Condylarthra is an informal group – previously considered an
order
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
– of extinct
placental mammal
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 distinguish ...
s, known primarily from the
Paleocene
The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...
and
Eocene
The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
epochs.
They are considered early, primitive
ungulates. It is now largely considered to be a
wastebasket taxon, having served as a dumping ground for classifying ungulates which had not been clearly established as part of either
Perissodactyla or
Cetartiodactyla, being composed thus of several unrelated lineages.
Taxonomic history
Condylarthra always was a problematic group. When Condylarthra was first described by ,
Phenacodontidae was the type and only family therein. , however, raised Condylarthra to an order and included a wide range of diverse placentals with generalized
dentitions and
postcranial skeletons. More recent researchers (i.e. post-WW2) have been more restrictive; either including only a limited number of taxa, or proposing that the term should be abandoned altogether.
Due to their primitive characteristics condylarths have been considered ancestral to several ungulate orders, including the living
Artiodactyla,
Cetacea,
Perissodactyla,
Hyracoidea,
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 ...
, and
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. Fr ...
, as well as the extinct
Desmostylia,
Embrithopoda,
Litopterna,
Notoungulata, and
Astrapotheria.
delimited condylarths as those having the following characters, but lacking the specializations present in more derived orders:
* superior ramus of
stapedial artery
In human anatomy, the stapedial branch of posterior auricular artery, or stapedial artery for short, is a small artery supplying the stapedius muscle in the inner ear.
Structure In humans
In humans, the stapedial artery is normally present in the ...
shifted to
petrosal or lost
*
mastoid foramen lost
*
bulla if present composed of
ectotympanic
* relatively
bunodont teeth with low cusp relief
*
trigonids of lower molars shortened anteroposteriorly
* large, posteriorly projecting
hypoconulid on M
3 (lower third molar)
* head of
astragalus
''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to te ...
is short and robust
Evolutionary history

The disappearance of the dinosaurs opened up an
ecological niche
In ecology, a niche is the match of a species to a specific environmental condition.
Three variants of ecological niche are described by
It describes how an organism or population responds to the distribution of resources and competitors (fo ...
for large mammalian
herbivore
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthp ...
s. Some condylarths evolved to fill the niche, while others remained insectivorous. This may explain, in part, the tremendous
evolutionary radiation
An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rap ...
of the condylarths that we can observe throughout the Paleocene, resulting in the different groups 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, giraff ...
s (or "
hoofed mammals") that form the
dominant herbivores in most
Cenozoic
The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configu ...
animal communities on land, except on the
island
An island or isle is a piece of subcontinental land completely surrounded by water. Very small islands such as emergent land features on atolls can be called islets, skerries, cays or keys. An island in a river or a lake island may be ...
continent of
Australia.
Among recent mammals,
Paenungulata (
hyraxes,
elephant
Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae ...
s, and
sea cows),
Perissodactyla (
horse
The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million ...
s,
rhinoceros
A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates in the family (biology), family Rhinocerotidae. (It can also refer to a member ...
es, and
tapir
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 inh ...
s),
Artiodactyla (
pigs,
deer
Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the ...
,
antelope
The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia.
Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mamm ...
,
cows,
camels,
hippos, and their relatives),
Cetacea (
whale
Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
s), and
Tubulidentata (
aardvarks) are traditionally regarded as members of the Ungulata.
Besides these, several extinct animals also belong to this group, especially the endemic
Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found els ...
South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the souther ...
n orders of ungulates, ( Meridiungulata). Although many ungulates have hooves, this feature does not define the Ungulata. Indeed, some condylarths had small hooves on their feet, but the most primitive forms are claw
A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tars ...
ed.
Recent molecular and DNA research has reorganised the picture of mammalian evolution. Paenungulates and tubulidentates are seen as 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 ...
ns, and no longer seen as closely related to the 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 ungula ...
n perissodactyls, artiodactyls, and cetaceans, implying that hooves were acquired independently (i.e. were analogous) by at least two different mammalian lineages, once in the Afrotheria and once in the Laurasiatheria. Condylarthra itself, therefore, is polyphyletic
A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
: the several condylarth groups are not closely related to each other at all. Indeed, Condylarthra is sometimes regarded as a 'wastebasket' taxon. True relationships remain in many cases unresolved.
In addition to meridiungulates and living ungulates, a condylarthran ancestry has been proposed for several other extinct groups of mammals, including Mesonychia and Dinocerata.
Taxonomy
*Family Arctocyonidae (possibly polyphyletic
A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
assemblage)
**Genus '' Arctocyon''
**Genus ''Chriacus
''Chriacus'' is an extinct genus of placental mammals that lived in what is now North America during the Paleocene epoch and died out after the early Eocene. In life, members of the genus would have looked something like a kinkajou or binturong, ...
''
*Family Periptychidae
**Genus ''Ectoconus
''Ectoconus'' (Greek: "outside" (ektos), "cone" (konos)) is an extinct genus of terrestrial herbivorous mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary ...
''
**Genus '' Oxyacodon''
*Family Hyopsodontidae (now established as within Perissodactyla)
**Subfamily '' Tricuspiodontinae''
***Genus '' Litomylus''
***Genus '' Paratricuspiodon''
***Genus '' Tricuspiodon''
**Genus '' Aletodon''
**Genus '' Decoredon''
**Genus '' Dipavali''
**Genus '' Dorraletes''
**Genus '' Haplaletes''
**Genus '' Haplomylus''
**Genus '' Hyopsodus''
**Genus '' Louisina''
**Genus ''Microhyus
''Microhyus'' is a genus of true weevils in the beetle family Curculionidae
The Curculionidae are a family of weevils, commonly called snout beetles or true weevils. They are one of the largest animal families, with 6,800 genera and 83,000 spe ...
''
**Genus '' Midiagnus''
**Genus '' Oxyprimus''
**Genus '' Palasiodon''
**Genus ''Paschatherium
''Paschatherium'' is a small extinct mammal of the Perissodactyla order, with an insectivore-like dentition.
Its morphology indicates an arboreal form, adapted climbing and running on trees. ''Paschatherium'' must have been extremely numerous in ...
''
**Genus '' Utemylus''
**Genus '' Yuodon''
*Family Mioclaenidae
*Family Phenacodontidae (established as stem- Perissodactyla)
**Subfamily Meniscotheriinae
''Meniscotherium'' is an extinct genus of dog-sized mammal which lived 54–38 million years ago. It was a herbivore and had hooves. Fossils have been found in Utah, New Mexico. and Colorado. Many individuals have been found together, indicating ...
***Genus '' Ectocion''
***Genus '' Meniscotherium''
**Genus '' Almogaver''
**Genus '' Copecion''
**Genus ''Eodesmatodon
''Eodesmatodon'' was an early herbivorous mammal that was part of the Aegialodontidae family
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by ...
''
**Genus '' Phenacodus''
*Family Pleuraspidotheriidae
Pleuraspidotheriidae is a family of "condylarths" that lived in Europe from the Palaeocene to the Mid Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the seco ...
**Genus '' Hilalia''
**Genus '' Orthaspidotherium''
**Genus '' Parabunodon''
**Genus ''Pleuraspidotherium
''Pleuraspidotherium'' is an extinct genus of condylarth of the family Pleuraspidotheriidae, whose fossils have been found in the Late Paleocene Marnes de Montchenot of France and the Tremp Formation of modern Spain
, image_flag = ...
''
*Family Didolodontidae (stem- Meridiungulata)
*Family Sparnotheriodontidae? ( Litopterna)
*Genus '' Tingamarra''? (non-descript therian mammal)
*Genus '' Protungulatum'' (either non-placental eutherian or basal artiodactyl).
*Genus ''Kharmerungulatum
Kharmerungulatum is an extinct genus of herbivorous mammal from the Late Cretaceous (Maastrichtian) Intertrappean Beds of Madhya Pradesh, India. Its specific epithet honors Leigh Van Valen
Leigh Van Valen (August 12, 1935 – October 16, 2010) ...
'' (a zhelestid
Zhelestidae is a lineage of extinct eutherian mammals. Occurring in the Late Cretaceous from the Turonian to the Maastrichtian, they were an extremely successful group, with representatives present in Europe, Asia, India (and subsequently in Mad ...
[James David Archibald · Alexander Olegovich Averianov]
Phylogenetic analysis, taxonomic revision, and dental ontogeny of the Cretaceous Zhelestidae (Mammalia: Eutheria)
Article · Feb 2012 · Zoological Journal of the Linnean Society)
See also
* Evolution of mammals
* List of prehistoric mammals
Notes
References
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External links
{{Authority control
Obsolete mammal taxa