and usually about the level of the second premolar; and by the distinct external division of the auditory bulla into its two elements either by a definite groove or, when rarely this is obliterated, by the depression of the tympanic bone in front of the swollen entotympanic. The typical dental formula is: 3.1.4.23.1.4.2, but the number may be reduced, although never to the same extent as in the Felidae.^[4]

Their flesh-shearing carnassial teeth are relatively undeveloped compared to those of other feliform carnivorans.^[6] Most viverrid species have a penis bone (a baculum).^[7]

Classification

Living species

In 1821, Gray defined this family as consisting of the genera Viverra, Genetta, Herpestes, and Suricata.^[3] Reginald Innes Pocock later redefined the family as containing a great number of highly diversified genera, and being susceptible of division into several subfamilies, based mainly on the structure of the feet and of some highly specialized scent glands, derived from the skin, which are present in most of the species and are situated in the region of the external generative organs. He subordinated the subfamilies Hemigalinae, Paradoxurinae, Prionodontinae, and Viverrinae to the Viverridae.^[4]

In 1833, Edward Turner Bennett described the Malagasy fossa (Cryptoprocta ferox) and subordinated the Cryptoprocta to the Viverridae.^[8] A molecular and morphological analysis based on DNA/DNA hybridization experiments suggests that Cryptoprocta does not belong within Viverridae, but is a member of the Eupleridae.^[9]

The African palm civet (Nandinia binotata) resembles the civets of the Viverridae, but is genetically distinct and belongs in its own monotypic family, the Nandiniidae. There is little dispute that the Poiana species are viverrids.^[1]

DNA analysis based on 29 carnivoran species, comprising 13 Viverrinae species and three species representing Paradoxurus, Paguma and Hemigalinae, confirmed Pocock's assumption that the African linsang Poiana represents the sister group of the genus Genetta. The placement of Prionodon as the sister group of the family Felidae is strongly supported, and it was proposed that the Asiatic linsangs be placed in the monogeneric family Prionodontidae.^[10]

Family Viverridae^[2][1][11]

Subfamily	Genus	Species	Image of type species
Viverrinae	Viverra Linnaeus, 1758[12]	Large Indian civet (V. zibetha) Linnaeus, 1758[12] Malayan civet (V. tangalunga) Gray, 1832[13] Malabar large-spotted civet (V. civettina) Blyth, 1862[14] Large-spotted civet (V. megaspila) Blyth, 1862[14]
	Viverricula Hodgson, 1838[15]	Small Indian civet (V. indica) (Geoffroy Saint-Hilaire, 1803)[16]
	Civettictis Pocock, 1915[17]	African civet (C. civetta) (Schreber, 1776)[18]
Hemigalinae Gray, 1864[19]
	Hemigalus Jourdan, 1837[20]	Banded palm civet (H. derbyanus) Jourdan, 1837[20]
	Cynogale Gray, 1836[21]	Otter civet (C. bennettii) Gray, 1836[21]
	Diplogale Thomas, 1912[22]	Hose's palm civet (D. hosei) (Thomas, 1892)[23]
	Macrogalidia Schwarz, 1910[24]	Sulawesi palm civet (M. musschenbroekii) (Schlegel, 1877)[25]
	Chrotogale Thomas, 1912[22]	Owston's palm civet (C. owstoni) Thomas, 1912[22]
Paradoxurinae Gray, 1864[19]	Paradoxurus Cuvier, 1822[26]	Asian palm civet (P. hermaphroditus) (Pallas, 1777)[27] Golden palm civet (P. zeylonensis) (Pallas, 1778)[28] Brown palm civet (P. jerdoni) Blanford, 1885[29]
	Arctictis Temminck, 1824[30]	Binturong (A. binturong) (Raffles, 1822)[31]
	Paguma Gray, 1831[32]	Masked palm civet (P. larvata) (Smith, 1827)[33]
	Arctogalidia Merriam, 1897[34]	Small-toothed palm civet (A. trivirgata) (Gray, 1832)[35]
Genettinae	Genetta Cuvier, 1816[36]	Common genet (G. genetta) (Linnaeus, 1758)[37] Cape genet (G. tigrina) (Schreber, 1778)[38] South African small-spotted genet (G. felina) (Thunberg, 1811)[39] Rusty-spotted genet (G. maculata) (Gray, 1828)[40] Pardine genet (G. pardina) Geoffroy Saint-Hilaire, 1832[41] Abyssinian genet (G. abyssinica) (Rüppell, 1835)[42] King genet (G. poensis) Waterhouse, 1838[43] Servaline genet (G. servalina) Pucheran, 1855 Angolan genet (G. angolensis) Bocage, 1882 Giant forest genet (G. victoriae) Thomas, 1901[44] Hausa genet (G. thierryi) Matschie, 1902[45] Letaba genet (G. letabae) Thomas and Schwann, 1906[46] Johnston's genet (G. johnstoni) (Pocock, 1908) Aquatic genet (G. piscivora) (Allen, 1919)[47] Crested servaline genet (G. cristata) Hayman, 1940 Schouteden's genet (G. schoutedeni) Crawford-Cabral, 1970 Bourlon's genet (G. bourloni) Gaubert, 2003[48]
	Poiana Gray, 1864[19]	Central African oyan (P. richardsonii) (Thomson, 1842)[49] West African oyan (P. leightoni) (Pocock, 1907)[50]

Phylogeny

The phylogenetic relationships of Viverridae are shown in the following cladogram:^[2][11][51]

Viverridae

Paradoxurinae Paradoxurus Golden palm civet P. zeylonensis Jerdon's palm civet P. jerdoni Asian palm civet P. hermaphroditus Paguma Masked palm civet P. larvata Arctictis Binturong A. binturong Arctogalidia Small-toothed palm civet A. trivirgata Hemigalinae Cynogale Otter civet C. bennettii Macrogalidia Sulawesi palm civet M. musschenbroekii Diplogale Hose's palm civet D. hosei Chrotogale Owston's palm civet C. owstoni Hemigalus Banded palm civet H. derbyanus Viverrinae Viverrinae Viverra Malabar large-spotted civet V. civettina Large-spotted civet V. megaspila Large Indian civet V. zibetha Malayan civet V. tangalunga Civettictis African civet C. civetta Viverricula Small Indian civet V. indica sensu stricto Genettinae Poiana West African oyan P. leightoni Central African oyan P. richardsonii Genetta Abyssinian genet G. abyssinica Haussa genet G. thierryi Giant forest genet G. victoriae Johnston's genet G. johnstoni Aquatic genet G. piscivora Servaline genet G. servalina Crested servaline genet G. cristata South African small-spotted genet G. felina Common genet G. genetta Cape genet G. tigrina Letaba genet G. letabae Schouteden's genet G. schoutedeni Rusty-spotted genet G. maculata Angolan genet G. angolensis Pardine genet G. pardina Bourlon's genet G. bourloni King genet G. poensis sensu lato

Extinct species

Subfamily	Genus	Species
Viverrinae	Viverra Linnaeus, 1758	Leakey's civet (V. leakeyi) Leakey, 1982
	Semigenetta Helbing 1927	†S. cadeoti Roman and Viret 1934 †S. elegans Dehm, 1950 †S. grandis Crusafont & Golpe, 1981 †S. laugnacensis De Bonis, 1973 †S. ripolli Petter, 1976 †S. sansaniensis Lartet, 1851
Paradoxurinae	Kichechia Savage, 1965[52]	†K. zamanae[52] †K. savagei[53]
	Tugenictis Morales & Pickford, 2005[54][55]	†T. ngororaensis[54] Morales & Pickford, 2005
	Kanuites Dehghani & Werdelin, 2008[56]	†K. lewisae[56] Dehghani & Werdelin, 2008
	Siamictis Grohé et al., 2020[57]	†S. carbonensis[57] Grohé et al., 2020

See also

• List of viverrids