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{{Short description|Extinct family of carnivores}}
{{Short description|Extinct family of carnivorous mammals}}
{{About|an extinct family of mammals|the extant family|Viverridae}}
{{About|an extinct family of carnivorous mammals|the extant family of carnivorans|Viverridae}}
{{Automatic taxobox
{{Automatic taxobox
|fossil_range = {{fossil range|66.043|33.9}} early [[Paleocene]] - late [[Eocene]]
|fossil_range = {{fossil range|66.043|33.9}} early [[Paleocene]] - late [[Eocene]]
|image =
|image = Viverravus minutus 2.jpg
|image_caption =
|image_caption = skull of ''[[Viverravus|Viverravus minutus]]''
|image2 = Didymictis protenus.jpg
|display_parents = 2
|image2_caption = skull of ''[[Didymictis|Didymictis protenus]]''
|display_parents = 3
|taxon = Viverravidae
|taxon = Viverravidae
|parent_authority = Wortman & Matthew, 1899
|parent_authority = Wortman & Matthew, 1899
|authority = {{Harvnb|Wortman|Matthew|1899}}<ref>{{Cite journal |last1=Wortman |first1=J. L. |last2=Matthew |first2=W. D. |author2-link=William Diller Matthew |title=The ancestry of certain members of the Canidae, Viverridae, and Procyonidae |year=1899 |journal=Bulletin of the American Museum of Natural History |volume=12 |pages=109–138 |oclc=46687698 |hdl=2246/1535}}<!-- {{Harvnb|Wortman|Matthew|1899}} --></ref>
|authority = Wortman & Matthew, 1899<ref>{{Cite journal |last1=Wortman |first1=J. L. |last2=Matthew |first2=W. D. |author2-link=William Diller Matthew |title=The ancestry of certain members of the Canidae, Viverridae, and Procyonidae |year=1899 |journal=Bulletin of the American Museum of Natural History |volume=12 |pages=109–138 |oclc=46687698 |hdl=2246/1535}}</ref>
|type_genus = †''[[Viverravus]]''
|type_genus_authority = Marsh, 1872
|subdivision_ranks = Genera
|subdivision_ranks = Genera
|subdivision = <center>[see [[#Classification|classification]]]</center>
|subdivision = {{center|[see [[#Classification|classification]]]}}
|synonyms =
|synonyms =
{{collapsible list |bullets=true |title=synonyms of family:
{{collapsible list |bullets=true |title=synonyms of family:
|Didymictida {{small|(Kretzoi, 1945)}}<ref>Miklos Kretzoi (1945.) "Bemerkungen uiber das Raubtiersystem." Anns. Hist.-Nat. Mus. Natn. Hung., Budapest, vol. 38, pp. 59-83.</ref>
|Didymictida {{small|(Kretzoi, 1945)}}<ref>Miklos Kretzoi (1945) "Bemerkungen über das Raubtiersystem." Annales Historico-Naturales Musei Nationalis Hungarici, Budapest, vol. 38, pp. 59–83.</ref>
|Didymictidae {{small|(Flynn & Galiano, 1982)}}<ref name=Flynn1982/>
|Didymictidae {{small|(Flynn & Galiano, 1982)}}<ref name="Flynn1982"/>
|Viverravidea
|Viverravidea
|Viverravinae {{small|(Matthew, 1909)}}<ref>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</ref>
|Viverravinae {{small|(Matthew, 1909)}}<ref>W. D. Matthew (1909) [https://digitallibrary.amnh.org/handle/2246/5744 "The Carnivora and Insectivora of the Bridger Basin, middle Eocene."] Memoirs of the American Museum of Natural History 9:289-567</ref>
}}
}}
}}
}}


'''Viverravidae''' ("[[civet]] ancestors") is an [[extinct]] [[monophyletic]] [[Family (biology)|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]].<ref>{{BioRef |fossilworks |title=†family Viverravidae (Wortman & Matthew, 1899) (placental) |id=40992 |access-date=29 June 2019}}</ref> They are once thought to be earliest carnivorans and ancestral to extant carnivorans, but now are placed outside the order [[Carnivora]] based on cranial [[morphology (biology)|morphology]] as a relatives (a [[Crown group|plesion-group]]) to extant carnivorans.<ref>{{Cite journal |last1=Wesley-Hunt |first1=G. D. |last2=Flynn |first2=J. J. |title=Phylogeny of the Carnivora: basal relationships among the carnivoramorphans, and assessment of the position of 'Miacoidea' relative to Carnivora |year=2005 |journal=Journal of Systematic Palaeontology |volume=3 |pages=1–28 |doi=10.1017/S1477201904001518 |s2cid=86755875}}</ref><ref name=Polly>{{cite journal |author=Polly, David, Gina D. Wesley-Hunt, Ronald E. Heinrich, Graham Davis and Peter Houde |year=2006 |title=Earliest known carnivoran auditory bulla and support for a recent origin of crown-clade carnivora (Eutheria, Mammalia) |journal=Palaeontology |volume=49 |issue=5 |pages=1019–1027 |url=http://mypage.iu.edu/~pdpolly/Papers/Polly%20et%20al,%202006,%20Viverravus.pdf |doi=10.1111/j.1475-4983.2006.00586.x |doi-access=free}}</ref>
'''Viverravidae''' ("ancestors of [[Viverridae|viverrids]]") is an [[extinct]] [[monophyletic]] [[Family (biology)|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]].<ref>{{BioRef |fossilworks |title=†family Viverravidae (Wortman & Matthew, 1899) (placental) |id=40992 |access-date=29 June 2019}}</ref><ref name="McKenna & Bell">Malcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'' in Columbia University Press, New York (1997), 631 Seiten. {{page|year=|isbn=0-231-11013-8|pages=}}</ref><ref name="Zack2012">{{cite journal |last1=Zack |first1=Shawn P. |title=Deciduous dentition of ''Didymictis'' (Carnivoramorpha: Viverravidae): implications for the first appearance of "Creodonta" |journal=Journal of Mammalogy |volume=93 |issue=3 |year=2012 |pages=808–817 |issn=0022-2372 |doi=10.1644/11-MAMM-A-245.1 |doi-access=free}}</ref> 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 (biology)|morphology]] as relatives (a [[Crown group|plesion-group]]) to extant carnivorans.<ref>{{Cite journal |last1=Wesley-Hunt |first1=G. D. |last2=Flynn |first2=J. J. |title=Phylogeny of the Carnivora: basal relationships among the carnivoramorphans, and assessment of the position of 'Miacoidea' relative to Carnivora |year=2005 |journal=Journal of Systematic Palaeontology |volume=3 |pages=1–28 |doi=10.1017/S1477201904001518 |s2cid=86755875}}</ref><ref name=Polly>{{cite journal |author=Polly, David, Gina D. Wesley-Hunt, Ronald E. Heinrich, Graham Davis and Peter Houde |year=2006 |title=Earliest known carnivoran auditory bulla and support for a recent origin of crown-clade carnivora (Eutheria, Mammalia) |journal=Palaeontology |volume=49 |issue=5 |pages=1019–1027 |url=http://mypage.iu.edu/~pdpolly/Papers/Polly%20et%20al,%202006,%20Viverravus.pdf |doi=10.1111/j.1475-4983.2006.00586.x |doi-access=free}}</ref>


==General characteristics==
==General characteristics==
[[Xiaoming Wang (paleontologist)|Wang]] and [[Richard H. Tedford|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.<ref name=wang2008>Wang, Xiaoming; Tedford, Richard H. (2008.) "Dogs: Their Fossil Relatives and Evolutionary History." New York: Columbia University Press</ref> In viverravids, the [[skull]] is elongated and the number of [[Molar (tooth)|molars]] is reduced to two (M1/m1 and M2/m2 are present and M3/m3 are absent).
[[Xiaoming Wang (paleontologist)|Wang]] and [[Richard H. Tedford|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.<ref name=wang2008>Wang, Xiaoming; Tedford, Richard H. (2008.) "Dogs: Their Fossil Relatives and Evolutionary History." New York: Columbia University Press</ref> In viverravids, the [[skull]] is elongated and the number of [[Molar (tooth)|molars]] is reduced to two (M1/m1 and M2/m2 are present and M3/m3 are absent).


==Classification and phylogeny==
==Classification and phylogeny==
===Classification===
===Classification===
Taxonomy retrieved from the [[Paleobiology Database]]<ref>Paleobiology Database. Retrieved with [http://fossilworks.org/bridge.pl?action=taxonInfo&taxon_no=40992 Fossilworks] (March 3, 2017)</ref>
Taxonomy retrieved from the [[Paleobiology Database]]<ref>Paleobiology Database. Retrieved with [http://www.fossilworks.org/cgi-bin/bridge.pl?action=taxonInfo&taxon_no=40992 Fossilworks] (March 3, 2017)</ref>
{|class="wikitable"
* Superfamily: †'''Viverravoidea''' <small>(Wortman & Matthew, 1899)</small>
|- style="vertical-align:top;"
** Family: †'''Viverravidae''' <small>(Wortman & Matthew, 1899)</small>
|style="text-align: center;"|Superfamily: †'''Viverravoidea''' {{small|(Wortman & Matthew, 1899)}}<br />Family: †'''Viverravidae''' {{small|(Wortman & Matthew, 1899)}}
*** Genus: †''[[Orientictis]]'' <small>(Huang & Zheng, 2005)</small>
----
**** †''Orientictis spanios'' <small>(Huang & Zheng, 2005)</small>
:{|
*** Genus: †''[[Pappictidops]]'' <small>(Qiu & Li, 1977)</small>
|- style="vertical-align:top;"
**** †''Pappictidops acies'' <small>(Wang, 1978)</small>
|
**** †''Pappictidops obtusus'' <small>(Wang, 1978)</small>
**** †''Pappictidops orientalis'' <small>(Qiu & Li, 1977)</small>
* Genus: †''[[Orientictis]]'' {{small|(Huang & Zheng, 2005)}}
*** Genus: †''[[Variviverra]]'' <small>(Tong & Wang, 2006)</small>
** †''Orientictis spanios'' {{small|(Huang & Zheng, 2005)}}
**** †''Variviverra vegetatus'' <small>(Tong & Wang, 2006)</small>
* Genus: †''[[Pappictidops]]'' {{small|(Qiu & Li, 1977)}}
*** Subfamily: †'''[[Didymictinae]]''' <small>(Flynn & Galiano, 1982)</small>
** †''Pappictidops acies'' {{small|(Wang, 1978)}}
**** Genus: †''[[Bryanictis]]'' <small>(MacIntyre, 1966)</small>
** †''Pappictidops obtusus'' {{small|(Wang, 1978)}}
***** †''Bryanictis microlestes'' <small>(Simpson, 1935)</small>
** †''Pappictidops orientalis'' {{small|(Qiu & Li, 1977)}}
***** †''Bryanictis paulus'' <small>(Meehan & Wilson, 2002)</small>
* Genus: †''[[Preonictis]]'' {{small|(Tong & Wang, 2006)}}
** †''Preonictis youngi'' {{small|(Tong & Wang, 2006)}}
**** Genus: †''[[Didymictis]]'' <small>(Cope, 1875)</small><ref name="Zack2012">{{cite journal |last1=Zack |first1=Shawn P. |title=Deciduous dentition of ''Didymictis'' (Carnivoramorpha: Viverravidae): implications for the first appearance of "Creodonta" |journal=Journal of Mammalogy |volume=93 |issue=3 |year=2012 |pages=808–817 |issn=0022-2372 |doi=10.1644/11-MAMM-A-245.1 |doi-access=free}}</ref>
***** †''Didymictis altidens'' <small>(Cope, 1880)</small>
* Genus: †''[[Variviverra]]'' {{small|(Tong & Wang, 2006)}}
***** †''Didymictis dellensis'' <small>(Dorr, 1952)</small>
** †''Variviverra vegetatus'' {{small|(Tong & Wang, 2006)}}
* ''[[Incertae sedis]]'':
***** †''Didymictis leptomylus'' <small>(Cope, 1880)</small>
** †[[Viverravidae sp. (CM 71188 & CM 71189)|Viverravidae sp. [''CM 71188'' & ''CM 71189'']]] {{small|(Friscia & Rasmussen, 2010)}}
***** †''Didymictis protenus'' <small>(Cope, 1874)</small>
***** †''Didymictis proteus'' <small>(Polly, 1997)</small>
* Subfamily: †'''[[Didymictinae]]''' {{small|(Flynn & Galiano, 1982)}}
***** †''Didymictis vancleveae'' <small>(Robinson, 1966)</small>
** Genus: †''[[Bryanictis]]'' {{small|(MacIntyre, 1966)}}
**** Genus: †''[[Intyrictis]]'' <small>(Gingerich & Winkler, 1985)</small>
*** †''Bryanictis microlestes'' {{small|(Simpson, 1935)}}
***** †''Intyrictis vanvaleni'' <small>(MacIntyre, 1966)</small>
*** †''Bryanictis paulus'' {{small|(Meehan & Wilson, 2002)}}
**** Genus: †''[[Protictis]]'' <small>(Matthew, 1937)</small>
*** †''Bryanictis terlinguae'' {{small|(Standhardt, 1986)}}
***** †''Protictis agastor'' <small>(Gingerich & Winkler, 1985)</small>
** Genus: †''[[Didymictis]]'' {{small|(Cope, 1875)}}
***** †''Protictis haydenianus'' <small>(Cope, 1882)</small>
*** †''Didymictis altidens'' {{small|(Cope, 1880)}}
***** †''Protictis minor'' <small>(Meehan & Wilson, 2002)</small>
*** †''Didymictis dellensis'' {{small|(Dorr, 1952)}}
***** †''Protictis paralus'' <small>(Holtzman, 1978)</small>
*** †''Didymictis leptomylus'' {{small|(Cope, 1880)}}
***** †''Protictis simpsoni'' <small>(Meehan & Wilson, 2002)</small>
*** †''Didymictis protenus'' {{small|(Cope, 1874)}}
**** Genus: †''[[Protictoides]]'' <small>(Flynn & Galiano, 1982)</small>
*** †''Didymictis proteus'' {{small|(Polly, 1997)}}
***** †''Protictoides aprophatos'' <small>(Flynn & Galiano, 1982)</small>
*** †''Didymictis vancleveae'' {{small|(Robinson, 1966)}}
*** †''Didymictis'' sp. [Erquelinnes, Hainaut, Belgium] {{small|(Dollo, 1909)}}
**** Genus: †''[[Raphictis]]'' <small>(Gingerich & Winkler, 1985)</small>
***** †''Raphictis gausion'' <small>(Gingerich & Winkler, 1985)</small>
** Genus: †''[[Intyrictis]]'' {{small|(Gingerich & Winkler, 1985)}}
***** †''Raphictis iota'' <small>(Scott, 2008)</small>
*** †''Intyrictis vanvaleni'' {{small|(MacIntyre, 1966)}}
***** †''Raphictis machaera'' <small>(Rankin, 2009)</small>
** Genus: †''[[Pristinictis]]'' {{small|(Fox & Youzwyshyn, 1994)}}
***** †''Raphictis nanoptexis'' <small>(Rankin, 2009)</small>
*** †''Pristinictis connata'' {{small|(Fox & Youzwyshyn, 1994)}}
** Genus: †''[[Protictis]]'' <sup>([[Paraphyly|paraphyletic]] genus)</sup> {{small|(Matthew, 1937)}}
*** Subfamily: †'''[[Ictidopappus|Ictidopappinae]]''' <small>(Van Valen, 1969)</small>
**** Genus: †''[[Ictidopappus]]'' <small>(Simpson, 1935)</small>
*** †''Protictis agastor'' {{small|(Gingerich & Winkler, 1985)}}
***** †''Ictidopappus mustelinus'' <small>(Simpson, 1935)</small>
*** †''Protictis haydenianus'' {{small|(Cope, 1882)}}
*** Subfamily: †'''[[Viverravinae]]''' <small>(Wortman & Matthew, 1899)</small>
*** †''Protictis minor'' {{small|(Meehan & Wilson, 2002)}}
**** Genus: †''[[Pristinictis]]'' <small>(Fox & Youzwyshyn, 1994)</small>
*** †''Protictis paralus'' {{small|(Holtzman, 1978)}}
***** †''Pristinictis connata'' <small>(Fox & Youzwyshyn, 1994)</small>
*** †''Protictis simpsoni'' {{small|(Meehan & Wilson, 2002)}}
**** Genus: †''[[Simpsonictis]]'' <small>(MacIntyre, 1962)</small>
*** Subgenus: †''[[Protictoides]]'' {{small|(Flynn & Galiano, 1982)}}
***** †''Simpsonictis jaynanneae'' <small>(Rigby, 1980)</small>
**** †''Protictis aprophatos'' {{small|(Flynn & Galiano, 1982)}}
***** †''Simpsonictis pegus'' <small>(Gingerich & Winkler, 1985)</small>
** Genus: †''[[Raphictis]]'' {{small|(Gingerich & Winkler, 1985)}}
***** †''Simpsonictis tenuis'' <small>(Simpson, 1935)</small>
*** †''Raphictis gausion'' {{small|(Gingerich & Winkler, 1985)}}
**** Genus: †''[[Viverravus]]'' <small>(Marsh, 1872)</small>
*** †''Raphictis iota'' {{small|(Scott, 2008)}}
***** †''Viverravus acutus'' <small>(Matthew & Granger, 1915)</small>
*** †''Raphictis machaera'' {{small|(Rankin, 2009)}}
***** †''Viverravus gracilis'' <small>(Marsh, 1872)</small>
*** †''Raphictis nanoptexis'' {{small|(Rankin, 2009)}}
** ''[[Incertae sedis]]'':
***** †''Viverravus lawsoni'' <small>(Hooker, 2010)</small>
***** †''Viverravus laytoni'' <small>(Gingerich & Winkler, 1985)</small>
*** †''[[Deltatherium durini|"Deltatherium" durini]]'' {{small|(Van Valen, 1978)}}
|
***** †''Viverravus lutosus'' <small>(Gazin, 1952)</small>
***** †''Viverravus minutus'' <small>(Wortman, 1901)</small>
* Subfamily: †'''[[Ictidopappus|Ictidopappinae]]''' {{small|(Van Valen, 1969)}}
***** †''Viverravus politus'' <small>(Matthew & Granger, 1915)</small>
** Genus: †''[[Ictidopappus]]'' {{small|(Simpson, 1935)}}
***** †''Viverravus rosei'' <small>(Polly, 1997)</small>
*** †''Ictidopappus mustelinus'' {{small|(Simpson, 1935)}}
***** †''Viverravus sicarius'' <small>(Matthew, 1909)</small>
* Subfamily: †'''[[Viverravinae]]''' {{small|(Wortman & Matthew, 1899)}}
**** Genus: †''[[Viverriscus]]'' <small>(Beard & Dawson, 2009)</small>
** Genus: †''[[Simpsonictis]]'' {{small|(MacIntyre, 1962)}}
***** †''Viverriscus omnivorus'' <small>(Beard & Dawson, 2009)</small>
*** †''Simpsonictis jaynanneae'' {{small|(Rigby, 1980)}}
*** †''Simpsonictis pegus'' {{small|(Gingerich & Winkler, 1985)}}
*** †''Simpsonictis tenuis'' {{small|(Simpson, 1935)}}
** Genus: †''[[Viverravus]]'' {{small|(Marsh, 1872)}}
*** †''Viverravus acutus'' {{small|(Matthew & Granger, 1915)}}
*** †''Viverravus gracilis'' {{small|(Marsh, 1872)}}
*** †''Viverravus lawsoni'' {{small|(Hooker, 2010)}}
*** †''Viverravus laytoni'' {{small|(Gingerich & Winkler, 1985)}}
*** †''Viverravus lutosus'' {{small|(Gazin, 1952)}}
*** †''Viverravus minutus'' {{small|(Wortman, 1901)}}
*** †''Viverravus politus'' {{small|(Matthew & Granger, 1915)}}
*** †''Viverravus rosei'' {{small|(Polly, 1997)}}
*** †''Viverravus sicarius'' {{small|(Matthew, 1909)}}
*** †''Viverravus'' sp. [''V11141''] {{small|(Meng, 1998)}}
*** †''Viverravus'' sp. [Locality Group 2, Washakie Basin, Wyoming] {{small|(Tomiya, 2021)}}
** Genus: †''[[Viverriscus]]'' {{small|(Beard & Dawson, 2009)}}
*** †''Viverriscus omnivorus'' {{small|(Beard & Dawson, 2009)}}
|}
|}


===Phylogeny===
===Phylogeny===
The phylogenetic relationships of Viverravidae are shown in the following cladogram:<ref name=Flynn1982>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</ref><ref>P. D. Gingerich and D. A. Winkler. (1985.) [https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48525/ID378.pdf?sequence=2&isAllowed=y "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</ref><ref>P. D. Polly. (1997.) [https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48638/ID504.pdf?sequence=2 "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</ref><ref name="McKenna & Bell">Malcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'' in Columbia University Press, New York (1997), 631 Seiten. {{page|year=|isbn=0-231-11013-8|pages=}}</ref><ref name="SoléSmith2016">{{cite journal |last1=Solé |first1=Floréal |last2=Smith |first2=Thierry |last3=De Bast |first3=Eric |last4=Codrea |first4=Vlad |last5=Gheerbrant |first5=Emmanuel |title=New carnivoraforms from the latest Paleocene of Europe and their bearing on the origin and radiation of Carnivoraformes (Carnivoramorpha, Mammalia) |journal=Journal of Vertebrate Paleontology |volume=36 |issue=2 |year=2016 |pages=e1082480 |issn=0272-4634 |doi=10.1080/02724634.2016.1082480|s2cid=87537565 }}</ref><ref>S. Faurby, L. Werdelin, A. Antonelli (2019.) [https://www.biorxiv.org/content/biorxiv/early/2019/09/05/755207.full.pdf "Dispersal ability predicts evolutionary success among mammalian carnivores"] Department of Biological and Environmental Sciences, University of Gothenburg, Box 461, SE</ref>
The phylogenetic relationships of family Viverravidae are shown in the following cladogram:<ref name=Flynn1982>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</ref><ref>P. D. Gingerich and D. A. Winkler. (1985.) [https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48525/ID378.pdf?sequence=2&isAllowed=y "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</ref><ref>P. D. Polly. (1997.) [https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48638/ID504.pdf?sequence=2 "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</ref><ref name="SoléSmith2016">{{cite journal |last1=Solé |first1=Floréal |last2=Smith |first2=Thierry |last3=De Bast |first3=Eric |last4=Codrea |first4=Vlad |last5=Gheerbrant |first5=Emmanuel |title=New carnivoraforms from the latest Paleocene of Europe and their bearing on the origin and radiation of Carnivoraformes (Carnivoramorpha, Mammalia) |journal=Journal of Vertebrate Paleontology |volume=36 |issue=2 |year=2016 |pages=e1082480 |issn=0272-4634 |doi=10.1080/02724634.2016.1082480|s2cid=87537565 }}</ref><ref>S. Faurby, L. Werdelin, A. Antonelli (2019.) [https://www.biorxiv.org/content/biorxiv/early/2019/09/05/755207.full.pdf "Dispersal ability predicts evolutionary success among mammalian carnivores"] Department of Biological and Environmental Sciences, University of Gothenburg, Box 461, SE</ref>
{{clade |style=font-size:85%;line-height:100%;
{{clade |style=font-size:85%;line-height:100%;
|label1=&nbsp;'''[[Carnivoramorpha]]'''&nbsp;
|label1=&nbsp;'''[[Carnivoramorpha]]'''&nbsp;
Line 95: Line 117:
|1='''[[Carnivoraformes]]''' [[File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XI).jpg|50px]]
|1='''[[Carnivoraformes]]''' [[File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XI).jpg|50px]]
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|2=†Carnivoramorpha sp. (''UALVP 31176'')
|2=†[[Carnivoramorpha sp. (UALVP 31176)|Carnivoramorpha sp. (''UALVP 31176'')]]
|label3=&nbsp;'''?'''&nbsp; |state3=dotted
|label3=&nbsp;'''?'''&nbsp; |state3=dotted
|3=†Carnivoramorpha sp. (''USNM 538395'')
|3=†[[Carnivoramorpha sp. (USNM 538395)|Carnivoramorpha sp. (''USNM 538395'')]]
|label4=&nbsp;'''?'''&nbsp; |state4=dotted
|label4=&nbsp;'''?'''&nbsp; |state4=dotted
|4=†''[[Sinopa insectivorus|"Sinopa" insectivorus]]''
|4=†''[[Sinopa insectivorus|"Sinopa" insectivorus]]''
Line 105: Line 127:
|1=†''[[Ravenictis]]''
|1=†''[[Ravenictis]]''
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|2=†Carnivoramorpha sp. (''UALVP 50993'' & ''UALVP 50994'')
|2=†[[Carnivoramorpha sp. (UALVP 50993 & UALVP 50994)|Carnivoramorpha sp. (''UALVP 50993'' & ''UALVP 50994'')]]
|label3=&nbsp;†'''Viverravoidea'''&nbsp;
|label3=&nbsp;†'''Viverravoidea'''&nbsp;
|3={{clade
|3={{clade
|thickness=2 |label1=&nbsp;†'''Viverravidae'''&nbsp;
|thickness=2 |label1=&nbsp;†'''Viverravidae'''&nbsp; |sublabel1=&nbsp;''sensu stricto''&nbsp;
|1={{clade
|1={{clade
|thickness=2
|thickness=2
Line 114: Line 136:
|thickness=2
|thickness=2
|1={{clade
|1={{clade
|thickness=2
|thickness=2 |label1=&nbsp;†''[[Pappictidops]]''&nbsp;
|1={{clade
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Variviverra]]''&nbsp;
|1=†''Variviverra vegetatus''
|label2=&nbsp;†''[[Preonictis]]''&nbsp;
|2=†''Preonictis youngi''
}}
|label2=&nbsp;†''[[Orientictis]]''&nbsp;
|2=†''Orientictis spanios''
|label3=&nbsp;†''[[Pappictidops]]''&nbsp;
|3={{clade
|thickness=2
|thickness=2
|1=†''Pappictidops orientalis''
|1={{clade
|2={{clade
|thickness=2
|thickness=2
|1=†''Pappictidops acies''
|1=†''Pappictidops acies''
|2=†''Pappictidops obtusus''
|2=†''Pappictidops obtusus''
}} }} }}
}}
|label2=&nbsp;†'''[[Ictidopappus|Ictidopappinae]]'''&nbsp;
|2=†''Pappictidops orientalis''
}}
|label2=&nbsp;†''[[Orientictis]]''&nbsp;
|2=†''Orientictis spanios''
|label3=&nbsp;†''[[Preonictis]]''&nbsp;
|3=†''Preonictis youngi''
|label4=&nbsp;†''[[Variviverra]]''&nbsp;
|4=†''Variviverra vegetatus''
}}
|2={{clade
|2={{clade
|thickness=2 |label1=&nbsp;†''[[Ictidopappus]]''&nbsp;
|thickness=2 |label1=&nbsp;†'''[[Ictidopappus|Ictidopappinae]]'''&nbsp;
|1=†''Ictidopappus mustelinus''
|1={{clade
}} }}
|thickness=2 |label1=&nbsp;†''[[Ictidopappus]]''&nbsp;
|1=†''Ictidopappus mustelinus''
}} }} }}
|2={{clade
|2={{clade
|thickness=2 |label1=&nbsp;†'''[[Viverravinae]]'''&nbsp;
|thickness=2 |label1=&nbsp;†'''[[Viverravinae]]'''&nbsp;
|1={{clade
|1={{clade
|thickness=2
|thickness=2 |label1=&nbsp;†''[[Viverravus]]''&nbsp;
|1={{clade
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Simpsonictis]]''&nbsp;
|thickness=2
|1={{clade
|1={{clade
|thickness=2
|thickness=2
|1={{clade
|1={{clade
|thickness=2
|thickness=2
|1=†''Simpsonictis pegus''
|1={{clade
|2=†''Simpsonictis tenuis''
|thickness=2
|1={{clade
|thickness=2
|1=†''Viverravus minutus''
|2=†''Viverravus sicarius''
|3=†''Viverravus'' sp. (Locality Group 2, Washakie Basin, Wyoming)
}}
|2=†''Viverravus gracilis''
|3=†''Viverravus lutosus''
}}
|2=†''Viverravus acutus''
|3=†''Viverravus rosei''
}}
}}
|2=†''Simpsonictis'' sp.
|2=†''Viverravus laytoni''
|3=†''Simpsonictis jaynanneae''
}}
}}
|label2=&nbsp;''[[Viverriscus]]''&nbsp;
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|2=†''Viverriscus omnivorus''
|2=†''Viverravus'' sp. (''V11141'')
|3=†''Viverravus lawsoni''
|4=†''Viverravus politus''
}}
}}
|2={{clade
|label2=&nbsp;†''[[Pristinictis]]''&nbsp; |sublabel2=&nbsp;'''?'''&nbsp; |state2=dotted
|2=†''Pristinictis connata''
|thickness=2 |label1=&nbsp;†''[[Viverriscus]]''&nbsp;
|label3=&nbsp;†''[[Viverravus]]''&nbsp;
|1=†''Viverriscus omnivorus''
|3={{clade
|label2=&nbsp;†''[[Simpsonictis]]''&nbsp;
|thickness=2
|2={{clade
|1={{clade
|thickness=2
|thickness=2
|1=†''Viverravus gracilis''
|1=†''Simpsonictis jaynanneae''
|2=†''Viverravus lutosus''
|2=†''Simpsonictis pegus''
|3=†''Viverravus minutus''
|3=†''Simpsonictis tenuis''
|4=†''Viverravus sicarius''
}} }} }}
}}
|2=†''Viverravus politus''
|3=†''Viverravus lawsoni''
|4={{clade
|thickness=2
|1=†''Viverravus laytoni''
|2={{clade
|thickness=2
|1=†''Viverravus acutus''
|2=†''Viverravus rosei''
}} }} }} }}
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|2=†Viverravidae sp. (''CM 71188'' & ''CM 71189'')
|2=†[[Viverravidae sp. (CM 71188 & CM 71189)|Viverravidae sp. (''CM 71188'' & ''CM 71189'')]]
|label3=&nbsp;†'''[[Didymictinae]]'''&nbsp;
|label3=&nbsp;†'''[[Didymictinae]]'''&nbsp;
|3={{clade
|3={{clade
|thickness=2
|thickness=2
|1={{clade
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Raphictis]]''&nbsp;
|thickness=2 |label1=&nbsp;†''[[Bryanictis]]''&nbsp;
|1={{clade
|1={{clade
|thickness=2
|thickness=2
|1=†''Raphictis gausion''
|1=†''Bryanictis microlestes''
|2=†''Raphictis iota''
|2=†''Bryanictis paulus''
|3=†''Raphictis machaera''
|3=†''Bryanictis terlinguae''
|4=†''Raphictis nanoptexis''
|5=†''Raphictis'' sp. (''UALVP 51558'')
|6=†''Raphictis'' sp. (''UALVP 8861'')
}}
}}
|2=†Didymictinae sp. (''UALVP 51560'')
|label2=&nbsp;†''[[Intyrictis]]''&nbsp;
|2=†''Intyrictis vanvaleni''
}}
}}
|label2=&nbsp;†''[[Didymictis]]''&nbsp;
|2={{clade
|2={{clade
|thickness=2 |caption={{Uline|c=green|&nbsp;†''[[Protictis]]''&nbsp;}} |captionstyle=text-align:center;font-weight:bold;padding:2px 10px 1px 1px; |color=green
|thickness=2
|1={{clade
|1={{Uline|c=green|&nbsp;†''[[Protictis|Protictis haydenianus]]''&nbsp;}}
|thickness=2
|2={{Uline|c=green|&nbsp;†''[[Protictis|Protictis minor]]''&nbsp;}}
|1=†''Didymictis leptomylus''
|3={{Uline|c=green|&nbsp;†''[[Protictis|Protictis simpsoni]]''&nbsp;}}
|2=†''Didymictis protenus''
|4={{clade
}}
|thickness=2 |color=green
|2=†''Didymictis vancleveae''
|3=†''Didymictis altidens''
|4=†''Didymictis'' sp. (''YPM VPPU 024902'')
|label5=&nbsp;†'''''Didymictis proteus'''''&nbsp; |sublabel5=&nbsp;''sensu lato''&nbsp;
|5={{clade
|thickness=2
|1=†''Didymictis proteus'' {{small|(''sensu stricto'')}}
|label2=&nbsp;'''?'''&nbsp; |state2=dotted
|2=†''Didymictis dellensis''
}} }}
|label3=&nbsp;†'''''Protictis'''''&nbsp; |sublabel3=&nbsp;''sensu lato''&nbsp;
|3={{clade
|thickness=2
|1={{clade
|thickness=2 |label1=&nbsp;†''[[Protictis]]''&nbsp; |sublabel1=&nbsp;''sensu stricto''&nbsp;
|1={{clade
|1={{clade
|thickness=2
|thickness=2 |label1=&nbsp;†''[[Raphictis]]''&nbsp; |color=green |color1=black
|1={{clade
|1={{clade
|thickness=2
|thickness=2
|1=†''Protictis paralus''
|1=†''Raphictis gausion''
|2=†''Protictis agastor''
|2=†''Raphictis iota''
|3=†''Raphictis machaera''
|4=†''Raphictis nanoptexis''
}}
}}
|2=†''Protictis simpsoni''
|2={{Uline|c=green|&nbsp;†''[[Protictis|Protictis paralus]]''&nbsp;}}
|3=†''Protictis minor''
|4=†''Protictis haydenianus''
}}
}}
|label2=&nbsp;†''[[Protictoides]]''&nbsp;
|2=†''Protictoides aprophatos''
}}
|label2=&nbsp;†'''''Bryanictis'''''&nbsp; |sublabel2=&nbsp;''sensu lato''&nbsp;
|2={{clade
|thickness=2 |label1=&nbsp;†''[[Intyrictis]]''&nbsp;
|1=†''Intyrictis vanvaleni''
|label2=&nbsp;†''[[Bryanictis]]''&nbsp; |sublabel2=&nbsp;''sensu stricto''&nbsp;
|2={{clade
|2={{clade
|thickness=2
|thickness=2 |color=green |color3=black |color4=black |color5=black
|1=†''Bryanictis microlestes''
|1={{Uline|c=green|&nbsp;†''[[Protictis|Protictis agastor]]''&nbsp;}}
|2=†''Bryanictis paulus''
|label2=&nbsp;(''[[Protictoides]]'')&nbsp;
|3=†''Bryanictis'' sp. (Dawson Creek)
|2={{Uline|c=green|&nbsp;†''[[Protictoides|Protictis aprophatos]]''&nbsp;}}
}} }} }} }} }} }} }} }} }} }}
|label3=&nbsp;†''[[Pristinictis]]''&nbsp;
|3=†''Pristinictis connata''
|label4=&nbsp;'''?'''&nbsp; |state4=dotted
|4=†''[[Deltatherium durini|"Deltatherium" durini]]''
|label5=&nbsp;†''[[Didymictis]]''&nbsp;
|5={{clade
|thickness=2
|1=†''Didymictis dellensis''
|2=†''Didymictis proteus''
|label3=&nbsp;'''?'''&nbsp; |state3=dotted
|3=†''Didymictis'' sp. (Erquelinnes, Hainaut, Belgium)
|4={{clade
|thickness=2
|1=†''Didymictis altidens''
|2=†''Didymictis leptomylus''
|3=†''Didymictis protenus''
|4=†''Didymictis vancleveae''
}} }} }} }} }} }} }} }} }} }} }} }}

==See also==
* [[Mammal classification]]
* [[Carnivoramorpha]]
* [[Miacoidea]]


==References==
==References==
{{Reflist}}
{{Reflist|2}}

{{Pan-Carnivora|C.|state=collapsed}}
{{Taxonbar|from=Q4889586}}
{{Taxonbar|from=Q4889586}}


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[[Category:Eocene extinctions]]
[[Category:Eocene extinctions]]
[[Category:Prehistoric mammal families]]
[[Category:Prehistoric mammal families]]



{{Paleo-carnivora-stub}}
{{Paleo-carnivora-stub}}

Latest revision as of 20:41, 19 January 2024

Viverravidae
Temporal range: 66.043–33.9 Ma early Paleocene - late Eocene
skull of Viverravus minutus
skull of Didymictis protenus
Scientific classification Edit this 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
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[edit]

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[edit]

Classification[edit]

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:
  • 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:
  • 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[edit]

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 

See also[edit]

References[edit]

  1. ^ 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.
  2. ^ Miklos Kretzoi (1945) "Bemerkungen über das Raubtiersystem." Annales Historico-Naturales Musei Nationalis Hungarici, Budapest, vol. 38, pp. 59–83.
  3. ^ 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
  4. ^ 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
  5. ^ "†family Viverravidae (Wortman & Matthew, 1899) (placental)". Fossilworks. Gateway to the Paleobiology Database. Retrieved 29 June 2019.
  6. ^ Malcolm C. McKenna, Susan K. Bell: Classification of Mammals: Above the Species Level in Columbia University Press, New York (1997), 631 Seiten. Viverravidae
  7. ^ 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.
  8. ^ 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.
  9. ^ 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)
  10. ^ Wang, Xiaoming; Tedford, Richard H. (2008.) "Dogs: Their Fossil Relatives and Evolutionary History." New York: Columbia University Press
  11. ^ Paleobiology Database. Retrieved with Fossilworks (March 3, 2017)
  12. ^ 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
  13. ^ 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
  14. ^ 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.
  15. ^ 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