Paenungulata (from

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

''paene'' "almost" + ''ungulātus'' "having hoofs") is a

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

of "sub-

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 ...

", which groups three

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 ...

mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...

orders Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * H ...

Proboscidea Proboscidea (; , ) is 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. Three l ...

(including

elephant Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...

s),

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 extant Sirenia comprise two distinct famili ...

( sea cows, including

dugong The dugong (; ''Dugong dugon'') is a marine mammal. It is one of four living species of the order Sirenia, which also includes three species of manatees. It is the only living representative of the once-diverse family Dugongidae; its closest ...

s and

manatee Manatees (, family (biology), family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivory, herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing t ...

s), and

Hyracoidea Hyraxes (), also called dassies, are small, stout, thickset, herbivorous mammals in the family (biology), family Procaviidae within the Order (biology), order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Modern hyraxes ...

(

hyrax Hyraxes (), also called dassies, are small, stout, thickset, herbivorous mammals in the family Procaviidae within the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Modern hyraxes are typically between in length a ...

es). At least two more possible orders are known only as fossils, namely

Embrithopoda Embrithopoda ("heavy-footed") is an Order (biology), 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; ...

and

Desmostylia The Desmostylia (from Greek δεσμά ''desma'', "bundle", and στῦλος ''stylos'', "pillar") are an extinct order of aquatic mammals native to the North Pacific from the early Oligocene (Rupelian) to the late Miocene (Tortonian) (). Desmos ...

. Molecular evidence indicates that Paenungulata (or at least its extant members) is part of the cohort

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 ...

, an ancient assemblage of mainly

Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

n mammals of great diversity. The other members of this cohort are the orders

Afrosoricida The clade Afrosoricida (a Latin-Greek compound name which means "looking like African shrews") contains the golden moles of Southern Africa, the otter shrews of equatorial Africa and the tenrecs of Madagascar. These three groups of small mamma ...

(

tenrec A tenrec () is a mammal belonging to any species within the afrotherian family Tenrecidae, which is endemic to Madagascar. Tenrecs are a very diverse group, as a result of adaptive radiation, and exhibit convergent evolution, some resemble hed ...

s and

golden mole Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family Chrysochloridae (the only family in the suborder Chrysochloridea) and as such they are taxonomically distinct from the true moles, f ...

s), Macroscelidea (

elephant shrew Elephant shrews, also called jumping shrews or sengis, are small insectivore, insectivorous mammals native to Africa, belonging to the family Macroscelididae, in the order (biology), order Macroscelidea. Their traditional common English name "ele ...

s) and Tubulidentata (

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 ...

s). Of the five orders, hyraxes are the most basal, followed by embrithopods; the remaining orders (sirenians and elephants) are more closely related. These latter three are grouped as the Tethytheria, because it is believed that their common ancestors lived on the shores of the prehistoric

Tethys Sea The Tethys Ocean ( ; ), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean during much of the Mesozoic Era and early-mid Cenozoic Era. It was the predecessor to the modern Indian Ocean, the Mediterranean Sea, and the Eurasia ...

; however, recent

myoglobin Myoglobin (symbol Mb or MB) is an iron- and oxygen-binding protein found in the cardiac and skeletal muscle, skeletal Muscle, muscle tissue of vertebrates in general and in almost all mammals. Myoglobin is distantly related to hemoglobin. Compar ...

studies indicate that even

had an aquatic ancestor.

History

In 1945,

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 ...

used traditional taxonomic techniques to group these spectacularly diverse mammals in the superorder he named Paenungulata ("almost

ungulate 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 ...

s"), but there were many loose threads in unravelling their genealogy. For example, hyraxes in his Paenungulata had some characteristics suggesting they might be connected to the

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 ...

(odd-toed ungulates, such as horses and rhinos). Indeed, early taxonomists placed the Hyracoidea closest to the rhinoceroses because of their dentition. When genetic techniques were developed for inspecting

amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...

differences among

haemoglobin Hemoglobin (haemoglobin, Hb or Hgb) is a protein containing iron that facilitates the transportation of oxygen in red blood cells. Almost all vertebrates contain hemoglobin, with the sole exception of the fish family Channichthyidae. Hemoglobi ...

sequences the most

parsimonious In philosophy, Occam's razor (also spelled Ockham's razor or Ocham's razor; ) is the problem-solving principle that recommends searching for explanations constructed with the smallest possible set of elements. It is also known as the principle o ...

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

s depicted Simpson's Paenungulata as an authentic

and as one of the first groups to diversify from the basal placental mammals (

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 ...

). The amino acid sequences reject a connection between extant paenungulates and perissodactyls (odd-toed ungulates). However, a 2014 cladistic analysis placed anthracobunids and desmostylians, two major extinct groups that have been considered to be non-African afrotheres, close to each other within

Phylogeny

Gallery

Extinct orders

Each of the

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 ...

orders, the

and

, was as unique in its members' ways of making a living as the three orders that survive. Embrithopods were

rhinoceros A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...

-like herbivorous mammals with

plantigrade 151px, Portion of a human skeleton, showing plantigrade habit In terrestrial animals, plantigrade locomotion means walking with the toes and metatarsals flat on the ground. It is one of three forms of locomotion adopted by terrestrial mammals. ...

feet, and desmostylians were

hippopotamus The hippopotamus (''Hippopotamus amphibius;'' ; : hippopotamuses), often shortened to hippo (: hippos), further qualified as the common hippopotamus, Nile hippopotamus and river hippopotamus, is a large semiaquatic mammal native to sub-Sahar ...

-like amphibious animals. Their walking posture and diet have been the subject of speculation, but tooth wear indicates that desmostylians browsed on terrestrial plants and had a posture similar to other large hoofed mammals.

Notes

References

Sources

* * * *

External links

* {{Taxonbar, from=Q131950 Paenungulatomorpha Mammal taxonomy Mammals of Africa Extant Selandian first appearances Mammal unranked clades