Viverravidae Temporal range: 66.043–33.9 Ma PreꞒ Ꞓ O S D C P T J K Pg N early Paleocene - late Eocene

skull of Viverravus minutus

skull of Didymictis protenus
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Pan-Carnivora
Clade:	Carnivoramorpha
Superfamily:	†Viverravoidea Wortman & Matthew, 1899
Family:	†Viverravidae Wortman & Matthew, 1899^[1]
Type genus
†Viverravus Marsh, 1872
Genera
[see classification]
Synonyms
synonyms of family: Didymictida (Kretzoi, 1945)^[2] Didymictidae (Flynn & Galiano, 1982)^[3] Viverravidea Viverravinae (Matthew, 1909)^[4]

Viverravidae ("ancestors of viverrids") is an extinct monophyletic family of mammals from extinct superfamily Viverravoidea within the clade Carnivoramorpha, that lived from the early Palaeocene to the late Eocene in North America, Europe and Asia.^[5]^[6]^[7] They were once thought to be the earliest carnivorans and ancestral to extant ones, but now are placed outside the order Carnivora based on cranial morphology as relatives (a plesion-group) to extant carnivorans.^[8]^[9]

General characteristics

Wang and Tedford propose that they arose in North America 66–60 million years ago, spread to Asia then later to Europe, and were the first carnivoramorphans and possessed the first true pair of carnassial teeth.^[10] In viverravids, the skull is elongated and the number of molars is reduced to two (M1/m1 and M2/m2 are present and M3/m3 are absent).

Classification and phylogeny

Classification

Taxonomy retrieved from the Paleobiology Database^[11]

Superfamily: †Viverravoidea (Wortman & Matthew, 1899)
Family: †Viverravidae (Wortman & Matthew, 1899)

Genus: †Orientictis (Huang & Zheng, 2005)
- †Orientictis spanios (Huang & Zheng, 2005)
Genus: †Pappictidops (Qiu & Li, 1977)
- †Pappictidops acies (Wang, 1978)
- †Pappictidops obtusus (Wang, 1978)
- †Pappictidops orientalis (Qiu & Li, 1977)
Genus: †Preonictis (Tong & Wang, 2006)
- †Preonictis youngi (Tong & Wang, 2006)
Genus: †Variviverra (Tong & Wang, 2006)
- †Variviverra vegetatus (Tong & Wang, 2006)
Incertae sedis:
- †Viverravidae sp. [CM 71188 & CM 71189] (Friscia & Rasmussen, 2010)
Subfamily: †Didymictinae (Flynn & Galiano, 1982)
- Genus: †Bryanictis (MacIntyre, 1966)
  - †Bryanictis microlestes (Simpson, 1935)
  - †Bryanictis paulus (Meehan & Wilson, 2002)
  - †Bryanictis terlinguae (Standhardt, 1986)
- Genus: †Didymictis (Cope, 1875)
  - †Didymictis altidens (Cope, 1880)
  - †Didymictis dellensis (Dorr, 1952)
  - †Didymictis leptomylus (Cope, 1880)
  - †Didymictis protenus (Cope, 1874)
  - †Didymictis proteus (Polly, 1997)
  - †Didymictis vancleveae (Robinson, 1966)
  - †Didymictis sp. [Erquelinnes, Hainaut, Belgium] (Dollo, 1909)
- Genus: †Intyrictis (Gingerich & Winkler, 1985)
  - †Intyrictis vanvaleni (MacIntyre, 1966)
- Genus: †Pristinictis (Fox & Youzwyshyn, 1994)
  - †Pristinictis connata (Fox & Youzwyshyn, 1994)
- Genus: †Protictis ^{(paraphyletic genus)} (Matthew, 1937)
  - †Protictis agastor (Gingerich & Winkler, 1985)
  - †Protictis haydenianus (Cope, 1882)
  - †Protictis minor (Meehan & Wilson, 2002)
  - †Protictis paralus (Holtzman, 1978)
  - †Protictis simpsoni (Meehan & Wilson, 2002)
  - Subgenus: †Protictoides (Flynn & Galiano, 1982)
    - †Protictis aprophatos (Flynn & Galiano, 1982)
- Genus: †Raphictis (Gingerich & Winkler, 1985)
  - †Raphictis gausion (Gingerich & Winkler, 1985)
  - †Raphictis iota (Scott, 2008)
  - †Raphictis machaera (Rankin, 2009)
  - †Raphictis nanoptexis (Rankin, 2009)
- Incertae sedis:
  - †"Deltatherium" durini (Van Valen, 1978)

Subfamily: †Ictidopappinae (Van Valen, 1969)
- Genus: †Ictidopappus (Simpson, 1935)
  - †Ictidopappus mustelinus (Simpson, 1935)
Subfamily: †Viverravinae (Wortman & Matthew, 1899)
- Genus: †Simpsonictis (MacIntyre, 1962)
  - †Simpsonictis jaynanneae (Rigby, 1980)
  - †Simpsonictis pegus (Gingerich & Winkler, 1985)
  - †Simpsonictis tenuis (Simpson, 1935)
- Genus: †Viverravus (Marsh, 1872)
  - †Viverravus acutus (Matthew & Granger, 1915)
  - †Viverravus gracilis (Marsh, 1872)
  - †Viverravus lawsoni (Hooker, 2010)
  - †Viverravus laytoni (Gingerich & Winkler, 1985)
  - †Viverravus lutosus (Gazin, 1952)
  - †Viverravus minutus (Wortman, 1901)
  - †Viverravus politus (Matthew & Granger, 1915)
  - †Viverravus rosei (Polly, 1997)
  - †Viverravus sicarius (Matthew, 1909)
  - †Viverravus sp. [V11141] (Meng, 1998)
  - †Viverravus sp. [Locality Group 2, Washakie Basin, Wyoming] (Tomiya, 2021)
- Genus: †Viverriscus (Beard & Dawson, 2009)
  - †Viverriscus omnivorus (Beard & Dawson, 2009)

Phylogeny

The phylogenetic relationships of family Viverravidae are shown in the following cladogram:^[3]^[12]^[13]^[14]^[15]

Carnivoramorpha

Carnivoraformes

?

†Carnivoramorpha sp. (UALVP 31176)

?

†Carnivoramorpha sp. (USNM 538395)

?

†"Sinopa" insectivorus

†Viverravidae

?

†Ravenictis

?

†Carnivoramorpha sp. (UALVP 50993 & UALVP 50994)

†Viverravoidea

†Viverravidae

†Pappictidops

	†Pappictidops acies

	†Pappictidops obtusus

†Pappictidops orientalis

†Orientictis

†Orientictis spanios

†Preonictis

†Preonictis youngi

†Variviverra

†Variviverra vegetatus

†Ictidopappinae

†Ictidopappus	†Ictidopappus mustelinus

†Viverravinae

†Viverravus

	†Viverravus minutus

	†Viverravus sicarius

	†Viverravus sp. (Locality Group 2, Washakie Basin, Wyoming)

†Viverravus gracilis

†Viverravus lutosus

†Viverravus acutus

†Viverravus rosei

†Viverravus laytoni

?

†Viverravus sp. (V11141)

†Viverravus lawsoni

†Viverravus politus

†Viverriscus

†Viverriscus omnivorus

†Simpsonictis

	†Simpsonictis jaynanneae

	†Simpsonictis pegus

	†Simpsonictis tenuis

?

†Viverravidae sp. (CM 71188 & CM 71189)

†Didymictinae

†Bryanictis

	†Bryanictis microlestes

	†Bryanictis paulus

	†Bryanictis terlinguae

†Intyrictis

†Intyrictis vanvaleni

†Protictis

†Protictis haydenianus

†Protictis minor

†Protictis simpsoni

†Raphictis

	†Raphictis gausion

	†Raphictis iota

	†Raphictis machaera

	†Raphictis nanoptexis

†Protictis paralus

†Protictis agastor

†(Protictoides)

†Protictis aprophatos

†Pristinictis

†Pristinictis connata

?

†"Deltatherium" durini

†Didymictis

†Didymictis dellensis

†Didymictis proteus

?

†Didymictis sp. (Erquelinnes, Hainaut, Belgium)

	†Didymictis altidens

	†Didymictis leptomylus

	†Didymictis protenus

	†Didymictis vancleveae

sensu stricto

sensu lato

References

^ Wortman, J. L.; Matthew, W. D. (1899). "The ancestry of certain members of the Canidae, Viverridae, and Procyonidae". Bulletin of the American Museum of Natural History. 12: 109–138. hdl:2246/1535. OCLC 46687698.
^ Miklos Kretzoi (1945) "Bemerkungen über das Raubtiersystem." Annales Historico-Naturales Musei Nationalis Hungarici, Budapest, vol. 38, pp. 59–83.
^ ^a ^b J. J. Flynn and H. Galiano. (1982.) "Phylogeny of Early Tertiary Carnivora, With a Description of a New Species of Protictis From the Middle Eocene of Northwestern Wyoming" American Museum Novitates 2725:1-64
^ W. D. Matthew (1909) "The Carnivora and Insectivora of the Bridger Basin, middle Eocene." Memoirs of the American Museum of Natural History 9:289-567
^ "†family Viverravidae (Wortman & Matthew, 1899) (placental)". Fossilworks. Gateway to the Paleobiology Database. Retrieved 29 June 2019.
^ Malcolm C. McKenna, Susan K. Bell: Classification of Mammals: Above the Species Level in Columbia University Press, New York (1997), 631 Seiten. Viverravidae
^ Zack, Shawn P. (2012). "Deciduous dentition of Didymictis (Carnivoramorpha: Viverravidae): implications for the first appearance of "Creodonta"". Journal of Mammalogy. 93 (3): 808–817. doi:10.1644/11-MAMM-A-245.1. ISSN 0022-2372.
^ Wesley-Hunt, G. D.; Flynn, J. J. (2005). "Phylogeny of the Carnivora: basal relationships among the carnivoramorphans, and assessment of the position of 'Miacoidea' relative to Carnivora". Journal of Systematic Palaeontology. 3: 1–28. doi:10.1017/S1477201904001518. S2CID 86755875.
^ Polly, David, Gina D. Wesley-Hunt, Ronald E. Heinrich, Graham Davis and Peter Houde (2006). "Earliest known carnivoran auditory bulla and support for a recent origin of crown-clade carnivora (Eutheria, Mammalia)" (PDF). Palaeontology. 49 (5): 1019–1027. doi:10.1111/j.1475-4983.2006.00586.x.((cite journal)): CS1 maint: multiple names: authors list (link)
^ Wang, Xiaoming; Tedford, Richard H. (2008.) "Dogs: Their Fossil Relatives and Evolutionary History." New York: Columbia University Press
^ Paleobiology Database. Retrieved with Fossilworks (March 3, 2017)
^ P. D. Gingerich and D. A. Winkler. (1985.) "Systematics of Paleocene Viverravidae (Mammalia, Carnivora) in the Bighorn Basin and Clark's Fork Basin, Wyoming." Contributions from the Museum of Paleontology, University of Michigan 27(4):87-128
^ P. D. Polly. (1997.) "Ancestry and Species Definition in Paleontology: A Stratocladistic Analysis of Paleocene-Eocene Viverravidae (Mammalia, Carnivora) from Wyoming." Contributions from the Museum of Paleontology, University of Michigan 30(1):1-53
^ Solé, Floréal; Smith, Thierry; De Bast, Eric; Codrea, Vlad; Gheerbrant, Emmanuel (2016). "New carnivoraforms from the latest Paleocene of Europe and their bearing on the origin and radiation of Carnivoraformes (Carnivoramorpha, Mammalia)". Journal of Vertebrate Paleontology. 36 (2): e1082480. doi:10.1080/02724634.2016.1082480. ISSN 0272-4634. S2CID 87537565.
^ S. Faurby, L. Werdelin, A. Antonelli (2019.) "Dispersal ability predicts evolutionary success among mammalian carnivores" Department of Biological and Environmental Sciences, University of Gothenburg, Box 461, SE

Mammals of clade Pan-Carnivora

Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Infraclass: Placentalia
Superorder: Laurasiatheria
Clade: Scrotifera
Grandorder: Ferungulata
Mirorder: Ferae

Pan-Carnivora

Carnivoramorpha	See Carnivoramorpha below ↓
†Hyaenodonta	See Hyaenodonta below ↓
†Oxyaenodonta	See Oxyaenodonta below ↓
†Wyolestidae	†Wyolestes
†Simidectes †Altacreodus †Tinerhodon

†Hyaenodonta

†Hyaenodontoidea

†Hyaenodontidae	†Alienetherium †Boritia †Cartierodon †Consobrinus †Cynohyaenodon †Eurotherium †Hyaenodon †Leonhardtina †Matthodon †Neosinopa †Oxyaenoides †Paenoxyaenoides †Paracynohyaenodon †Praecodens †Preregidens †Prodissopsalis †Propterodon †Protoproviverra †Quercytherium

†Proviverridae

†Allopterodon
†Lesmesodon
†Minimovellentodon
†Morlodon
†Parvagula
†Proviverra

†Afro‑Arabian
clade

†Parvavorodon
†Tritemnodon
†Koholia

†Indohyaenodontidae

†Glibzegdouia
†Indohyaenodon
†Yarshea

†Hyainailouroidea

†Hyainailouridae

†Maocyon/Orienspterodon clade

†Maocyon
†Orienspterodon

†Apterodontinae

†Apterodon
†Quasiapterodon

†Hyainailourinae

†Leakitherium †Megistotherium †Mlanyama †Metapterodon †Pakakali †Paroxyaena †Simbakubwa
†Akhnatenavus clade	†Akhnatenavus †Hemipsalodon †Ischnognathus
†Hyainailourini	†Exiguodon †Falcatodon †Hyainailouros †Parapterodon †Sectisodon †Sivapterodon †Isohyaenodon †Kerberos †Pterodon

†Prionogalidae

†Namasector
†Prionogale
†Thereutherium

†Teratodontinae

†Anasinopa
†Brychotherium
†Buhakia
†Dissopsalis
†Ekweeconfractus
†Furodon
†Kyawdawia
†Masrasector
†Metasinopa
†Paratritemnodon
†Teratodon

other representatives

†Eoproviverra †Gazinocyon †Pyrocyon
†Arfia clade	†Arfia
†Galecyon clade	†Galecyon
†Lahimia clade	†Boualitomus †Lahimia
†Limnocyonidae	†Iridodon †Limnocyon †Oxyaenodon †Prolaena †Prolimnocyon †Thinocyon
†Sinopidae	†Acarictis †Prototomus †Sinopa

ichnotaxa
of Hyaenodonta

†Creodontipus
†Dischidodacylus
†Sarcotherichnus
†Zanclonychopus
†Hyaenodontipus
†Quiritipes
†Sarjeantipes

†Oxyaenodonta

†Oxyaenidae

†Machaeroidinae	†Apataelurus †Diegoaelurus †Isphanatherium †Machaeroides
†Oxyaeninae	†Argillotherium †Dipsalidictis †Malfelis †Oxyaena †Patriofelis †Protopsalis †Sarkastodon
†Palaeonictinae	†Ambloctonus †Dipsalodon †Palaeonictis
†Tytthaeninae	†Tytthaena

Carnivoramorpha

Carnivoraformes
(Clade "A")

Clade "B"

†Messelogale †Miocyon †Simamphicyon †Uintacyon †Xinyuictis †Zodiocyon
†Quercygalidae	†Quercygale
†Gracilocyon/Oodectes clade	†Eogale †Gracilocyon †Oodectes †Paramiacis †Paroodectes

Clade "C"

Clade "D"

†Ceruttia
†Harpalodon
†Lycarion
†Neovulpavus
†Procynodictis
†Prodaphaenus
†Tapocyon
†Walshius

Carnivora
(Clade "E")

Caniformia	(See Caniformia)
Feliformia	(See Feliformia)

other representatives

†Africtis
†Dawsonicyon
†Miacis

ichnotaxa of
Carnivoraformes

†Falcatipes

†Viverravoidea

†Viverravidae

†Orientictis †Pappictidops †Preonictis †Variviverra
†Didymictinae	†Bryanictis †Didymictis †Intyrictis †Pristinictis †Protictis †Raphictis
†Ictidopappinae	†Ictidopappus
†Viverravinae	†Simpsonictis †Viverravus †Viverriscus

Incertae sedis

†Ravenictis

General characteristics

Classification and phylogeny

Classification

Phylogeny

See also

References