Ferae ( , , "wild beasts") is a

mirorder Order ( la, wikt:ordo#Latin, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomi ...

of placental mammalsMalcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'' in Columbia University Press, New York (1997), 631 Seiten. that groups together clades Pan-Carnivora and

Pholidotamorpha Pholidotamorpha ("pangolin-shaped") is a clade of mammals that includes the orders Palaeanodonta and Pholidota (the pangolins). In the past both orders were formerly classified with various other orders of ant-eating mammals, most notably Xenar ...

. The Ferae is a sister group to the clade Pan-Euungulata and together they make grandorder Ferungulata.

Classification and phylogeny

Taxonomy

Phylogeny

The phylogenetic relationships of mirorder Ferae are shown in the following cladogram, reconstructed from mitochondrial and nuclear DNA and protein characters, as well as the fossil record.O’Leary, M. A., Bloch JI, Flynn, J. J., Gaudin, T. J., Giallombardo, A., Giannini, N. P., Goldber, S. L, Kraatz, B. P., Luo, Z-X, Jin Meng, Xijun Ni, Novacek, M. J., Perini, F. A., Randall, Z. S., Rougier, G. W., Sargis, E. J., Silcox, M. T., Simmons, N. B., Spaulding, M. Velazco, P. M., Weksler, M., Wible, J. R. Cirranello, A. L. (2013.
"The Placental Mammal Ancestor and the Post–K-Pg Radiation of Placentals."
Science 339:6120:662-667.Conference abstract (p. 99)
. Explanation and conclusions: .Prevosti, F. J., & Forasiepi, A. M. (2018)
"Introduction. Evolution of South American Mammalian Predators During the Cenozoic: Paleobiogeographic and Paleoenvironmental Contingencies"
/ref>

Position of pangolins

Pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', ''Phataginus'', and '' Smuts ...

s were long thought to be the closest relatives of

Xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. ...

(armadillos, anteaters, and sloths), forming to the

polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...

group Edentata. Research based on immunodiffusion technique and comparison of protein and DNA sequences revealed the close relationships between pangolins and carnivorans. Living

pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', ''Phataginus'', and '' Smuts ...

s and carnivorans also share a few unusual derived morphological and anatomical traits, such as the ossified

tentorium cerebelli The cerebellar tentorium or tentorium cerebelli ( Latin for "tent of the cerebellum") is an extension of the dura mater that separates the cerebellum from the inferior portion of the occipital lobes. Structure The cerebellar tentorium is an arc ...

and the fusion of the

scaphoid The scaphoid bone is one of the carpal bones of the wrist. It is situated between the hand and forearm on the thumb side of the wrist (also called the lateral or radial side). It forms the radial border of the carpal tunnel. The scaphoid bone i ...

and

lunate Lunate is a crescent or moon-shaped microlith. In the specialized terminology of lithic reduction, a lunate flake is a small, crescent-shaped flake removed from a stone tool during the process of pressure flaking. In the Natufian period, a lu ...

bones in the wrist. The

last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...

of extant Ferae is supposed to have diversified c. 78.9 million years ago.

Sister groups to Ferae

According to recent studies (reflected in the diagram below), the closest living relatives of Ferae are members of mirorder

Euungulata Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ca ...

(group of mammals which includes order Perissodactyla and

Artiodactyla The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...

). An alternate phylogeny holds that the closest relatives to the Ferae are the Perissodactyla and

Chiroptera Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bir ...

(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.

Fossil members

Position of creodonts

While there has been strong support in the inclusion of order

Creodonta Creodonta ("meat teeth") is a former order of extinct carnivorous placental mammals that lived from the early Paleocene to the late Miocene epochs in North America, Europe, Asia and Africa. Originally thought to be a single group of animals ance ...

into Ferae, they were usually recovered as sister taxon to

Carnivora Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...

. Diagnostic traits shared by creodonts and carnivorans include the presence of carnassial teeth. 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

(pangolins and their stem-relatives). However, recent studies have show Creodonta is invalid

taxon. Members of this group are part of clade Pan-Carnivora and sister taxa to

Carnivoramorpha Carnivoramorpha ("carnivoran-like forms") is a clade of placental mammals that includes the modern order Carnivora and its extinct stem-relatives.Bryant, H.N., and M. Wolson (2004“Phylogenetic Nomenclature of Carnivoran Mammals.”''First Inte ...

(carnivorans and their stem-relatives). They are split in two groups: order Oxyaenodonta on one side and order

Hyaenodonta Hyaenodonta (" hyena teeth") is an extinct order of hypercarnivorous placental pan-carnivoran mammals from mirorder Ferae. Hyaenodonts were important mammalian predators that arose during the early Paleocene in Europe and persisted well into ...

plus its stem-relatives ('' Altacreodus'' and '' Tinerhodon'') on the other.

Alternative classification and possible members

In ''Halliday et al. (2015)'' various enigmatic Palaeocene mammals have been proposed to be possible members to Ferae, like members of suboders

Pantodonta Pantodonta is an extinct suborder (or, according to some, an order) of eutherian mammals. These herbivorous mammals were one of the first groups of large mammals to evolve (around 66 million years ago) after the end of the Cretaceous. The last ...

and Taeniodonta, and families Didelphodontidae,

Nyctitheriidae Nyctitheriidae is a family of extinct eulipotyphlan insectivores known from the Paleocene and Eocene epochs of North America and Asia and persisting into the Oligocene of Europe. Several genera, including '' Nyctitherium'', '' Paradoxonycteris' ...

, Oxyclaenidae,

Palaeoryctidae Palaeoryctidae or Palaeoryctoidea ("old/stony digger", from Greek: ὀρύκτης, ''oryctes'') is an extinct group of relatively non-specialized non-placental eutherian mammals that lived in North America during the late Cretaceous and took par ...

Pantolestidae Pantolestidae is an extinct family of semi-aquatic, non-placental eutherian mammals. Forming the core of the equally extinct suborder Pantolesta, the pantolestids evolved as a series of increasingly otter-like forms, ranging from the Middle P ...

, Pentacodontidae,

Periptychidae Periptychidae is a family of Paleocene placental mammals, known definitively only from North America. The family is part of a radiation of early herbivorous and omnivorous mammals formerly classified in the extinct order "Condylarthra", which ma ...

and Triisodontidae. In addition various "hoofed mammals" like the

mesonychia Mesonychia ("middle claws") is an extinct taxon of small- to large-sized carnivorous ungulates related to artiodactyls. Mesonychids first appeared in the early Paleocene, went into a sharp decline at the end of the Eocene, and died out entirely ...

ns and arctocyonids (usually considered as a stem-

artiodactyl The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...

s) also placed in this group. Mesonychians are placed as the sister group to carnivoramorphs, while arctocyonids are polyphyletic with ''

Arctocyon ''Arctocyon'' ('bear dog') is an extinct genus of ungulate mammals. ''Arctocyon'' was a "ground dwelling omnivore", that lived from 61.3-56.8 Ma. Synonyms of ''Arctocyon'' include ''Claenodon'', and ''Neoclaenodon''. ''Arctocyon'' was likely plant ...

'' and '' Loxolophus'' sister to

pantodont Pantodonta is an extinct suborder (or, according to some, an order) of eutherian mammals. These herbivorous mammals were one of the first groups of large mammals to evolve (around 66 million years ago) after the end of the Cretaceous. The last ...

s and periptychids, '' Goniacodon'' and ''

Eoconodon ''Eoconodon'' is an extinct genus of triisodontid mesonychid that existed during the early Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of t ...

'' 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

, even though recent studies despute this classification.Frank Zachos (2020.
"Mammalian Phylogenetics: A Short Overview of Recent Advances"
In book: "Mammals of Europe - Past, Present, and Future" (pp.31-48)Xue Lv, Jingyang Hu, Yiwen Hu, Yitian Li, Dongming Xu, Oliver A. Ryder, David M. Irwin, Li Yu (2021.
"Diverse phylogenomic datasets uncover a concordant scenario of laurasiatherian interordinal relationships"
Molecular Phylogenetics and Evolution, Volume 157

References

{{Taxonbar, from=Q20868 Mammal unranked clades