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Temporal range: 78.9–0 Ma[1]
Late Cretaceous - present
Order Carnivora.jpg
High diversity of carnivorans
Pangolin borneo.jpg
Low species diversity in Pholidotes: a pangolin
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Ferungulata
Clade: Ferae
Linnaeus, 1758[2]

Possible members:[3]

Ferae (/ˈfɪər/ FEER-ee, Latin[ˈfɛrae̯]) 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). An alternate name, Ostentoria, has also been proposed for a grouping of the Carnivora and Pholidota.[4] The last common ancestor of extant Ferae is supposed to have diversified ca. 78.9 million years ago.[1] Several extinct orders, relatives of the Pholidota, such as Creodonts,[3] are members of Ferae as well.

Phylogeny of extant taxa[edit]

Position of pangolins[edit]

Pangolins were long thought to be the closest relatives of Xenarthra (armadillos, anteaters, and sloths), contributing to the polyphyletic group of Edentata. Research based on immunodiffusion technique[5] and comparison of protein and DNA sequences[6][7][8] revealed the close relationships between pangolins and carnivorans. Living pangolins and carnivorans also share 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.[9]

Sister groups to Ferae[edit]

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).[10][11] An alternate phylogeny (less supported) holds that the closest relatives to the Ferae are the Perissodactyla and Chiroptera (bats), not Cetartiodactyla.[12] 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.

 (primates, colugos, treeshrews, rodents, rabbits)  Lepus timidus - 1700-1880 - Print - Iconographia Zoologica -(white background).jpg


 (hedgehogs, shrews, moles, solenodons)  Erinaceus europaeus - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg


 (bats and flying foxes)  Flying fox at botanical gardens in Sydney (cropped and flipped).jpg


 (pangolins)  Pangolin Hardwicke (white background).jpg

 (cats, hyenas, dogs, bears, seals, etc.)  Dogs, jackals, wolves, and foxes (Plate XI).jpg


  (horses, tapirs, rhinos, etc.) Equus quagga (white background).jpg

 (camels, pigs, ruminants, hippos, whales, etc.)  The deer of all lands (1898) Hangul white background.png

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

Fossil members[edit]

Position of Creodonta[edit]

Various enigmatic Palaeocene mammals have been found to belong to Ferae. While there has been strong support in the inclusion of creodonts into Ferae, they were usually recovered as sister taxon to Carnivoramorpha (carnivorans and their stem-relatives).[13] The 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).[3]

Phylogenetic hypothesis for the position of Creodonta within Ferae.

Euungulata Equus quagga (white background).jpg


Pholidota (sensu stricto) Pangolin Hardwicke (white background).jpg

Palaeanodonta Metacheiromys DB152-2.jpg

 (Pholidota sensu lato) 

Oxyaenidae Patriofelis ferox by R. B. Horsfall (coloured).png




Hyaenodonta (sensu stricto) Hyaenodon horridus by R. B. Horsfall (coloured).jpg

 sensu lato 




Carnivoraformes Dogs, jackals, wolves, and foxes (Plate XI).jpg

 (Carnivora sensu lato) 

Possible members[edit]

In addition various supposedly "hoofed mammals" like the mesonychians and arctocyonids (usually considered as stem-artiodactyls[16]) also belong to the group. In addition, Mesonychians might be 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, and Protungulatum not a placental mammal.[3] This enlarged Ferae was also found to be the sister group to Chiroptera.[3]

Below is a phylogeny of the interrelationships within Ferae among the various extant and fossil groups, as well as their sister placement to Chiroptera after Halliday et al. (2015)[3]:




































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