South American native ungulates, commonly abbreviated as SANUs, are
extinct 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 ...
-like
mammals
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), fu ...
of controversial affinities that were indigenous to
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 ...
prior to the
Great American Biotic Interchange. They comprise five major groups conventionally ranked as
orders
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 ...
—
Astrapotheria,
Litopterna
Litopterna (from grc, λῑτή πτέρνα "smooth heel") is an extinct order of fossil hoofed mammals from the Cenozoic era. The order is one of the five great orders of South American ungulates that were endemic to the continent, until the G ...
,
Notoungulata
Notoungulata is an extinct order of mammalian ungulates that inhabited South America from the early Paleocene to the Holocene, living from approximately 61 million to 11,000 years ago. Notoungulates were morphologically diverse, with forms resem ...
,
Pyrotheria, and
Xenungulata
Xenungulata ("strange ungulates") is an order of extinct and primitive South American hoofed mammals that lived from the Late Paleocene to Early Eocene ( Itaboraian to Casamayoran in the SALMA classification). Fossils of the order are known fr ...
—as well as some other taxa, such as
Didolodontidae and
Kollpaniidae. It has been proposed that some or all of the members of this group form a clade, named Meridiungulata, though the relationships of South American ungulates remain largely unresolved. The two largest groups of South American ungulates, the notoungulates and the litopterns, were the only groups to persist beyond the mid
Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...
. Only a few of the largest species of notoungulates and litopterns survived until the end-
Pleistocene extinctions
The Quaternary period (from 2.588 ± 0.005 million years ago to the present) has seen the extinctions of numerous predominantly megafaunal species, which have resulted in a collapse in faunal density and diversity and the extinction of key ecolog ...
.
Though most SANUs lived in South America, astrapotheres and litopterns are known from
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'', " ...
aged deposits in the
Antarctic Peninsula
The Antarctic Peninsula, known as O'Higgins Land in Chile and Tierra de San Martín in Argentina, and originally as Graham Land in the United Kingdom and the Palmer Peninsula in the United States, is the northernmost part of mainland Antarctic ...
. and the notoungulate ''
Mixotoxodon'' spread as far north as what is now Texas during the Great American Biotic Interchange.
Taxonomy
Meridiungulata might have originated in
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 ...
from a
North American
condylarth ancestor,
and they may be members of the clade
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 ...
, related to other ungulates, including
artiodactyl
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poste ...
s and
perissodactyl
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) ...
s.
It has, however, been suggested the Meridiungulata are part of a different macro-group of placental mammals called
Atlantogenata
Atlantogenata is a proposed clade of mammals containing the cohorts or superorders Xenarthra and Afrotheria. These groups originated and radiated in the South American and African continents, respectively, presumably in the Cretaceous. Togeth ...
.
Much of the evolution of meridiungulates occurred in isolation from other ungulates, a great example of
convergent evolution
Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. However, the argument that meridiungulates are related to artiodactyls and perissodactyls needs support from molecular sequencing. Some
paleontologists
Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
have also challenged the
monophyly
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...
of Meridiungulata by suggesting that the pyrotheres are more closely related to other mammals, such as
Embrithopoda (an African order possibly related to
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), than to other South American ungulates.
Molecular sequence data from both collagen and mitochondrial DNA supports a placement of litopterns within the
total group of
Perissodactyla,
Panperissodactyla
Mesaxonia (near-synonymous with Panperissodactyla) is a clade of ungulates whose weight is distributed on the third toe on all legs through the plane symmetry of their feet. For a while it was often seen to only contain the order Perissodactyla ...
, which has also been supported by some analyses of morphology. However, other morphological analyses have placed Litopterna elsewhere within Laurasiatheria. Didolodontids may be closely related to litopterns, and it has been proposed that they should be classified within Litopterna, but some analyses do not find them to be close relatives.
Molecular sequence data from collagen supports a close relationship between Notoungulata and Litopterna within Panperissodactyla, suggesting that at least part of Meridiungulata is monophyletic. By contrast, morphology-based analyses have found a range of possible positions for notoungulates. They have been found to be elsewhere within Laurasiatheria, within
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 ...
, and as stem-group
atlantogenata
Atlantogenata is a proposed clade of mammals containing the cohorts or superorders Xenarthra and Afrotheria. These groups originated and radiated in the South American and African continents, respectively, presumably in the Cretaceous. Togeth ...
ns. A position within Afrotheria has been argued to be unlikely on biogeographic grounds, and some of the afrotherian characteristics present in notoungulates have been refuted.
Litopterns and notoungulates are the only South American ungulates to have gone extinct recently enough for molecular data to be available, so the relationships of astrapotheres, pyrotheres, and xenungulates must be determined based on morphology alone.
The clade Sudamericungulata has been proposed to encompass astrapotheres, notoungulates, pyrotheres, and xenoungulates but not litopterns. Such a clade had been found in previous studies, but left unnamed. The study proposing the name Sudamericungulata found them to be afrotheres.
Notes
References
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{{Taxonbar, from=Q735466
Cenozoic mammals of South America
Mammal unranked clades
Prehistoric mammals of South America
Ungulates
Taxa named by Malcolm McKenna