Ferungulata ("

wild beasts Wild Beasts were an English indie rock band, formed in 2002 in Kendal. They released their first single, "Brave Bulging Buoyant Clairvoyants", on Bad Sneakers Records in November 2006, and subsequently signed to Domino Records. They have rele ...

and

ungulates 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, cam ...

") is a

grandorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...

placental mammals Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...

that groups together

mirorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...

Ferae Ferae ( , , "wild beasts") is a mirorder 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-Ca ...

and clade Pan-Euungulata. It has existed in two guises, a traditional one based on morphological analysis and a revised one taking into account more recent molecular analyses. The Fereungulata is a sister group to the order

Chiroptera Bats are mammals of the Order (biology), 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 fli ...

(bats) and together they make clade

Scrotifera Scrotifera ("beasts with scrotum") is a clade of Placentalia, placental mammals that groups together grandorder Ferungulata, clade Bat, Apo-Chiroptera (aka Order (biology), order Bat, Chiroptera), other extinct members and their common ancestors ...

Characteristics

According to 2022 study of Anne E. Kort, member of grandorder Ferungulata, in addition to their genetic similarities, share common

synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...

in the

lumbar vertebrae The lumbar vertebrae are, in human anatomy, the five vertebrae between the rib cage and the pelvis. They are the largest segments of the vertebral column and are characterized by the absence of the foramen transversarium within the transverse p ...

(the S-shaped

postzygapophysis The articular processes or zygapophyses (Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "process") of a vertebra are projections of the vertebra that serve the purpose of fitting with an adjacent vertebr ...

). In extant ferungulate mammals, this is known only within members of order

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 ...

. However, this feature is found also in many extinct members of Ferungulata, including

hyaenodonts Hyaenodontidae ("hyena teeth") is a family of extinct predatory mammals from extinct superfamily Hyaenodontoidea within extinct order Hyaenodonta. Hyaenodontids arose during the early Eocene and persisted well into the early Miocene. Fossils of t ...

, oxyaenids,

mesonychids 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 ...

, arctocyonids and the stem-relatives of perissodactyls.Anne E. Kort (2022.) "Bizarre Backbones: A Synapomorphy in the Lumbar Vertebrae for Ferungulata", in
"The Society of Vertebrate Paleontology 82th annual meeting"
' This synapomorphy suggests that this may be basal to all Ferungulata and secondarily lost in modern members, like

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 ...

ns and crown Perissodactyla. Previous study has shown that these S-shaped zygapophyses prevent torsion between vertebrae. As said in this study, it is possible this feature evolved in response to a need for stabilization in posterior spine as ribs became reduced. Also, in this and two other 2022 studies the extinct

eutheria Eutheria (; from Greek , 'good, right' and , 'beast'; ) is the clade consisting of all therian mammals that are more closely related to placentals than to marsupials. Eutherians are distinguished from noneutherians by various phenotypic tra ...

pantodonts 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 las ...

, tillodonts and genus '' Deltatherium'' are recognised as crown-group placental mammals and members of Ferungulata, who also possess S-shaped postzygapophysis.Thomas E. Williamson, Sarah L. Shelley, Gregory Funston, John R. Wible, Stephen L. Brusatte (2022.) "Triangular beast: New fossils shed light on ''Deltatherium'', an enigmatic early Paleocene mammal from New Mexico", in
"The Society of Vertebrate Paleontology 82nd annual meeting"
'

Classification and phylogeny

History of phylogeny

The traditional Ferungulata was established by

George Gaylord Simpson George Gaylord Simpson (June 16, 1902 – October 6, 1984) was an American paleontologist. Simpson was perhaps the most influential paleontologist of the twentieth century, and a major participant in the Modern synthesis (20th century), modern ...

in 1945. It grouped together the

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

orders

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 ...

, Perissodactyla and

with the

Tubulidentata Orycteropodidae is a family of afrotherian mammals. Although there are many fossil species, the only species surviving today is the aardvark, ''Orycteropus afer''. Orycteropodidae is recognized as the only family within the order Tubulidentata ...

and the

superorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...

Paenungulata Paenungulata (from Latin ''paene'' "almost" + ''ungulātus'' "having Hoof, hoofs") is a clade of "sub-Ungulate, ungulates", which groups three Extant taxon, extant mammal Order (biology), orders: Proboscidea (including elephants), Sirenia (sirenia ...

, as well as a number of orders known only from

fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...

. Simpson established the grouping on the basis of morphological criteria, but this traditional understanding of Ferungulata has been challenged by a more recent classification, relying upon genetic criteria. These studies separated his ungulate orders into two distinct placental groups, within

Afrotheria Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a clade of mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephant shrews (also know ...

and

Laurasiatheria Laurasiatheria ("laurasian beasts") is a superorder of placental mammals that groups together true insectivores ( eulipotyphlans), bats ( chiropterans), carnivorans, pangolins ( pholidotes), even-toed ungulates (artiodactyls), odd-toed ungulates ...

, respectively. The 'true' ungulates (mirorder Euungulata), Perissodactyla and Artiodactyla, are included in the revised group, along with the Carnivora, and with the addition of pangolins (order

Pholidota 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 ''Smutsia ...

), but the Tubulidentata and paenungulates are excluded. Although Simpson placed whales (

Cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...

) in a separate cohort, recent evidence linking them to Artiodactyla would mean that they belong here as well. To reflect this difference, the revised clade is usually referred to as Fereungulata.

Taxonomy

Phylogeny

The phylogenetic relationships of grandorder Ferungulata 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: .Frank Zachos (2020.
"Mammalian Phylogenetics: A Short Overview of Recent Advances"
/ref>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

Mammal taxonomy {{Mammal-stub