Ferae

Ferae [/ˈfɪəriː/ 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). 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.

Ferae Temporal range: 78.9–0 Ma[1] PreꞒ Ꞓ O S D C P T J K Pg N Late Cretaceous to present
High diversity of carnivorans
Low species diversity in Pholidotes
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Ferungulata
Clade:	Ferae Linnaeus, 1758[2]
Subgroups
[see classification]
Synonyms
list of synonyms: Carnaria (Haeckel, 1866) Carnassia (Haeckel, 1895) Carnivora (Zagorodniuk, 2008)[3] Carnivoramorpha (Kalandadze & Rautian, 1992) Ferina (Newman, 1843) Ostentoria (Amrine-Madsen, 2003)[4] Rapacia (Newman, 1843) Sarcotheria (Haeckel, 1895)

Classification and phylogeny

Traditional classification Clade: Ferae [6][7] Order: Carnivora (carnivorans) Order: Cimolesta [including pangolins] Order: †Creodonta

Revised classification Clade: Ferae [4][8][9][10] Clade: Pan-Carnivora Clade: Carnivoramorpha Order: †Hyaenodonta Order: †Oxyaenodonta (unranked): †Altacreodus/Tinerhodon clade Genus: †Altacreodus Genus: †Tinerhodon Clade: Pholidotamorpha Order: Pholidota (pangolins) Order: †Palaeanodonta (stem-pangolins)

Position of pangolins

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[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 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 Ferae

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)

Boreoeutheria     Euarchontoglires  (primates, colugos, treeshrews, rodents, rabbits)     Laurasiatheria     Eulipotyphla  (hedgehogs, shrews, moles, solenodons)     Scrotifera     Chiroptera  (bats and flying foxes)     Ferungulata   Ferae     Pholidota  (pangolins)       Carnivora  (cats, hyenas, dogs, bears, seals, etc.)       Euungulata     Perissodactyla   (horses, tapirs, rhinos, etc.)      Cetartiodactyla  (camels, pigs, ruminants, hippos, whales, etc.)

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

Position of Creodonta

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).[19] 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, split in two groups: order Oxyaenodonta on one side and order Hyaenodonta plus stem-relatives (Altacreodus and Tinerhodon) on the other.[20][8][9][10][21]

Phylogenetic position of "Creodonta" within Ferae.[20][8][9][10][21]

Ferungulata    Pan-Euungulata    Ferae   Pholidotamorpha    Pholidota (sensu stricto)     †Palaeanodonta    (Pholidota sensu lato)   Pan-Carnivora  †"Creodonta"    †Oxyaenodonta  †Oxyaenidae        †Hyaenodonta      †Altacreodus     †Tinerhodon   †Altacreodus/Tinerhodon clade     †Hyaenodonta (sensu stricto)    sensu lato    Carnivoramorpha (Carnivora sensu lato)

Possible members

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[22]) 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]

...

Chiroptera    Ferae        Carnivoramorpha     †Mesonychia      †Triisodontidae    †Goniacodon     †Eoconodon                 †Pentacodontidae     †Periptychidae       †Pantodonta      †Arctocyonidae    †Arctocyon     †Loxolophus                 †Taeniodonta     †Escavadodontidae         †Wyolestidae     †Nyctitheriidae           †Pantolestidae    Pholidotamorpha    †Palaeanodonta     Pholidota            †Oxyclaenidae        †Chriacus     †Thryptacodon         †Claenodon     †Anacodon         †Oxyclaenus         †Palaeoryctidae     †Didelphodus     †Acmeodon     †Gelastops    †Creodonta    †Oxyaenidae     †Hyaenodonta

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