Ferae ( , , "wild beasts") is a mirorder of placental mammals in grandorder

Ferungulata Ferungulata (" wild beasts and ungulates") is a grandorder of placental mammals that groups together mirorder Ferae and clade Pan-Euungulata. It has existed in two guises, a traditional one based on morphological analysis and a revised one ta ...

, that groups together clades Pan-Carnivora (that includes

carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...

ns and their fossil relatives) and Pholidotamorpha (

pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smutsia''. ''Manis'' comprises four species found in Asia, while ' ...

s and their fossil relatives), and

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

order Pantolesta.

General characteristics

In mirorder Ferae

The common feature for members of this mirorder is ossified

tentorium cerebelli The cerebellar tentorium or tentorium cerebelli (Latin for "tent of the cerebellum") is one of four dural folds that separate the cranial cavity into four (incomplete) compartments. The cerebellar tentorium separates the cerebellum from the cere ...

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 ...

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 lithic flake, flake removed from a stone tool during the process of pressure flaking. In the Natufian cu ...

bones in the wrist.Floréal Solé & Thierry Smith (2013.
"Dispersals of placental carnivorous mammals (Carnivoramorpha, Oxyaenodonta & Hyaenodontida) near the Paleocene-Eocene boundary: a climatic and almost worldwide story"
Geologica Belgica 16/4: 254–261

In clade Pan-Carnivora

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

heterodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. Human dentition is heterodont and diphyodont as an example. In vertebrates, heterodont pertains to animals wher ...

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

from the other carnivorous placental mammals. However, these mammals are distinguished between themselves 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.

Classification and phylogeny

Sister groups to Ferae

According to recent studies, the closest relatives of Ferae are members of clade

Pan-Euungulata Pan-Euungulata (literally: "all true ungulates"; also known as: Euungulata ensu latoref name="Av"/>) is a clade of placental mammals from grandorder Ferungulata, consisting of the taxon Euungulata ensu strictoand all taxa (species) more clos ...

(group that includes mirorder

Euungulata Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with Hoof, hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined ...

and extinct genus ''

Protungulatum ''Protungulatum'' ('first ungulate') is an extinct genus of eutherian mammals within extinct family Protungulatidae, and is possibly one of the earliest known placental mammals in the fossil record, that lived in North America from the Late Cre ...

''). Together they form grandorder

. An alternate phylogeny holds that the closest relative to Ferae is order

Perissodactyla Perissodactyla (, ), or odd-toed ungulates, is an order of ungulates. The order includes about 17 living species divided into three families: Equidae (horses, asses, and zebras), Rhinocerotidae (rhinoceroses), and Tapiridae (tapirs). They t ...

, with whom they form a clade Zooamata. Together, clade Zooamata and order

Chiroptera Bats are flying mammals of the order Chiroptera (). 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 birds, flying with their very long spread-out ...

form clade Pegasoferae, and Pegasoferae is sister taxon to order

Artiodactyl Artiodactyls are placental mammals belonging to the order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof). The other t ...

a within clade Scrotifera. However, subsequent molecular studies have generally failed to support this proposal.

Position of pangolins and creodonts within clade

Pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smutsia''. ''Manis'' comprises four species found in Asia, while ' ...

s were long thought to be the closest relatives of

aardvark Aardvarks ( ; ''Orycteropus afer'') are medium-sized, burrowing, nocturnal mammals native to Africa. Aardvarks are the only living species of the family Orycteropodidae and the order Tubulidentata. They have a long proboscis, similar to a pi ...

and

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

ns, 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

ns, with whom they also share a few unusual derived morphological and anatomical traits, such as the ossified

and the fusion of the

and

bones in the wrist. The

last common ancestor A most recent common ancestor (MRCA), also known as a last common ancestor (LCA), is the most recent individual from which all organisms of a set are inferred to have descended. The most recent common ancestor of a higher taxon is generally assu ...

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 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 order Carnivora. The ''Halliday et al. (2015)'' phylogenetic analysis of hundreds of morphological characters of

Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...

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 A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...

taxon. Members of this group are now part of clade Pan-Carnivora and sister taxa to Carnivoramorpha (carnivorans and their stem-relatives), split in two groups: order Oxyaenodonta on one side and on the other side order

plus its stem-relatives, like family Wyolestidae (that only contains genus '' Wyolestes''), and genera '' Simidectes''S. P. Zack and S. Tomiya (2016.) "New postcrania clarify the affinities of the unusual Eocene mammal ''Simidectes''", i
"The Society of Vertebrate Paleontology 76th annual meeting"
/ref> and ''

Altacreodus ''Altacreodus'' (" creodont from Alberta") is an extinct genus of eutherian mammals. Fossils have been found in North America where they first appeared during the Late Cretaceous, and they died out prior to the start of the Paleocene. It is poss ...

''.Prevosti, F. J., & Forasiepi, A. M. (2018)
"Introduction. Evolution of South American Mammalian Predators During the Cenozoic: Paleobiogeographic and Paleoenvironmental Contingencies"
/ref>

Taxonomy

Alternative classification and possible fossil members

In ''Halliday et al. (2015)'' various enigmatic Palaeocene

eutheria Eutheria (from Greek , 'good, right' and , 'beast'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians ...

n mammals have been proposed to be possible members of Ferae, like members of orders

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

Pantodonta Pantodonta is an extinct suborder (or, according to some, an Order (biology), 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 K-T boundary, en ...

and

Taeniodonta Taeniodonta ("banded teeth") is an extinct order of eutherian mammals, that lived in North America and Europe from the late Cretaceous (Maastrichtian) to the middle Eocene. Taeniodonts evolved quickly into highly specialized digging animals ...

, and families Arctocyonidae, Didelphodontidae,

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

Palaeoryctidae Palaeoryctidae ("ancient diggers") is an extinct family of non-specialized eutherian mammals from extinct order Palaeoryctida, that lived in North America, Europe, Asia and Africa from the late Cretaceous to the middle Eocene.Gregg F. Gunnell, Th ...

, Periptychidae and Triisodontidae. Mesonychians are proposed to be a sister group to carnivoramorphs, while arctocyonids were

, with genera '' Arctocyon'' and '' Loxolophus'' as a sister taxa to pantodonts and periptychids, '' Goniacodon'' and '' Eoconodon'' sister to the Carnivoramorpha-Mesonychia clade, and other genera allied with creodonts and palaeoryctids. This enlarged Ferae was also found to be the sister group to order

, even though recent studies dispute 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