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Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Cladus: Synapsida
Cladus: Eupelycosauria
Cladus: Sphenacodontia
Cladus: Sphenacodontoidea
Ordo: Therapsida
Cladus: Theriodontia
Subordo: Cynodontia
Cladus: Mammaliaformes
Classis: Mammalia
Subclassis: Trechnotheria
Infraclassis: Zatheria
Supercohort: Theria
Cohort: Eutheria
Cohort: Placentalia
Cladus: Boreoeutheria
Superordo: Laurasiatheria
Cladus: Ferae
Ordo: Carnivora
Subordo: †Miacoidea
Familia: †Viverravidae
Genera: †Bryanictis – †Didymictis – †Ictidopappus – †Pristinictis – †Protictis – †Raphictis – †Simpsonictis – †Viverravus

Viverravidae ("civet ancestors") 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] They were once thought to be earliest carnivorans and ancestral to extant carnivorans, but now are placed outside the order Carnivora based on cranial morphology as a relatives (a plesion-group) to extant carnivorans.[6][7]
Contents

1 General characteristics
2 Classification and phylogeny
2.1 Classification
2.2 Phylogeny
3 References

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.[8] 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[9]

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)
Subfamily: †Didymictinae (Flynn & Galiano, 1982)
Genus: †Bryanictis (MacIntyre, 1966)
†Bryanictis microlestes (Simpson, 1935)
†Bryanictis paulus (Meehan & Wilson, 2002)
Genus: †Didymictis (Cope, 1875)[10]
†Didymictis altidens (Cope, 1880)
†Didymictis dellensis (Dorr, 1952)
†Didymictis leptomylus (Cope, 1880)
†Didymictis protenus (Cope, 1874)
†Didymictis proteus (Polly, 1997)
†Didymictis vancleveae (Robinson, 1966)
Genus: †Intyrictis (Gingerich & Winkler, 1985)
†Intyrictis vanvaleni (MacIntyre, 1966)
Genus: †Pristinictis (Fox & Youzwyshyn, 1994)
†Pristinictis connata (Fox & Youzwyshyn, 1994)
Genus: †Protictis (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)
Genus: †Protictoides (Flynn & Galiano, 1982)
†Protictoides 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)
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)
Genus: †Viverriscus (Beard & Dawson, 2009)
†Viverriscus omnivorus (Beard & Dawson, 2009)

Phylogeny

The phylogenetic relationships of Viverravidae are shown in the following cladogram:[3][11][12][13][14][15]
Carnivoramorpha

orpha 

Carnivoraformes Dogs, jackals, wolves, and foxes (Plate XI).jpg

 ? 

†Carnivoramorpha sp. (UALVP 31176)

 ? 

†Carnivoramorpha sp. (USNM 538395)

 ? 

"Sinopa" insectivorus

 †Viverravidae 
 ? 

Ravenictis

 ? 

†Carnivoramorpha sp. (UALVP 50993 & UALVP 50994)

 †Viverravoidea 
 †Viverravidae 
 †Variviverra 

Variviverra vegetatus

 †Preonictis 

Preonictis youngi

 †Orientictis 

Orientictis spanios

 †Pappictidops 

Pappictidops orientalis

Pappictidops acies

Pappictidops obtusus

 †Ictidopappinae 
 †Ictidopappus 

Ictidopappus mustelinus

 †Viverravinae 
 †Simpsonictis 

Simpsonictis pegus

Simpsonictis tenuis

Simpsonictis sp.

Simpsonictis jaynanneae

 †Viverriscus 

Viverriscus omnivorus

 †Pristinictis 

Pristinictis connata

 ? 
 †Viverravus 

Viverravus gracilis

Viverravus lutosus

Viverravus minutus

Viverravus sicarius

Viverravus politus

Viverravus lawsoni

Viverravus laytoni

Viverravus acutus

Viverravus rosei

 ? 

†Viverravidae sp. (CM 71188 & CM 71189)

 †Didymictinae 
 †Raphictis 

Raphictis gausion

Raphictis iota

Raphictis machaera

Raphictis nanoptexis

Raphictis sp. (UALVP 51558)

Raphictis sp. (UALVP 8861)

†Didymictinae sp. (UALVP 51560)

 †Didymictis 

Didymictis leptomylus

Didymictis protenus

Didymictis vancleveae

Didymictis altidens

Didymictis sp. (YPM VPPU 024902)

 †Didymictis proteus 

Didymictis proteus (sensu stricto)

 ? 

Didymictis dellensis

 sensu lato 
 †Protictis 
 †Protictis 

Protictis paralus

Protictis agastor

Protictis simpsoni

Protictis minor

Protictis haydenianus

 sensu stricto 
 †Protictoides 

Protictoides aprophatos

 †Bryanictis 
 †Intyrictis 

Intyrictis vanvaleni

 †Bryanictis 

Bryanictis microlestes

Bryanictis paulus

Bryanictis sp. (Dawson Creek)


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 uiber das Raubtiersystem." Anns. Hist.-Nat. Mus. Natn. Hung., Budapest, vol. 38, pp. 59-83.
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. Retrieved 29 June 2019 from the Paleobiology Database.
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.
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)
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.
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
Malcolm C. McKenna, Susan K. Bell: Classification of Mammals: Above the Species Level in Columbia University Press, New York (1997), 631 Seiten. Viverravidae
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

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