Ferae (/ˈfɪər/ FEER-ee, Latin: [ˈfɛrae̯], "wild beasts") is a clade of mammals, consisting of the orders Carnivora (wolves, cats, seals, and so on; over 260 species around the globe) and Pholidota (eight species of pangolins in tropical Africa and Asia). Another name, Ostentoria, has also been proposed for a grouping of the Carnivora and Pholidota.[3] The last common ancestor of extant Ferae is supposed to have diversified c. 78.9 million years ago.[4] Several extinct orders such as creodonts[5] are members of Ferae as well.

Temporal range: 78.9–0 Ma early Late Cretaceous to present
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Grandorder: Ferungulata
Mirorder: Ferae
Linnaeus, 1758[1]
Living subgroups

Others, see text

list of synonyms:
  • Carnaria (Haeckel, 1866)
  • Carnassia (Haeckel, 1895)
  • Carnivora (Zagorodniuk, 2008)[2]
  • Carnivoramorpha (Kalandadze & Rautian, 1992)
  • Ferina (Newman, 1843)
  • Ostentoria (Amrine-Madsen, 2003)[3]
  • Rapacia (Newman, 1843)
  • Sarcotheria (Haeckel, 1895)

Classification and phylogenyEdit

Traditional classificationEdit

Revised classificationEdit

Phylogeny of extant taxaEdit

Position of pangolinsEdit

Pangolins were long thought to be the closest relatives of Xenarthra (armadillos, anteaters, and sloths), forming to the polyphyletic group Edentata. Research based on immunodiffusion technique[11] and comparison of protein and DNA sequences[12][13][14] revealed the close relationships between pangolins and carnivorans. Living pangolins and carnivorans also share a few unusual derived morphological and anatomical traits, such as the ossified tentorium cerebelli and the fusion of the scaphoid and lunate bones in the wrist.[15]

Sister groups to FeraeEdit

According to recent studies (reflected in the diagram below), the closest relatives of Ferae are Perissodactyla (horses, tapirs, and rhinos) and Cetartiodactyla (which combines Artiodactyla—camels, pigs, ruminants and hippos—with Cetacea—whales and dolphins).[16][17] An alternate phylogeny (less supported) holds that the closest relatives to the Ferae are the Perissodactyla and Chiroptera (bats), not Cetartiodactyla.[18] Ferae together with Perissodactyla has been called Zooamata. Ferae, Perissodactyla, and Chiroptera together has been called Pegasoferae.

Phylogenetic position of the Ferae in the context of the order-level cladogram of Boreoeutheria
(only living groups)

 (primates, colugos, treeshrews, rodents, rabbits)   


 (hedgehogs, shrews, moles, solenodons)   


 (bats and flying foxes)   



 (cats, hyenas, dogs, bears, seals, etc.)   


  (horses, tapirs, rhinos, etc.)  

 (camels, pigs, ruminants, hippos, whales, etc.)   

The cladogram has been reconstructed from mitochondrial and nuclear DNA and protein characters.

Fossil membersEdit

Position of CreodontaEdit

While there has been strong support in the inclusion of creodonts into Ferae, they were usually recovered as sister taxon to Carnivora.[6] The Halliday et al. (2015) phylogenetic analysis of hundreds of morphological characters of Paleocene placentals found instead that creodonts might be the sister group to Pholidotamorpha (pangolins and their stem-relatives).[5] However, recent studies have show Creodonta is polyphyletic clade and sister taxon to Carnivoramorpha (carnivorans and their stem-relatives), split in two groups: order Oxyaenodonta on one side and order Hyaenodonta plus stem-relatives (Altacreodus and Tinerhodon) on the other.[19][8][9][10][20]

Phylogenetic position of "Creodonta" within Ferae.[19][8][9][10][20]



Pholidota (sensu stricto)  


 (Pholidota sensu lato) 





Altacreodus/Tinerhodon clade

Hyaenodonta (sensu stricto)  

 sensu lato 

Carnivoramorpha (Carnivora sensu lato)  

Alternative classification and possible membersEdit

In Halliday et al. (2015) various enigmatic Palaeocene mammals have been proposed to be possible members to Ferae, like members of suboders Pantodonta and Taeniodonta, and families Didelphodontidae, Nyctitheriidae, Oxyclaenidae, Palaeoryctidae, Pantolestidae, Pentacodontidae, Periptychidae, Triisodontidae and Wyolestidae.[5] In addition various "hoofed mammals" like the mesonychians and arctocyonids (usually considered as a stem-artiodactyls[21]) also placed in this group. In addition, Mesonychians are placed as the sister group to carnivoramorphs, while arctocyonids are polyphyletic with Arctocyon and Loxolophus sister to pantodonts and periptychids, Goniacodon and Eoconodon sister to the Carnivoramorpha-Mesonychia clade, most other genera allied with creodonts and palaeoryctidans.[5] This enlarged Ferae was also found to be the sister group to Chiroptera,[5] even though recent studies despute this classification.[22][23]

Below is a phylogeny of the interrelationships within Ferae among the various extant and fossil groups after Halliday et al. (2015):[5]





































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