Ferungulata ("
wild beasts and
ungulates
Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined to b ...
") is a
grandorder of
placental mammals that groups together
mirorder Ferae and 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 ...
. 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 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 ...
(bats) and together they make clade
Scrotifera.
General characteristics
According to a 2022 study by Anne E. Kort, members of grandorder Ferungulata, in addition to their genetic similarities, share common
synapomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
in the
lumbar vertebrae
The lumbar vertebrae are located between the thoracic vertebrae and pelvis. They form the lower part of the back in humans, and the tail end of the back in quadrupeds. In humans, there are five lumbar vertebrae. The term is used to describe t ...
(the S-shaped
postzygapophysis). In extant ferungulate mammals, this is known only within members of order
Artiodactyla
Artiodactyls are placental mammals belonging to the order (biology), 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 ...
. However, this feature is found also in many extinct members of Ferungulata, including
hyaenodonts,
oxyaenids,
mesonychids,
arctocyonids and the stem-relatives of perissodactyls (like ''
Cambaytherium'').
[Anne E. Kort (2022.) "Bizarre Backbones: A Synapomorphy in the Lumbar Vertebrae for Ferungulata", in ]
"The Society of Vertebrate Paleontology 82nd annual meeting"
' This synapomorphy suggests that this may be basal to all Ferungulata and secondarily lost in modern members, like
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 crown
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 ...
. 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 several other 2022 studies the extinct
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 ...
ns
pantodonts
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 ...
,
tillodonts and genus ''
Deltatherium'' (family Deltatheriidae) are recognised as crown-group placental mammals,
[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"
' who also possess S-shaped postzygapophysis like other members of Ferungulata.
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, contributing '' Tempo ...
in 1945.
[ It grouped together the ]extant
Extant or Least-concern species, least concern 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
* Exta ...
orders 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 ...
, 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 ...
and Artiodactyla
Artiodactyls are placental mammals belonging to the order (biology), 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 ...
with the Tubulidentata and the superorder
Order () 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 recognized ...
Paenungulata, 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 ...
.
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.[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 These studies separated his ungulate orders into two distinct placental groups, within Afrotheria
Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a superorder of placental mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephan ...
and Laurasiatheria
Laurasiatheria (; "Laurasian beasts") is a superorder of Placentalia, placental mammals that groups together true insectivores (eulipotyphlans), bats (chiropterans), carnivorans, pangolins (Pholidota, pholidotes), even-toed ungulates (Artiodacty ...
, 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 (). The one Neontology, extant family, the Manidae, has three genera: ''Manis'', ''Phataginus'', and ''Smutsia''. ''Manis'' comprises four species found in Asia, ...
), but the Tubulidentata and paenungulates are excluded. Although Simpson placed whales (Cetacea
Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
) 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.
The karyotype
A karyotype is the general appearance of the complete set of chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is discerned by de ...
s of ferungulates are highly conserved: "the fereungulatan ancestor chromosomes (n = 23 + X) ... differ from those of the antecedent scrotiferan ancestor by five rearrangements that occurred over 1 My (''citation excluded''). For the fereungulatan ancestor, we discovered four ancestral syntenies..."
Point coloration has been observed in a wide variety of ferungulates, including cats, foxes, horses, and dogs.
Taxonomy
See also
* Mammal classification
References
{{Taxonbar, from=Q5445608, from2=Q5444079
Mammal taxonomy