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 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

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)
    • 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

 PanCarnivora 

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.
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