Carnivoramorpha

Carnivoramorpha ("shape of carnivoran") is a clade of mammals that includes the modern order Carnivora and its extinct stem-relatives.

Carnivoramorpha Temporal range: 66.043–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N early Paleocene to present
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Ferae
Clade:	Carnivoramorpha Bryant, 1996
Subgroups
[see classification]

Life reconstruction of Tapocyon robustus

Classification and phylogeny

Traditional classification

• Clade: Carnivoramorpha (basal to Carnivora but without Creodonts)
  • Superfamily: †Miacoidea
    • Family: †Miacidae
      • Genera: †Chailicyon, †Eosictis, †Miacis, †Miocyon, †Oodectes, †Palaearctonyx, †Paramiacis, †Paroodectes, †Prodaphaenus, †Quercygale, †Tapocyon, †Uintacyon, †Vassacyon, †Vulpavus and †Xinyuictis
    • Family: †Viverravidae
      • Genera: †Bryanictis, †Didymictis, †Ictidopappus, †Palaeogale, †Pappidictops, †Pristinictis, †Protictis, †Raphictis, †Simpsonictis and †Viverravus
  • Order: Carnivora (carnivorans)
    • Suborder: Caniformia [or Canoidea] ("dog-like" carnivorans)
    • Suborder: Feliformia [or Feloidea] ("cat-like" carnivorans)

Revised classification

As 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.[1] 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).

• Clade: Carnivoramorpha (Bryant, 1996)
  • Superfamily: †Viverravoidea (Wortman & Matthew, 1899)
    • Family: †Viverravidae (Wortman & Matthew, 1899)
  • Incertae sedis:
    • Genus: †Ravenictis (Fox & Youzwyshyn, 1994)
      • †Ravenictis krausei (Fox & Youzwyshyn, 1994)
  • Clade: Carnivoraformes (Flynn, 2010)
    • Genus: †Africtis (Mattingly, 2020)
      • †Africtis sirtensis (Mattingly, 2020)
    • Genus: †Ceruttia (Tomiya, 2013)
      • †Ceruttia sandiegoensis (Tomiya, 2013)
    • Genus: †Dawsonicyon (Spaulding, Flynn & Stucky, 2010)
      • †Dawsonicyon isami (Spaulding, Flynn & Stucky, 2010)
    • Genus: †Miacis (Cope, 1872)
      • †Miacis parvivorus (Cope, 1872)
    • Genus: †Prodaphaenus (Wortman & Matthew, 1899)
      • †Prodaphaenus uintensis (Osborn, 1895)
      • †Prodaphaenus vulpinus (Scott & Osborn, 1887)
    • Genus: †Walshius (Tomiya, 2013)
      • †Walshius pacificus (Tomiya, 2013)
    • (unranked): †Lycarion/Neovulpavus clade
      • Genus: †Lycarion (Matthew, 1909)
        • †Lycarion hargeri (Wortman, 1901)
        • †Lycarion medius (Matthew, 1909)
      • Genus: †Neovulpavus (Wortman, 1901)
        • †Neovulpavus washakius (Wortman, 1901)
    • (unranked): †Oodectes clade
      • Genus: †Eogale (Beard & Dawson, 2009)
        • †Eogale parydros (Beard & Dawson, 2009)
      • Genus: †Gracilocyon (Smith & Smith, 2010)
        • †Gracilocyon igniculus (Beard & Dawson, 2009)
        • †Gracilocyon rosei (Heinrich, 2008)
        • †Gracilocyon rundlei (Hooker, 2010)
        • †Gracilocyon solei (Smith & Smith, 2010)
        • †Gracilocyon winkleri (Gingerich, 1983)
      • Genus: †Oodectes (Wortman, 1901)
        • †Oodectes herpestoides (Wortman, 1901)
        • †Oodectes jepseni (Guthrie, 1967)
      • Genus: †Paramiacis (Mathis, 1985)
        • †Paramiacis exilis (Filhol, 1876)
        • †Paramiacis teilhardi (Mathis, 1987)
      • Genus: †Paroodectes (Springhorn, 1980)
        • †Paroodectes feisti (Springhorn, 1980)
    • (unranked): †Uintacyon clade
      • Genus: †Miocyon (Matthew, 1909)
        • †Miocyon bathygnathus (Scott, 1888)
        • †Miocyon magnus (Bryant, 1992)
        • †Miocyon major (Matthew, 1909)
        • †Miocyon scotti (Wortman & Matthew, 1899)
        • †Miocyon vallisrubrae (Friscia & Rasmussen, 2010)
      • Genus: †Simamphicyon (Viret, 1942)
        • †Simamphicyon helveticus (Pictet & Humbert, 1869)
      • Genus: †Uintacyon (Leidy, 1873)
        • †Uintacyon acutus (Thorpe, 1923)
        • †Uintacyon asodes (Gazin, 1952)
        • †Uintacyon edax (Leidy, 1873)
        • †Uintacyon gingerichi (Heinrich, 2008)
        • †Uintacyon hookeri (Solé, 2014)
        • †Uintacyon jugulans (Matthew, 1909)
        • †Uintacyon massetericus (Cope, 1882)
        • †Uintacyon rudis (Rose, 1981)
        • †Uintacyon vorax (Leidy, 1873)
        • Subgenus: †Xinyuictis (Zheng, 1975)
          • †Uintacyon tenuis (Zheng, 1975)
      • Genus: †Zodiocyon (Tong & Wang, 2006)
        • †Zodiocyon zetesios (Tong & Wang, 2006)
      • Incertae sedis:
        • †"Miacis" boqinghensis (Huang, 1999)
        • †"Miacis" invictus (Matthew & Granger, 1925)
        • †"Miacis" lushiensis (Chow, 1975)
        • †"Miacis" thailandicus (Ducrocq, 1992)
    • (unranked): †Vulpavus clade
      • Genus: †Dormaalocyon (Solé, 2014)
        • †Dormaalocyon latouri (Quinet, 1966)
      • Genus: †Vassacyon (Matthew, 1909)
        • †Vassacyon bowni (Heinrich, 2008)
        • †Vassacyon prieuri (Solé, 2016)
        • †Vassacyon promicrodon (Wortman & Matthew, 1899)
        • †Vassacyon taxidiotis (Solé, 2013)
      • Genus: †Vulpavus (Marsh, 1871)
        • †Vulpavus australis (Matthew & Granger, 1915)
        • †Vulpavus canavus (Cope, 1881)
        • †Vulpavus completus (Matthew, 1909)
        • †Vulpavus farsonensis (Gunnell, 1998)
        • †Vulpavus palustris (Marsh, 1871)
        • †Vulpavus profectus (Matthew, 1909)
        • Subgenus: †Palaearctonyx (Matthew, 1909)
          • †Vulpavus meadi (Matthew, 1909)
        • Subgenus: †Phlaodectes (Matthew, 1909)
          • †Vulpavus ovatus (Matthew, 1909)
      • Incertae sedis:
        • †"Miacis" deutschi (Gingerich, 1983)
        • †"Miacis" exiguus (Matthew & Granger, 1915)
    • Incertae sedis:
      • †"Miacis" hookwayi (Stock, 1934)
      • †"Miacis" latidens (Matthew & Granger, 1915)
      • †"Miacis" petilus (Gingerich, 1983)
    • Order: Carnivora (Bowdich, 1821) (carnivorans)
      • Suborder: Caniformia (Kretzoi, 1943) ("dog-like" carnivorans)
      • Suborder: Feliformia (Kretzoi, 1945) ("cat-like" carnivorans)

Phylogenetic tree

The phylogenetic relationships of Carnivoramorpha are shown in the following cladogram:[2][1][3][4][5][6][7][8][9][10]

Ferae

Pholidotamorpha    Pan‑Carnivora    †Oxyaenodonta      †Hyaenodonta    †Hyaenodonta (sensu stricto)       †Altacreodus     †Tinerhodon   †Altacreodus/Tinerhodon clade    sensu lato   Carnivoramorpha   †Viverravidae   †Viverravoidea  †Viverravidae (sensu stricto)    ?  †carnivoramorph sp. (UALVP 50993 & UALVP 50994)    ?  †Ravenictis    sensu lato   ?  †carnivoramorph sp. (UALVP 31176)    ?  †carnivoramorph sp. (WW-84: USNM 538395)    Carnivoraformes      †Neovulpavus    ?  †carnivoraform undet. Genus A (UCMP 110072)     †Lycarion   †Lycarion/Neovulpavus clade    ?  †carnivoraform undet. Genus B (SDSNH 56335)    ?  †Ceruttia               †Dormaalocyon     †"Miacis" exiguus         †Vassacyon     †Vulpavus         †"Miacis" deutschi   †Vulpavus clade         †Miocyon     †Simamphicyon       †Zodiocyon       †Uintacyon    ?  †"Miacis" boqinghensis    ?  †"Miacis" lushiensis    ?  †"Miacis" thailandicus    ?  †"Miacis" invictus     †Uintacyon clade     †Miacis     †carnivoraform sp. (PM 3868)     †Dawsonicyon    ?  †Africtis     †"Miacis" petilus     †"Miacis" latidens    ?  †"Miacis" hookwayi       †Paroodectes     †Paramiacis     †Eogale       †Gracilocyon     †Oodectes     †Gracilocyon/Oodectes clade         †Prodaphaenus    ?  †Walshius    Carnivora   Feliformia        Aeluroidea     †Nimravidae      ?  †Eosictis    ?  †Chailicyon    †Palaeogalidae    †Palaeogale     †Cryptailurus      †Quercygalidae    †Messelogale     †Quercygale         †Tapocyon      Caniformia    †Harpalodon       †Procynodictis       †Amphicyonoidea       †Lycophocyon     Canoidea            sensu stricto         (Carnivora [sensu lato])

References

1. John J. Flynn; John A. Finarelli; Michelle Spaulding (2010). "Phylogeny of the Carnivora and Carnivoramorpha, and the use of the fossil record to enhance understanding of evolutionary transformations". In Anjali Goswami; Anthony Friscia (eds.). Carnivoran evolution. New views on phylogeny, form and function. Cambridge University Press. pp. 25–63. doi:10.1017/CBO9781139193436.003. ISBN 9781139193436.
2. Bryant, H.N., and M. Wolson (2004) “Phylogenetic Nomenclature of Carnivoran Mammals.” First International Phylogenetic Nomenclature Meeting. Paris, Museum National d’Histoire Naturelle
3. Michelle Spaulding; John J. Flynn; Richard K. Stucky (2010). "A new basal Carnivoramorphan (Mammalia) from the 'Bridger B' (Black's Fork member, Bridger Formation, Bridgerian Nalma, middle Eocene) of Wyoming, USA". Palaeontology. 53 (4): 815–832. doi:10.1111/j.1475-4983.2010.00963.x.
4. Susumu Tomiya (2011). "A new basal caniform (Mammalia: Carnivora) from the Middle Eocene of North America and remarks on the phylogeny of early carnivorans". PLOS One. 6 (9): e24146. doi:10.1371/journal.pone.0024146. PMC 3173397. PMID 21935380.
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.
6. Wang, Jian; Zhang, Zhaoqun (2015). "Phylogenetic analysis on Palaeogale (Palaeogalidae, Carnivora) based on specimens from Oligocene strata of Saint-Jacques, Nei Mongol". Vertebrata PalAsiatica. 53 (4): 310–334.
7. 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.
8. Tomiya, Susumu; Tseng, Zhijie Jack (2016). "Whence the beardogs? Reappraisal of the Middle to Late Eocene 'Miacis' from Texas, USA, and the origin of Amphicyonidae (Mammalia, Carnivora)". Royal Society Open Science. 3 (10): 160518. Bibcode:2016RSOS....3p0518T. doi:10.1098/rsos.160518. ISSN 2054-5703. PMC 5098994. PMID 27853569.
9. Solé, F. & Ladevèze, S. (2017). "Evolution of the hypercarnivorous dentition in mammals (Metatheria, Eutheria) and its bearing on the development of tribosphenic molars." Evolution & Development, 19(2), 56–68.
10. Prevosti, F. J., & Forasiepi, A. M. (2018). "Introduction. Evolution of South American Mammalian Predators During the Cenozoic: Paleobiogeographic and Paleoenvironmental Contingencies"

• Archibold, J.D., and K.D. Rose (eds.). (2005). “The Rise of Placental Mammals: Origins and Relationships of the Major Clades.” Baltimore and London: The Johns Hopkins University Press. ISBN 0-8018-8022-X
• Archibold, J.D., and K.D. Rose. (2005). “Womb with a View: the Rise of Placentals.”
• Benton, Michael J. and Philip C.J. Donoghue. (2007). “Paleontological Evidence to Date the Tree of Life.” Dating the Tree of Life 31.
• Bryant, H.N., and M. Wolson (2004) “Phylogenetic Nomenclature of Carnivoran Mammals.” First International Phylogenetic Nomenclature Meeting. Paris, Museum National d’Histoire Naturelle, July 6–9, 2004.
• Flynn, John J. and Gina D. Wesley-Hunt. 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. Abstract:
• Haaramo, Mikko. (2005). “Mikko's Phylogeny Archive Carnivoramorpha - Carnivores and relatives.” Retrieved February 17, 2007, from:
• 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, Gina D. (2006). “The Morphological Diversification of Carnivores in North America.” Paleobiology. Vol. 31, Issue 1, pp. 35–55.
• 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 0-387-97853-4.
• John J. Flynn; John A. Finarelli; Michelle Spaulding (2010). "Phylogeny of the Carnivora and Carnivoramorpha, and the use of the fossil record to enhance understanding of evolutionary transformations". In Anjali Goswami; Anthony Friscia (eds.). Carnivoran evolution. New views on phylogeny, form and function. Cambridge University Press. pp. 25–63. doi:10.1017/CBO9781139193436.003. ISBN 9781139193436.