Ferae

Ferae (, , "wild beasts") is a mirorder of placental mammals from grandorder Ferungulata, that groups together clades Pan-Carnivora, which includes modern carnivorans, and Pholidotamorpha, which includes pangolins.

In mirorder Ferae
The common feature for members of this mirorder is ossified tentorium cerebelli and the fusion of the scaphoid and lunate bones in the wrist.

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

Carnassials are feature that allows distinguishing the Carnivoramorpha, Oxyaenodonta and Hyaenodonta from the other carnivorous placental mammals. However, these mammals are distinguished between them self 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.

Sister groups to Ferae and position of pangolins and creodonts
According to recent studies (reflected in the diagram below), the closest living relatives of Ferae are members of mirorder Euungulata (group of mammals which includes order Perissodactyla and Artiodactyla).

An alternate phylogeny holds that the closest relatives to the Ferae are the Perissodactyla and Chiroptera (bats), not Artiodactyla. Ferae together with Perissodactyla has been called Zooamata. Ferae, Perissodactyla, and Chiroptera together has been called Pegasoferae. Subsequent molecular studies have generally failed to support the proposal.

Pangolins were long thought to be the closest relatives of aardvark and xenarthrans (armadillos, anteaters, and sloths), forming to the now obsolete order Edentata. Research based on immunodiffusion technique and comparison of protein and DNA sequences  revealed the close relationships between pangolins and carnivorans, with whom they 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. The last common ancestor of extant Ferae is supposed to have diversified c. 79.47 million years ago.

While there has been strong support in the inclusion of order Creodonta into Ferae, they were usually recovered as sister taxon to Carnivora. 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). However, recent studies have shown that Creodonta is an invalid polyphyletic taxon. Members of this group are part of clade Pan-Carnivora and sister taxa to Carnivoramorpha (carnivorans and their stem-relatives). They are split in two groups: order Oxyaenodonta on one side and order Hyaenodonta plus its stem-relatives (family Wyolestidae and genera Altacreodus, Simidectes and Tinerhodon) on the other.

Taxonomy


! Traditional classification: !! Revised classification:
 * - style="vertical-align:top;"
 * Clade: Ferae (Linnaeus, 1758)
 * Order: Carnivora (Bowdich, 1821) (carnivorans)
 * Order: Cimolesta (McKenna, 1975) [a group with pangolins as its members at that time]
 * Order: †Creodonta (Cope, 1875) (false carnivorans)
 * Mirorder: Ferae (Linnaeus, 1758)
 * Clade: Pan-Carnivora (Flynn, Wyss & Wolsan, 2020)
 * Clade: Carnivoramorpha (Wyss & Flynn, 1993) (carnivorans and stem-relatives)
 * Order: †Hyaenodonta (Van Valen, 1967)
 * Order: †Oxyaenodonta (Van Valen, 1971)
 * Family: †Oxyaenidae (Cope, 1877)
 * Family: †Wyolestidae (Gingerich, 1981)
 * Genus: †Simidectes (Stock, 1933)
 * (unranked): †Altacreodus/Tinerhodon clade
 * Genus: †Altacreodus (Fox, 2015)
 * Genus: †Tinerhodon (Gheerbrant, 1995)
 * Clade: Pholidotamorpha (Gaudin, 2009)
 * Order: Pholidota (Weber, 1904) (pangolins)
 * Order: †Palaeanodonta (Matthew, 1918) (stem-pangolins)
 * }
 * Order: †Palaeanodonta (Matthew, 1918) (stem-pangolins)
 * }

Alternative classification and possible fossil members
In Halliday et al. (2015) various enigmatic Palaeocene mammals have been proposed to be possible members to Ferae, like members of orders Pantodonta and Taeniodonta, and families Didelphodontidae, Nyctitheriidae, Oxyclaenidae, Palaeoryctidae, Pantolestidae, Pentacodontidae, Periptychidae and Triisodontidae. In addition various "hoofed mammals" like the mesonychians and arctocyonids (usually considered as a stem-artiodactyls ) also placed in this group. 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. This enlarged Ferae was also found to be the sister group to Chiroptera, even though recent studies dispute this classification.