Carnivoramorpha

Carnivoramorpha; Temporal range: 66.043–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N early Paleocene to present
	Diversity of Carnivoramorpha
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Infraclass:	Placentalia
Mirorder:	Ferae
Clade:	Pan-Carnivora
Clade:	Carnivoramorpha; Wyss & Flynn, 1993
Subgroups
	[see classification]
Synonyms
	Carnivora (Rose, 2012); Carnivoramoepha (Matsui & Kimura, 2022); Carnivoramomorpha (Wyss & Flynn, 1993);

Carnivoramorpha ("carnivoran-like forms") is a clade of placental mammals within the larger clade Pan-Carnivora from the mirorder Ferae, that includes the modern order Carnivora as well as extinct stem-group relatives that are more closely related to carnivorans proper than to "creodonts".^[4]^[5] However, recent analyses suggest carnivoramorphans was polyphyletic with Viverravidae being basal to the clade containing hyaenodonts, oxyaenids and carnivoraforms.^[6]^[7]^[8]

General characteristics

The common feature for members of this clade is the presence of the carnassial teeth. The carnassial teeth of the Carnivoramorpha are upper premolar P4 and lower molar m1.^[9]

Comparison of carnassial teeth of a carnivoran (wolf), a hyaenodontid (Hyaenodon) and an oxyaenid (Oxyaena)

diagram of a wolf skull with key features labelled

skull of Viverravus minutus

skull of Vulpavus profectus

Classification and phylogeny

Traditional classification

Clade: Carnivoramorpha (Wyss & Flynn, 1993) [carnivorans + all basal clades to Carnivora, but without creodonts] Order: Carnivora (Bowdich, 1821) (carnivorans) Suborder: Caniformia (Kretzoi, 1943) ("dog-like" carnivorans) Suborder: Feliformia (Kretzoi, 1945) ("cat-like" carnivorans) Incertae sedis: Genus: †Palaeogale (von Meyer, 1846) Superfamily: †Miacoidea (Cope, 1880) Family: †Miacidae (Cope, 1880) Family: †Viverravidae (Wortman & Matthew, 1899) Incertae sedis: †"Sinopa" insectivorus (Cope, 1872) Incertae sedis: Genus: †Ravenictis (Fox & Youzwyshyn, 1994) †Carnivoramorpha sp. [UALVP 31176] (Fox & Youzwyshyn, 1994)

Revised classification

Recent phylogenetic studies indicate that the superfamily Miacoidea and family Miacidae are paraphyletic, with "miacids" being more closely related to carnivorans than to viverravids. In 2010 Flynn, Finarelli & Spaulding named a new clade Carnivoraformes within Carnivoramorpha, containing carnivorans and "miacids" but not viverravids.^[10] The authors defined Carnivoraformes as the clade containing Carnivora and all taxa that are more closely related to Carnivora (represented by Canis lupus) than to viverravids (represented by Viverravus gracilis). More recent analysis have questioned the relation of viverravids to carnivoraforms, suggesting carnivoramorphans wasn't a monophyletic group.^[6]^[7]^[8]

Clade: Carnivoramorpha (Wyss & Flynn, 1993) [= Carnivora (sensu lato)] Clade: Carnivoraformes (Flynn, 2010) [= Clade "A"] Genus: †Africtis (Mattingly, 2020) Genus: †Dawsonicyon (Spaulding, Flynn & Stucky, 2010) Genus: †Miacis (Cope, 1872) (unranked): Clade "B" Family: †Quercygalidae (Kretzoi, 1945) (unranked): †Gracilocyon/Oodectes clade Genus: †Eogale (Beard & Dawson, 2009) Genus: †Gracilocyon ^{(paraphyletic genus)} (Smith & Smith, 2010) Genus: †Oodectes ^{(paraphyletic genus)} (Wortman, 1901) Genus: †Paramiacis (Mathis, 1985) Genus: †Paroodectes (Springhorn, 1980) Incertae sedis: †"Miacis" sp. [CM 67873 & CM 77299] (Beard & Dawson, 2009) Genus: †Messelogale (Springhorn, 2000) Genus: †Miocyon (Matthew, 1909) Genus: †Simamphicyon (Viret, 1942) Genus: †Uintacyon ^{(paraphyletic genus)} (Leidy, 1872) Genus: †Xinyuictis (Zheng, 1975) Genus: †Zodiocyon (Tong & Wang, 2006) (unranked): Clade "C" Genus: †Dormaalocyon (Solé, 2014) (unranked): †Vulpavus clade Genus: †Palaearctonyx (Matthew, 1909) Genus: †Vassacyon (Matthew, 1909) Genus: †Vulpavus ^{(paraphyletic genus)} (Marsh, 1871) Incertae sedis: †"Miacis" deutschi (Gingerich, 1983) †"Miacis" exiguus (Matthew & Granger, 1915) (unranked): Clade "D" Order: Carnivora [sensu stricto] (Bowdich, 1821) (carnivorans) [= Clade "E"] Genus: †Ceruttia (Tomiya, 2013) Genus: †Harpalodon (Marsh, 1872) Genus: †Lycarion (Matthew, 1909) Genus: †Neovulpavus (Wortman, 1901) Genus: †Procynodictis (Wortman & Matthew, 1899) Genus: †Prodaphaenus (Wortman & Matthew, 1899) Genus: †Tapocyon (Stock, 1934) Genus: †Walshius (Tomiya, 2013) Incertae sedis: †"Miacis" gracilis (Clark, 1939) †"Miacis" hargeri (Wortman, 1901) †"Miacis" invictus (Matthew & Granger, 1925) †"Miacis" lushiensis (Chow, 1975) †Carnivoraformes sp. [SDSNH 49600] (Poust & Tomiya, 2024) Incertae sedis: †"Miacis" boqinghensis (Huang, 1999) †"Miacis" hookwayi (Stock, 1934) †"Miacis" latidens (Matthew & Granger, 1915) †"Miacis" petilus (Gingerich, 1983) †Carnivoraformes undet. Genus A (Tomiya, 2013) †Carnivoraformes undet. Genus B (Tomiya, 2013) ichnotaxa of Carnivoraformes: Ichnogenus: †Falcatipes (Sargeant & Langston, 1994) (unranked): †Viverravidae [sensu lato] Superfamily: †Viverravoidea (Wortman & Matthew, 1899) Family: †Viverravidae [sensu stricto] (Wortman & Matthew, 1899) Genus: †Ravenictis (Fox & Youzwyshyn, 1994) Incertae sedis: †Carnivoramorpha sp. [UALVP 50993 & UALVP 50994] (Fox, Scott & Rankin, 2010) Incertae sedis: †"Sinopa" insectivorus (Cope, 1872) †Carnivoramorpha sp. [UALVP 31176] (Fox & Youzwyshyn, 1994) †Carnivoramorpha sp. [USNM 538395] (Rose, 2012)

References

↑ Wyss, A. R. & Flynn, J. J. (1993.) "A Phylogenetic Analysis and Definition of the Carnivora." in "Mammal Phylogeny – Placentals", Szalay, F. S., M. J. Novacek and M. C. McKenna (eds.). ISBN 978-0-387-97853-6
↑ K. D. Rose, A. E. Chew, R. H. Dunn, M. J. Kraus, H. C. Fricke and S. P. Zack (2012.) "Earliest Eocene mammalian fauna from the Paleocene-Eocene Thermal Maximum at Sand Creek Divide, southern Bighorn Basin, Wyoming." University of Michigan Papers on Paleontology 36:1-122
↑ Matsui, K.; Kimura, Y. (2022). "A "Mammalian-like" Pycnodont Fish: Independent Acquisition of Thecodont Implantation, True Vertical Replacement, and Carnassial Dentitions in Carnivorous Mammals and a Peculiar Group of Pycnodont Fish". Life. 12 (2) 250. Bibcode:2022Life...12..250M. doi:10.3390/life12020250. PMC 8878644. PMID 35207537.
↑ Bryant, H.N., and M. Wolson (2004) "Phylogenetic Nomenclature of Carnivoran Mammals." First International Phylogenetic Nomenclature Meeting. Paris, Museum National d'Histoire Naturelle
↑ Solé, Floréal; Smith, Richard; Coillot, Tiphaine; de Bast, Eric; Smith, Thierry (2014). "Dental and tarsal anatomy of Miacis latouri and a phylogenetic analysis of the earliest carnivoraforms (Mammalia, Carnivoramorpha)". Journal of Vertebrate Paleontology. 34 (1): 1–21. doi:10.1080/02724634.2013.793195. ISSN 0272-4634. S2CID 86207013.
1 2 Zack, Shawn P.; Rose, Kenneth D.; O'Leary, Maureen A. (2025-06-24). "New Cranial and Postcranial Remains of the Once Enigmatic Early Eocene Mammal Wyolestes (Mammalia, Ferae, Hyaenodonta) from North America, and Phylogenetic Evidence for its Interordinal Relationships". Bulletin of the American Museum of Natural History. 2025 (475). doi:10.1206/0003-0090.475.1.1. ISSN 0003-0090.
1 2 Zack, Shawn P. (2019). "A skeleton of a Uintan machaeroidine 'creodont' and the phylogeny of carnivorous eutherian mammals". Journal of Systematic Palaeontology. 19 (8): 653–689. doi:10.1080/14772019.2018.1466374.
1 2 Zack, Shawn (2026). "A new species of Simidectes (Mammalia: Ferae) from the early Uintan of California and new evidence of the affinities of the genus". Historical Biology. doi:10.1080/08912963.2026.2646998.
↑ Floréal Solé & Thierry Smith (2013.) "Dispersals of placental carnivorous mammals (Carnivoramorpha, Oxyaenodonta & Hyaenodontida) near the Paleocene-Eocene boundary: a climatic and almost worldwide story" Geologica Belgica 16/4: 254-261
↑ Flynn, John J.; Finarelli, John A.; Spaulding, Michelle (2010). "Phylogeny of the Carnivora and Carnivoramorpha, and the use of the fossil record to enhance understanding of evolutionary transformations". In Goswami, Anjali; Friscia, Anthony (eds.). Carnivoran evolution. New views on phylogeny, form and function. Cambridge University Press. pp. 25–63. doi:10.1017/CBO9781139193436.003. ISBN 978-1-139-19343-6.

C. M. Janis, J. A. Baskin, A. Berta, J. J. Flynn, G. F. Gunnell, R. M. Hunt jr., L. D. Martin, and K. Munthe (1998.) "Carnivorous mammals." In C. M. Janis, K. M. Scott, and L. L. Jacobs (eds.) "Evolution of Tertiary Mammals of North America. Volume 1: Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals." Cambridge University Press, Cambridge. ISBN 9780521355193
K. D. Rose and J. D. Archibold (2005) "The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades", Baltimore and London, Johns Hopkins University Press ISBN 978-0-8018-8022-3
K. D. Rose and J. D. Archibold (2005) "Womb with a View: the Rise of Placentals." In: K. D. Rose and J. D. Archibold "The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades", Baltimore and London, Johns Hopkins University Press ISBN 978-0-8018-8022-3
Welsey-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): 1–28. doi:10.1017/S1477201904001518. S2CID 86755875.
Stiles, David P. (2005) "An investigation of the Vulpes and Urocyon phylogenetic classification: Feliformia or Caniformia?" Fox Phylogeny. Vertebrate Evolution – Fall 2005, The Evergreen State College, Olympia, WA.
Wesley-Hunt, G. D.; Werdelin, L. (2005). "Basicranial morphology and phylogenetic position of the upper Eocene carnivoramorphan Quercygale". Acta Palaeontologica Polonica. 50 (4): 837–846.
Wesley-Hunt, Gina D. (2005) "The Morphological Diversification of Carnivores in North America." Paleobiology. Vol. 31, Issue 1, pp. 35–55.
Benton, Michael J. and Philip C. J. Donoghue (2007) "Paleontological Evidence to Date the Tree of Life.", Molecular Biology and Evolution, Volume 24, Issue 1, Pages 26–53
Spaulding, M.; O'Leary, M. A.; Gatesy, J. (2009). Farke, Andrew Allen (ed.). "Relationships of Cetacea (Artiodactyla) Among Mammals: Increased Taxon Sampling Alters Interpretations of Key Fossils and Character Evolution". PLOS ONE. 4 (9) e7062. Bibcode:2009PLoSO...4.7062S. doi:10.1371/journal.pone.0007062. PMC 2740860. PMID 19774069.
Jackson, S.; Jackson, S. M.; Groves, C. (2015). Taxonomy of Australian mammals. Csiro Publishing. p. 238. ISBN 978-1-4863-0013-6.
Susumu Tomiya, Shawn P. Zack, Michelle Spaulding and John J. Flynn (2019.) "Carnivorous mammals from the Middle Eocene Washakie formation, Wyoming, U.S.A., and their diversity trajectory in a post-warming world", in "The Society of Vertebrate Paleontology 79th annual meeting"
Solé, F.; Fischer, V.; Le Verger, K.; Mennecart, B.; Speijer, R. P.; Peigné, S.; Smith, T. (2022). "Evolution of European carnivorous mammal assemblages through the Palaeogene". Biological Journal of the Linnean Society. 135 (4): 734–753. doi:10.1093/biolinnean/blac002.

[1] Wyss, A. R. & Flynn, J. J. (1993.) "A Phylogenetic Analysis and Definition of the Carnivora." in "Mammal Phylogeny – Placentals", Szalay, F. S., M. J. Novacek and M. C. McKenna (eds.). ISBN 978-0-387-97853-6

[2] K. D. Rose, A. E. Chew, R. H. Dunn, M. J. Kraus, H. C. Fricke and S. P. Zack (2012.) "Earliest Eocene mammalian fauna from the Paleocene-Eocene Thermal Maximum at Sand Creek Divide, southern Bighorn Basin, Wyoming." University of Michigan Papers on Paleontology 36:1-122

[3] Matsui, K.; Kimura, Y. (2022). "A "Mammalian-like" Pycnodont Fish: Independent Acquisition of Thecodont Implantation, True Vertical Replacement, and Carnassial Dentitions in Carnivorous Mammals and a Peculiar Group of Pycnodont Fish". Life. 12 (2) 250. Bibcode:2022Life...12..250M. doi:10.3390/life12020250. PMC 8878644. PMID 35207537.

[Bryant2004-4] Bryant, H.N., and M. Wolson (2004) "Phylogenetic Nomenclature of Carnivoran Mammals." First International Phylogenetic Nomenclature Meeting. Paris, Museum National d'Histoire Naturelle

[SoléSmith2014-5] Solé, Floréal; Smith, Richard; Coillot, Tiphaine; de Bast, Eric; Smith, Thierry (2014). "Dental and tarsal anatomy of Miacis latouri and a phylogenetic analysis of the earliest carnivoraforms (Mammalia, Carnivoramorpha)". Journal of Vertebrate Paleontology. 34 (1): 1–21. doi:10.1080/02724634.2013.793195. ISSN 0272-4634. S2CID 86207013.

[:1-6] 1 2 Zack, Shawn P.; Rose, Kenneth D.; O'Leary, Maureen A. (2025-06-24). "New Cranial and Postcranial Remains of the Once Enigmatic Early Eocene Mammal Wyolestes (Mammalia, Ferae, Hyaenodonta) from North America, and Phylogenetic Evidence for its Interordinal Relationships". Bulletin of the American Museum of Natural History. 2025 (475). doi:10.1206/0003-0090.475.1.1. ISSN 0003-0090.

[:0-7] 1 2 Zack, Shawn P. (2019). "A skeleton of a Uintan machaeroidine 'creodont' and the phylogeny of carnivorous eutherian mammals". Journal of Systematic Palaeontology. 19 (8): 653–689. doi:10.1080/14772019.2018.1466374.

[:2-8] 1 2 Zack, Shawn (2026). "A new species of Simidectes (Mammalia: Ferae) from the early Uintan of California and new evidence of the affinities of the genus". Historical Biology. doi:10.1080/08912963.2026.2646998.

[9] Floréal Solé & Thierry Smith (2013.) "Dispersals of placental carnivorous mammals (Carnivoramorpha, Oxyaenodonta & Hyaenodontida) near the Paleocene-Eocene boundary: a climatic and almost worldwide story" Geologica Belgica 16/4: 254-261

[asdff-10] Flynn, John J.; Finarelli, John A.; Spaulding, Michelle (2010). "Phylogeny of the Carnivora and Carnivoramorpha, and the use of the fossil record to enhance understanding of evolutionary transformations". In Goswami, Anjali; Friscia, Anthony (eds.). Carnivoran evolution. New views on phylogeny, form and function. Cambridge University Press. pp. 25–63. doi:10.1017/CBO9781139193436.003. ISBN 978-1-139-19343-6.

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