Altungulata or Pantomesaxonia (''

sensu ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

'' and later authors) is an invalid

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

(

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 ...

) of

ungulate 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 ...

mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...

s comprising the

perissodactyls Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...

hyracoids Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simila ...

, and tethytheres (

sirenians The Sirenia (), commonly referred to as sea-cows or sirenians, are an order (biology), order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently compris ...

proboscideans The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. From ...

, and related extinct taxa). The name "Pantomesaxonia" was originally introduced by , a German zoologist and racial theorist. It was resurrected by by including sirenians and excluding South American ungulates, phenacodontids, and meniscotheriids from the original concept. The name "Altungulata", introduced by and revised by , was erected as an alternative because the updated concept of "Pantomesaxonia" was regarded too deviant from the original concept. Both names are still in use, and, to add to the confusion, various authors assign different ranks to the involved taxa. For example, according to ,

Phenacodonta Phenacodontidae is an extinct family of large herbivorous mammals traditionally placed in the “wastebasket taxon” Condylarthra, which may instead represent early-stage perissodactyls. They lived in the Paleocene and Eocene epochs (about 60 ...

(Phenacodontidae and Meniscotheriidae) and Pantomesaxonia (Sirenia, Desmostylia, Proboscidea, Hyracoidea, and Perissodactyla) are sister groups together making up the superorder Paenungulata. Altungulata is not supported by molecular evidence unless perissodactyls are excluded (thus dividing Altungulata into

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 ...

and

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 the validity of the following uniting

synapomorphies 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 ...

remain disputed: * bilophodonty, two lophs or crests running transversally across the crown of the tooth * large third molars * molarization of posterior premolars * elongated thoracic region with at least 19 vertebrae * clavicle absent * similar development of fetal membranes Recent studies on '' Abdounodus'' showcase that dental synapomorphies between both groups arose independently, further discrediting the Altungulata hypothesis.

Classification

The classification below is from .

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 ...

together with Macroscelidea,

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 lipotyphlan families Tenrecidae and

Chrysochloridae Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family Chrysochloridae and as such they are taxonomically distinct from the true moles, family Talpidae, and other mole-like families, all o ...

compose

. With the exclusion of the better known ''Radinskya'' and ''Minchenella'' from Phenacolophidae, their affinities to Embrithropoda are suspect, and they were regarded as Altungulata ''incertae sedis'' by Mao et al. (2015).Fang-Yuan Mao, Yuan-Qing Wang, Qian Li & Xun Jin (2015): New records of archaic ungulates from the Lower Eocene of Sanshui Basin, Guangdong, China, Historical Biology: An International Journal of Paleobiology, DOI: 10.1080/08912963.2015.1034120 Altungulata Prothero and Schoch 1989 * †'' Radinskya'', * †''

Olbitherium ''Olbitherium'' is an extinct mammal of the order Perissodactyla (an early ancestor of modern-day horses, rhinos, and tapirs) from the early Eocene epoch of Shandong, China. Description and classification ''Olbitherium'' was originally described ...

'' * † Phenacolophidae Zhang 1978 ** †'' Ganolophus'' Zhang 1979 ** †'' Phenacolophus'' Matthew and Granger 1925 (= ''Procoryphodon'' Flerow 1957) ** †'' Sanshuilophus'' Mao et al. 2015 ** †'' Tieshanilophus'' Tong 1979 ** †'' Yuelophus'' Zhang 1978 ** †'' Zaisanolophus'' Gabunia 1998 * Order Perissodactyla ** Suborder

Hippomorpha Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...

*** Superfamily Equoidea ****

Equidae Equidae (sometimes known as the horse family) is the taxonomic family of horses and related animals, including the extant horses, asses, and zebras, and many other species known only from fossils. All extant species are in the genus '' Equus'', ...

**** †

Palaeotheriidae Palaeotheriidae is an extinct family of herbivorous perissodactyl mammals related to equids. They ranged across Europe and Asia from the Eocene through to the early Oligocene 55–33 Ma, existing for approximately . Living in dense forests ...

** Suborder

Tapiromorpha Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) ...

*** †

Isectolophidae Isectolophidae is a potentially paraphyletic family of browsing, herbivorous, mammals in the Perissodactyla suborder Ancylopoda that show long, curved and cleft claws A claw is a curved, pointed appendage found at the end of a toe or fin ...

*** Infraorder

Ceratomorpha Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) ...

**** Superfamily

Tapiroidea Tapiroidea is a superfamily of perissodactyls which includes the modern tapir. Members of the superfamily are small to large browsing mammals, roughly pig-like in shape, with short, prehensile snouts. Their closest relatives are the other odd-toe ...

***** †

Helaletidae The Helaletidae is an extinct family of tapiroid closely related and likely ancestral to the true tapirs, which contain ''Protapirus ''Protapirus'' (Latin: "before" (pro), + Brazilian Indian: "tapir" (tapira)) is an extinct genus of tapir know ...

***** †

Deperetellidae Deperetellidae is an extinct family of herbivorous odd-toed ungulates containing the genera '' Bahinolophus'', '' Deperetella'', '' Irenolophus'', and ''Teleolophus''. Their closest living relatives are tapirs Tapirs ( ) are large, herbivoro ...

***** † Lophialetidae *****

Tapiridae Tapirs ( ) are large, herbivorous mammals belonging to the family Tapiridae. They are similar in shape to a pig, with a short, prehensile nose trunk. Tapirs inhabit jungle and forest regions of South and Central America, with one species inhabi ...

**** Superfamily

Rhinocerotoidea Rhinocerotoidea is a superfamily (taxonomy), superfamily consisting of three family groups of odd-toed ungulates, three of which, the Amynodontidae, Hyracodontidae, and Paraceratheriidae, are extinct. The only extant family group is the Rhinocer ...

***** † Hyrachyidae (

Hyracodontidae The Hyracodontidae are an extinct family of rhinoceroses endemic to North America, Europe, and Asia during the Eocene through early Oligocene, living from 48.6 to 26.3 million years ago (Mya), existing about . The Hyracodontidae thrived in the r ...

?) ***** †

***** †

Amynodontidae Amynodontidae ("defensive tooth") is a family of extinct perissodactyls related to true rhinoceroses. They are commonly portrayed as semiaquatic hippo-like rhinos but this description only fits members of the Metamynodontini; other groups of a ...

(

Rhinocerotidae A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species ...

?) *****

*** Infraorder †

Ancylopoda Ancylopoda is a group of browsing, herbivorous, mammals in the Perissodactyla that show long, curved and cleft claws. Morphological evidence indicates the Ancylopoda diverged from the tapirs, rhinoceroses and horses ( Euperissodactyla) after t ...

**** †

Eomoropidae Eomoropidae is a family of odd-toed ungulates, a group which also includes horses, rhinoceroses, and tapirs. They were most closely related to the extinct chalicotheres, which they greatly resemble, and may have been their immediate ancestors. The ...

**** †

Chalicotheriidae Chalicotheres (from Greek '' chalix'', "gravel" and '' therion'', "beast") are an extinct clade of herbivorous, odd-toed ungulate (perissodactyl) mammals that lived in North America, Eurasia, and Africa from the Middle Eocene until the Early Pl ...

**** †

Lophiodontidae Lophiodontidae is a family of browsing, herbivorous, mammals in the Perissodactyla suborder Ancylopoda that show long, curved and cleft claws. They lived in southern Europe during the Eocene epoch. Previously thought to be related to tapir T ...

(

?) ** Suborder † Titanotheriomorpha *** Superfamily † Brontotherioidea **** †

Brontotheriidae Brontotheriidae is a family of extinct mammals belonging to the order Perissodactyla, the order that includes horses, rhinoceroses, and tapirs. Superficially, they looked rather like rhinos, although they were actually more closely related to ...

( equoids) **** † Anchilophidae ( palaeotheriid equoids?) * Order

Uranotheria 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 ...

) ** Suborder

Hyracoidea Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simil ...

(sister taxon of Perissodactyla?) *** †

Pliohyracidae Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simi ...

***

Procaviidae Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simi ...

** Suborder

Tethytheria Tethytheria is a clade of mammals that includes the sirenians and proboscideans, as well as the extinct order Embrithopoda. Though there is strong anatomical and molecular support for the monophyly of Tethytheria, the interrelationships between ...

*** Infraorder †

Embrithopoda Embrithopoda ("heavy-footed") is an order of extinct mammals known from Asia, Africa and Eastern Europe. Most of the embrithopod genera are known exclusively from jaws and teeth dated from the late Paleocene to the late Eocene; however, the ord ...

**** †

Arsinoitheriidae Arsinoitheriidae is a family of hoofed mammals belonging to the extinct order Embrithopoda. Remains have been found in the Middle East, Africa, Asia and Romania. When alive, they would have borne a strong but superficial resemblance to modern rh ...

*** Infraorder

Sirenia The Sirenia (), commonly referred to as sea-cows or sirenians, are an order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently comprise two distinct f ...

**** †

Prorastomidae Prorastomidae is a family of extinct sirenians from Jamaica, related to the extant manatees and dugong. The family includes the oldest known fossils of Sirenians, represented in two genera: *''Pezosiren'' ''Pezosiren'' comprises one known species ...

**** †

Protosirenidae Protosirenidae is an extinct primitive family of the order Sirenia. Protosirenids are thought to have been amphibious quadrupeds, meaning that they spent their time both on land and in the water and had four legs. See also *Dugongidae * Evoluti ...

(

Dugongidae Dugongidae is a family in the order of Sirenia. The family has one surviving species, the dugong (''Dugong dugon''), one recently extinct species, Steller's sea cow (''Hydrodamalis gigas''), and a number of extinct genera known from fossil rec ...

?) ****

****

Trichechidae Trichechidae is a family of sirenians that includes all living manatees and several extinct genera. Systematics TRICHECHIDAE *MiosireninaeM. Voss. 2014. On the invalidity of Halitherium schinzii Kaup, 1838 (Mammalia, Sirenia), with comments on ...

*** Infraorder †

Desmostylia The Desmostylia (from Greek δεσμά ''desma'', "bundle", and στῦλος ''stylos'', "pillar") are an extinct order of aquatic mammals that existed from the early Oligocene (Rupelian) to the late Miocene (Tortonian) (). Desmostylians are t ...

**** †

Desmostylidae Desmostylidae is an extinct family (biology), family of herbivorous marine mammals belonging to the order of Desmostylia. They lived in the coastal waters of the northern Pacific Ocean from the Early Oligocene (Rupelian) through the Late Miocene ...

***Infraorder

Proboscidea The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. From ...

**** †

Anthracobunidae Anthracobunidae is an extinct family of stem perissodactyls that lived in the early to middle Eocene period. They were originally considered to be a paraphyletic family of primitive proboscideans possibly ancestral to the Moeritheriidae and the ...

(tethytheres?) **** † Phosphatheriidae (

Numidotheriidae Numidotheriidae is an extinct family of primitive proboscidean that lived from the late Paleocene to the early Oligocene periods of North Africa. Fragmentary fossils (mainly teeth) of the early Eocene genera, ''Daouitherium'' and ''Phosphatheri ...

?) **** †

**** †

Moeritheriidae ''Moeritherium'' ("the beast from Lake Moeris") is an extinct genus of primitive proboscideans. These prehistoric mammals are related to the elephant and, more distantly, sea cows and hyraxes. They lived during the Eocene epoch. Description ''Mo ...

**** †

Barytheriidae Barytheriidae (meaning "heavy beasts") is an extinct family of primitive proboscideans that lived during the late Eocene and early Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to ...

**** †

Deinotheriidae Deinotheriidae ("terrible beasts") is a family of prehistoric elephant-like proboscideans that lived during the Cenozoic era, first appearing in Africa, then spreading across southern Asia (Indo-Pakistan) and Europe. During that time, they cha ...

**** † Palaeomastodontidae **** † Phiomiidae **** † Hemimastodontidae **** †

Mammutidae Mammutidae is an extinct family of proboscideans that appeared during the Oligocene epoch and survived until the start of the Holocene. The family was first described in 1922, classifying fossil specimens of the type genus ''Mammut'' (mastodons) ...

**** †

Gomphotheriidae Gomphotheres are any members of the diverse, extinct taxonomic family Gomphotheriidae. Gomphotheres were elephant-like proboscideans, but do not belong to the family Elephantidae. They were widespread across Afro-Eurasia and North America during ...

****

Elephantidae Elephantidae is a family (biology), family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are terrestrial animal, terrestrial large mammals with a snout modified into a Elephant#Trunk, trunk and teeth ...

Notes

References

* * * * * * * {{Taxonbar, from=Q16556658 Obsolete mammal taxa