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Condylarthra is an informal group – previously considered an
order 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 ...
– 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 (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
and
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
epochs. They are considered early, primitive
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 ...
and is now largely considered to be a
wastebasket taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists to refer to a taxon that has the purpose of classifying organisms that do not fit anywhere else. They are typically defined by e ...
, having served as a dumping ground for classifying ungulates which had not been clearly established as part of either
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 ...
or
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 ...
, being composed thus of several unrelated lineages.


Taxonomic history

Condylarthra always was a problematic group. When 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
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
s 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 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 ...
,
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 ...
,
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 ...
,
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 ...
,
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 ...
, and
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 ...
, as well as the extinct Desmostylia,
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; ...
,
Litopterna Litopterna (from "smooth heel") is an extinction, extinct order of South American native ungulates that lived from the Paleocene to the Pleistocene-Holocene around 62.5 million to 12,000 years ago (or possibly as late as 3,500 years ago), and we ...
,
Notoungulata Notoungulata is an extinct order of ungulates that inhabited South America from the early Paleocene to the end of the Pleistocene, living from approximately 61 million to 11,000 years ago. Notoungulates were morphologically diverse, with forms re ...
, and
Astrapotheria Astrapotheria is an extinct order of South American and Antarctic hoofed mammals that existed from the late Paleocene to the Middle Miocene, ."The uruguaytheriine Astrapotheriidae from the rich middle Miocene Honda Group of the upper Magdalena R ...
. 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 ...
shifted to petrosal or lost *
mastoid foramen The mastoid foramen is a hole in the posterior border of the temporal bone. It transmits an emissary vein between the sigmoid sinus and the suboccipital venous plexus, and a small branch of the occipital artery, the posterior meningeal artery t ...
lost * bulla if present composed of
ectotympanic The ectotympanic, or tympanicum, is a bony structure found in all mammals, located on the tympanic part of the temporal bone, which holds the tympanic membrane (eardrum) in place. In catarrhine primates (including humans), it takes a tube-shape. ...
* relatively bunodont teeth with low cusp relief *
trigonid The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone tooth ...
s of lower molars shortened anteroposteriorly * large, posteriorly projecting hypoconulid on M3 (lower third molar) * head of
astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
is short and robust


Evolutionary history

The disappearance of the non-avian 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 Resource (biology), resources an ...
for large mammalian
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
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. A significantly large and diverse radiation within ...
of the condylarths that we can observe throughout the Paleocene, resulting in the different groups of
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 (or "
hoof The hoof (: hooves) is the tip of a toe of an ungulate mammal, which is covered and strengthened with a thick and horny keratin covering. Artiodactyls are even-toed ungulates, species whose feet have an even number of digits; the ruminants with ...
ed mammals") that form the dominant herbivores in most
Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...
animal communities on land, except on the
island An island or isle is a piece of land, distinct from a continent, completely surrounded by water. There are continental islands, which were formed by being split from a continent by plate tectonics, and oceanic islands, which have never been ...
continent of
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
. Among recent mammals,
Paenungulata Paenungulata (from Latin ''paene'' "almost" + ''ungulātus'' "having hoofs") is a clade of "sub- ungulates", which groups three extant mammal orders: Proboscidea (including elephants), Sirenia ( sea cows, including dugongs and manatees), and ...
(
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,
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, and sea cows),
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 ...
(
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 mi ...
s,
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 ...
es, and
tapir Tapirs ( ) are large, herbivorous mammals belonging to the family Tapiridae. They are similar in shape to a Suidae, pig, with a short, prehensile nose trunk (proboscis). Tapirs inhabit jungle and forest regions of South America, South and Centr ...
s),
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 ...
(
pig The pig (''Sus domesticus''), also called swine (: swine) or hog, is an omnivorous, domesticated, even-toed, hoofed mammal. It is named the domestic pig when distinguishing it from other members of the genus '' Sus''. Some authorities cons ...
s,
deer A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) ...
,
antelope The term antelope refers to numerous extant or recently extinct species of the ruminant artiodactyl family Bovidae that are indigenous to most of Africa, India, the Middle East, Central Asia, and a small area of Eastern Europe. Antelopes do ...
,
cow Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are called co ...
s,
camel A camel (from and () from Ancient Semitic: ''gāmāl'') is an even-toed ungulate in the genus ''Camelus'' that bears distinctive fatty deposits known as "humps" on its back. Camels have long been domesticated and, as livestock, they provid ...
s,
hippos A hippo or hippopotamus is either of two species of large African mammal which live mainly in and near water: * Hippopotamus * Pygmy hippopotamus Hippo or Hippos may also refer to: Toponymy * The ancient city of Hippo Regius (modern Annaba, Alg ...
, and their relatives),
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 ...
(
whale Whales are a widely distributed and diverse group of fully Aquatic animal, aquatic placental mammal, placental marine mammals. As an informal and Colloquialism, colloquial grouping, they correspond to large members of the infraorder Cetacea ...
s), and
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, ...
(
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) are traditionally regarded as members of the Euungulata. Besides these, several extinct animals also belong to this group, especially the
endemic Endemism is the state of a species being found only 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 foun ...
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
n orders of ungulates, (
Meridiungulata South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals that were indigenous to South America from the Paleocene (from at least 63 million years ago) until the end of the Late Pleistocene (~12,000 years a ...
). Although many ungulates have hooves, this feature does not define the Euungulata. 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 Arthro ...
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 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 ...
ns, and no longer seen as closely related to the
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 ...
n perissodactyls, artiodactyls, and cetaceans, implying that hooves were acquired independently (i.e. were
analogous Analogy is a comparison or correspondence between two things (or two groups of things) because of a third element that they are considered to share. In logic, it is an inference or an argument from one particular to another particular, as oppose ...
) 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 that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
: 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 Mesonychia ("middle claws") is an extinct taxon of small to large-sized carnivorous ungulates related to artiodactyls. Mesonychians first appeared in the early Paleocene, went into a sharp decline at the end of the Eocene, and died out entirely w ...
and
Dinocerata Dinocerata, from Ancient Greek (), "terrible", and (), "horn", or Uintatheria, is an extinct order of large herbivorous hoofed mammals with horns and protuberant canine teeth, known from the Paleocene and Eocene of Asia and North America. With ...
.


Taxonomy

*Family Arctocyonidae (possibly
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
assemblage) **Genus '' Arctocyon'' **Genus '' Chriacus'' *Family Periptychidae **Genus '' Ectoconus'' **Genus '' Oxyacodon'' *Family Hyopsodontidae (now established as within
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 ...
) **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 spec ...
'' **Genus '' Midiagnus'' **Genus '' Oxyprimus'' **Genus '' Palasiodon'' **Genus '' Paschatherium'' **Genus '' Utemylus'' **Genus '' Yuodon'' *Family Mioclaenidae *Family Phenacodontidae (established as stem-
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 ...
) **Subfamily Meniscotheriinae ***Genus '' Ectocion'' ***Genus '' Meniscotherium'' **Genus '' Almogaver'' **Genus '' Copecion'' **Genus '' Eodesmatodon'' **Genus '' Phenacodus'' *Family Pleuraspidotheriidae **Genus '' Hilalia'' **Genus '' Orthaspidotherium'' **Genus '' Parabunodon'' **Genus '' Pleuraspidotherium'' *Family Didolodontidae (stem-
Meridiungulata South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals that were indigenous to South America from the Paleocene (from at least 63 million years ago) until the end of the Late Pleistocene (~12,000 years a ...
) *Family Sparnotheriodontidae? (
Litopterna Litopterna (from "smooth heel") is an extinction, extinct order of South American native ungulates that lived from the Paleocene to the Pleistocene-Holocene around 62.5 million to 12,000 years ago (or possibly as late as 3,500 years ago), and we ...
) *Genus '' Tingamarra''? (non-descript therian mammal) *Genus ''
Protungulatum ''Protungulatum'' ('first ungulate') is an extinct genus of eutherian mammals within extinct family Protungulatidae, and is possibly one of the earliest known placental mammals in the fossil record, that lived in North America from the Late Cre ...
'' (either non-placental eutherian or basal artiodactyl). *Genus '' Kharmerungulatum'' (a zhelestidJames 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 The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian (geology), Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synaps ...
*
List of prehistoric mammals This is an incomplete list of prehistoric mammals. It does not include List of mammals, extant mammals or List of extinct mammals, recently extinct mammals. For extinct primate species, see: list of fossil primates.Mikko's Phylogeny Archiv Ma ...


Notes


References

* * * * * * * * * * *


External links

{{Authority control Obsolete mammal taxa