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 such as creodonts[5] are members of Ferae as well.

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

Phylogeny of extant taxaEdit

Position of pangolinsEdit

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[6] and comparison of protein and DNA sequences[7][8][9] 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.[10]

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).[11][12] An alternate phylogeny (less supported) holds that the closest relatives to the Ferae are the Perissodactyla and Chiroptera (bats), not Cetartiodactyla.[13] 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 clades)

 (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 Carnivoramorpha (carnivorans and their stem-relatives).[14] 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]

Phylogenetic position of Creodonta within Ferae.[15][16]



Pholidota (sensu stricto)  


 (Pholidota sensu lato) 





Altacreodus/Tinerhodon clade

Hyaenodonta (sensu stricto)  

 sensu lato 

Viverravidae (sensu stricto)


carnivoramorph sp. (UALVP 50993 & UALVP 50994)



 sensu lato 

carnivoramorph sp. (UALVP 31176)


carnivoramorph sp. (WW-84: USNM 538395)


 (Carnivora sensu lato) 

Possible membersEdit

After Halliday et al. (2015) various enigmatic Palaeocene mammals have been found 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 supposedly "hoofed mammals" like the mesonychians and arctocyonids (usually considered as stem-artiodactyls[17]) 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.[5] This enlarged Ferae was also found to be the sister group to Chiroptera.[5]

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):[5]




































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