# Ferae

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Clade of mammals consisting of carnivorans and pholidotes

Ferae Temporal range: 79.47–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Late Cretaceous to present[1] Scientific classification Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Placentalia Grandorder: Ferungulata Mirorder: Ferae Linnaeus, 1758[2] Main subgroups Pholidotamorpha (pangolins and extinct relatives) Pan-Carnivora †Hyaenodonta †Oxyaenidae Carnivoramorpha For other groups, see text Synonyms list of synonyms: Carnaria (Haeckel, 1866)[3] Carnassia (Haeckel, 1895)[4] Carnivora (Zagorodniuk, 2008)[5] Carnivoramorpha (Kalandadze & Rautian, 1992)[6] Ferina (Newman, 1843)[7] Ostentoria (Amrine-Madsen, 2003)[8] Rapacia (Newman, 1843) Sarcotheria (Haeckel, 1895)

**Ferae** ([/ˈfɪəriː/](https://en.wikipedia.org/wiki/Help:IPA/English) [*FEER-ee*](https://en.wikipedia.org/wiki/Help:Pronunciation_respelling_key), Latin: [\[ˈfɛrae̯\]](https://en.wikipedia.org/wiki/Help:IPA/Latin), "wild beasts") is a [mirorder](/source/Mirorder) of [placental mammals](/source/Placentalia)[9][10] in grandorder [Ferungulata](/source/Ferungulata), that groups together clades **Pan-Carnivora** (that includes [carnivorans](/source/Carnivora) and their fossil relatives, such as [hyaenodonts](/source/Hyaenodonta)) and [Pholidotamorpha](/source/Pholidotamorpha) ([pangolins](/source/Pangolin) and their fossil relatives).

## General characteristics

### In mirorder Ferae

The common features for members of this mirorder are:[11][12]

- ossified [tentorium cerebelli](/source/Tentorium_cerebelli),

- fusion of the [scaphoid](/source/Scaphoid) and [lunate](/source/Lunate_bone) bones in the wrist,

- and the a network of [diploic venous](/source/Diploic_veins) channels throughout their [cranial vault](/source/Cranial_vault).[13]

forefoot of *[Patriomanis americana](/source/Patriomanis)*

forefoot of *[Vulpavus profectus](/source/Vulpavus)*

forefoot of
*[Thinocyon medius](https://en.wikipedia.org/w/index.php?title=Thinocyon&action=edit&redlink=1)*

forefoot of *[Patriofelis ferox](/source/Patriofelis)*

### In clade Pan-Carnivora

The common features for members of clade Pan-Carnivora are:

- [binocular vision](/source/Binocular_vision),

- sharp claws,

- [heterodont](/source/Heterodont) teeth that are sharp and for cutting meat,

- canine teeth that are usually large, conical, pointed, thick and stress resistant,

- and presence of the [carnassial teeth](/source/Carnassial).

Carnassials are feature that allows distinguishing the [Carnivoramorpha](/source/Carnivoramorpha), [Oxyaenodonta](/source/Oxyaenidae) and [Hyaenodonta](/source/Hyaenodonta) from the other carnivorous placental mammals.[11] However, these mammals are distinguished between themselves based on the position of the carnassial teeth and the number of molars. The carnassial teeth of the Carnivoramorpha are located in P4 and m1, in Oxyaenodonta are M1 and m2, and in Hyaenodonta and close relatives are M2 and m3. This appears to be a case of a possible evolutionary convergent adaptation toward similar diet.[11]

skeleton of [wolf](/source/Wolf)

skeleton of *[Hyaenodon horridus](/source/Hyaenodon)*

skeleton of *[Patriofelis ferox](/source/Patriofelis)*

diagram of a wolf skull with key features labelled

Comparison of carnassial teeth of a carnivoran (wolf), a hyaenodontid (*[Hyaenodon](/source/Hyaenodon)*) and an oxyaenid (*[Oxyaena](/source/Oxyaena)*)

skull and jaws of
*[Simidectes merriami](https://en.wikipedia.org/w/index.php?title=Simidectes&action=edit&redlink=1)*

## Classification and phylogeny

### Sister groups to Ferae

According to recent studies, the closest relatives of Ferae are members of clade [Pan-Euungulata](/source/Pan-Euungulata) (group that includes mirorder [Euungulata](/source/Ungulate)[14][15] and extinct genus *[Protungulatum](/source/Protungulatum)*). Together they form grandorder [Ferungulata](/source/Ferungulata).

An alternate phylogeny holds that the closest relative to Ferae is order [Perissodactyla](/source/Perissodactyla), with whom they form a clade [Zooamata](/source/Zooamata). Together, clade Zooamata and order [Chiroptera](/source/Bat) form clade [Pegasoferae](/source/Pegasoferae),[16] and Pegasoferae is sister taxon to order [Artiodactyla](/source/Artiodactyl) within clade [Scrotifera](/source/Scrotifera). However, subsequent molecular studies have generally failed to support this proposal.[17][18][19][20][21]

### Position of pangolins and creodonts within clade

[Pangolins](/source/Pangolin) were long thought to be the closest relatives of [aardvark](/source/Aardvark) and [xenarthrans](/source/Xenarthra), forming to the now obsolete order [Edentata](/source/Edentata). Research based on immunodiffusion technique[22] and comparison of protein and DNA sequences[23][24][25] revealed the close relationships between pangolins and [carnivorans](/source/Carnivora), with whom they also share a few unusual derived morphological and anatomical traits, such as the ossified [tentorium cerebelli](/source/Tentorium_cerebelli) and the fusion of the [scaphoid](/source/Scaphoid) and [lunate](/source/Lunate_bone) bones in the wrist. The [last common ancestor](/source/Last_common_ancestor) of extant Ferae is supposed to have diversified c. 79.47 million years ago.[1]

While there has been strong support in the inclusion of order [Creodonta](/source/Creodonta) into Ferae, they were usually recovered as sister taxon to order Carnivora.[9] The *Halliday et al. (2015)* phylogenetic analysis of hundreds of morphological characters of [Paleocene](/source/Paleocene) placentals found instead that creodonts might be the sister group to [Pholidotamorpha](/source/Pholidotamorpha) (pangolins and their stem-relatives).[26] However, recent studies have shown that Creodonta is an invalid [polyphyletic](/source/Polyphyletic) taxon. Members of this group are now part of clade Pan-Carnivora and sister taxa to [Carnivoramorpha](/source/Carnivoramorpha) (carnivorans and their stem-relatives), split in two groups: order [Oxyaenodonta](/source/Oxyaenidae) on one side and on the other side order [Hyaenodonta](/source/Hyaenodonta) plus its stem-relatives, genera *[Simidectes](https://en.wikipedia.org/w/index.php?title=Simidectes&action=edit&redlink=1)*[27][28] and *[Altacreodus](/source/Altacreodus)*.[29][30][31][32][33]

### Taxonomy

Former classification (McKenna & Bell, 1997):[10] Current classification: Grandorder: Ferae (Linnaeus, 1758) Order: Carnivora (Bowdich, 1821) (carnivorans) Order: Cimolesta (McKenna, 1975) Suborder: Pholidota (Weber, 1904) (pangolins) Suborder: †Apatotheria (Scott & Jepsen, 1936) Suborder: †Didelphodonta (McKenna, 1975) Suborder: †Ernanodonta (Ding, 1987) Suborder: †Pantodonta (Cope, 1873) Suborder: †Pantolesta (McKenna, 1975) Suborder: †Taeniodonta (Cope, 1876) Suborder: †Tillodontia (Marsh, 1875) Family: †Palaeoryctidae (Winge, 1917) "Subfamily:" †Wyolestinae (Gingerich, 1981) Genus: †Alostera (Fox, 1989) Genus: †Avitotherium (Cifelli, 1990) Genus: †Pararyctes (Van Valen, 1966) Genus: †Ravenictis (Fox & Youzwyshyn, 1994) Order: †Creodonta (Cope, 1875) (false carnivorans) Family: †Hyaenodontidae (Leidy, 1869) Family: †Oxyaenidae (Cope, 1877) "Subfamily:" †Koholiinae (Crochet, 1988) Genus: †Prionogale (Schmidt-Kittler & Heizmann, 1991) Mirorder: Ferae (Linnaeus, 1758) Clade: Pan-Carnivora (Flynn, Wyss & Wolsan, 2020)[34] (carnivorans and stem-relatives) Clade: Carnivoramorpha (Wyss & Flynn, 1993) (carnivoran-like mammals) [= Carnivora (sensu lato)] Order: †Oxyaenodonta (Van Valen, 1971) Family: †Oxyaenidae (Cope, 1877) (unranked): †Hyaenodonta [sensu lato] Order: †Hyaenodonta [sensu stricto] (Van Valen, 1967) Genus: †Altacreodus (Fox, 2015) Genus: †Simidectes (Stock, 1933) Clade: Pholidotamorpha (Gaudin, 2009) (pangolin-like mammals) Order: Pholidota (Weber, 1904) (pangolins) Order: †Afredentata (Szalay & Schrenk, 1994) Order: †Palaeanodonta (Matthew, 1918) (stem-pangolins) Genus: †Euromanis (Gaudin, 2009) (?) Order: †Pantolesta (McKenna, 1975)

#### Alternative classification and possible fossil members

In *Halliday et al. (2015)* various enigmatic Palaeocene [eutherian](/source/Eutheria) mammals have been proposed to be possible members of Ferae, like members of orders [Mesonychia](/source/Mesonychia), [Pantodonta](/source/Pantodonta) and [Taeniodonta](/source/Taeniodonta), and families [Arctocyonidae](/source/Arctocyonidae), [Didelphodontidae](https://en.wikipedia.org/w/index.php?title=Didelphodontidae&action=edit&redlink=1), [Nyctitheriidae](/source/Nyctitheriidae), [Palaeoryctidae](/source/Palaeoryctidae), [Periptychidae](/source/Periptychidae) and [Triisodontidae](/source/Triisodontidae).[26] Mesonychians are proposed to be a sister group to carnivoramorphs, while arctocyonids were [polyphyletic](/source/Polyphyletic), with genera *[Arctocyon](/source/Arctocyon)* and *[Loxolophus](/source/Loxolophus)* as a sister taxa to pantodonts and periptychids, *[Goniacodon](/source/Goniacodon)* and *[Eoconodon](/source/Eoconodon)* sister to the Carnivoramorpha-Mesonychia clade, and other genera allied with creodonts and palaeoryctids.[26] This enlarged Ferae was also found to be the sister group to order [Chiroptera](/source/Bat),[26] even though recent studies dispute this classification.[35][36]

## See also

- [Mammal classification](/source/Mammal_classification)

- [Ferungulata](/source/Ferungulata)

## References

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v t e Mammals of clade Pan-Carnivora Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Placentalia Superorder: Laurasiatheria Clade: Scrotifera Grandorder: Ferungulata Mirorder: Ferae Pan-Carnivora Pan-Carnivora Carnivoramorpha See Carnivoramorpha below ↓ †Hyaenodonta See Hyaenodonta below ↓ †Oxyaenodonta See Oxyaenodonta below ↓ †Wyolestidae Wyolestes †Simidectes †Altacreodus †Ravenictis †Tinerhodon †Hyaenodonta Hyaenodonta Arfia Boualitomus Eoproviverra Galecyon Gazinocyon Koholia Lahimia Parvavorodon Pyrocyon Tritemnodon Hyaenodontidae Alienetherium Boritia Cartierodon Consobrinus Cynohyaenodon Eurotherium Hyaenodon Leonhardtina Matthodon Neosinopa Oxyaenoides Paenoxyaenoides Paracynohyaenodon Praecodens Preregidens Prodissopsalis Propterodon Protoproviverra Quercytherium Hyainailouridae Apterodontinae Apterodon Quasiapterodon Hyainailourinae Akhnatenavus Bastetodon Exiguodon Falcatodon Hemipsalodon Hyainailouros Ischnognathus Isohyaenodon Kerberos Leakitherium Maocyon Megistotherium Mlanyama Metapterodon Orienspterodon Pakakali Parapterodon Paroxyaena Pterodon Sectisodon Sekhmetops Simbakubwa Sivapterodon Indohyaenodontidae Glibzegdouia Indohyaenodon Yarshea Limnocyonidae Iridodon Limnocyon Oxyaenodon Prolaena Prolimnocyon Thinocyon Prionogalidae Namasector Prionogale Thereutherium Proviverridae Allopterodon Lesmesodon Minimovellentodon Morlodon Parvagula Proviverra Sinopinae Acarictis Prototomus Sinopa Teratodontidae Anasinopa Brychotherium Buhakia Dissopsalis Ekweeconfractus Furodon Kyawdawia Masrasector Metasinopa Paratritemnodon Teratodon ichnotaxa of Hyaenodonta Creodontipus Dischidodacylus Sarcotherichnus Zanclonychopus Hyaenodontipus Quiritipes Sarjeantipes †Oxyaenodonta Oxyaenidae Machaeroidinae Apataelurus Diegoaelurus Isphanatherium Machaeroides Oxyaeninae Argillotherium Dipsalidictis Malfelis Oxyaena Patriofelis Protopsalis Sarkastodon Palaeonictinae Ambloctonus Dipsalodon Palaeonictis Tytthaeninae Tytthaena Carnivoramorpha Carnivoraformes †Africtis †Ceruttia †Dawsonicyon †Dormaalocyon †Eogale †Gracilocyon †Harpalodon †Lycarion †Messelogale †Miacis †Miocyon †Neovulpavus †Oodectes †Palaearctonyx †Paramiacis †Paroodectes †Procynodictis †Prodaphaenus †Quercygale †Simamphicyon †Tapocyon †Uintacyon †Vassacyon †Vulpavus †Walshius †Xinyuictis †Zodiocyon Carnivora Caniformia (See Caniformia) Feliformia (See Feliformia) ichnotaxa of Carnivoraformes †Falcatipes †Viverravidae Bryanictis Didymictis Ictidopappus Intyrictis Orientictis Pappictidops Preonictis Pristinictis Protictis Raphictis Simpsonictis Variviverra Viverravus Viverriscus

v t e Mammals of clade Pholidotamorpha Kingdom: Animalia Phylum: Chordata Class: Mammalia Clade: Ferae Clade: Pholidotamorpha Pholidotamorpha Order: Pholidota Suborder: Eupholidota Superfamily: Manoidea Family: Manidae See in template: Manidae Family: †Patriomanidae †Cryptomanis †Patriomanis Incertae sedis †Necromanis Superfamily: †Eomanoidea Family: †Eomanidae †Eomanis Incertae sedis †Pholidota sp. (BC 16’08) Order: †Afredentata Family: †Eurotamanduidae †Eurotamandua Order: †Palaeanodonta Family: †Epoicotheriidae (paraphyletic family) †Alocodontulum †Amelotabes †Auroratherium †Dipassalus †Myrmecoboides †Tubulodon Subfamily: †Epoicotheriinae (paraphyletic subfamily) †Pentapassalus †Tetrapassalus †Epoicotherium/Xenocranium clade †Epoicotherium †Molaetherium †Xenocranium Family: †Ernanodontidae †Asiabradypus †Ernanodon Family: †Escavadodontidae †Escavadodon Family: †Metacheiromyidae (paraphyletic family) †Brachianodon †Mylanodon Subfamily: †Metacheiromyinae (paraphyletic subfamily) †Metacheiromys †Palaeanodon Subfamily: †Propalaeanodontinae †Propalaeanodon Incertae sedis †Arcticanodon †Melaniella other representatives †Euromanis

Taxon identifiers Ferae Wikidata: Q20868 Wikispecies: Ferae Paleobiology Database: 159623

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Adapted from the Wikipedia article [Ferae](https://en.wikipedia.org/wiki/Ferae) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Ferae?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.