Meridiungulata Temporal range: | |
---|---|
Toxodon , a notoungulate | |
Thesodon , a litoptern | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Infraclass: | Placentalia |
Clade: | † Meridiungulata McKenna 1975 |
Orders | |
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 ago). They represented a dominant element of South America's Cenozoic terrestrial mammal fauna prior to the arrival of living unguate groups in South America during the Pliocene and Pleistocene as part of the Great American Interchange. They comprise five major groups conventionally ranked as orders—Astrapotheria, Litopterna, Notoungulata, Pyrotheria, and Xenungulata—as well as the primitive "condylarth" groups Didolodontidae and Kollpaniinae. 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. Only a few (mostly large) species of notoungulates and litopterns survived until the end-Pleistocene extinction event around 12,000 years ago where they became extinct with most other large mammals in the Americas, shortly after the first arrival of humans into the region.
Though most SANUs lived in South America, astrapotheres and litopterns are known from Eocene aged deposits in the Antarctic Peninsula [1] and the notoungulate Mixotoxodon spread as far north as what is now Texas during the Pleistocene as part of the Great American Biotic Interchange. [2]
The relationships of SANUs to living mammals has been historically uncertain, though analysis of DNA and collagen suggests that at least notoungulates and litopterns are members of Laurasiatheria, and closely related to living perissodactyls.
Meridiungulata might have originated in South America from a North American condylarth ancestor, [3] and they may be members of the clade Laurasiatheria, related to other ungulates, including artiodactyls and perissodactyls. [4] It has, however, been suggested the Meridiungulata are part of a different macro-group of placental mammals called Atlantogenata. [5] [1]
Much of the evolution of meridiungulates occurred in isolation from other ungulates, a great example of convergent evolution. However, the argument that meridiungulates are related to artiodactyls and perissodactyls needs support from molecular sequencing. Some paleontologists have also challenged the monophyly of Meridiungulata by suggesting that the pyrotheres are more closely related to other mammals, such as Embrithopoda (an African order possibly related to elephants), than to other South American ungulates. [6]
Molecular sequence data from both collagen [7] [8] and mitochondrial DNA [9] supports that litopterns and notoungulates are most closely related to Perissodactyla (the group containing equids, rhinoceroses, and tapirs) among living mammals, as part of the clade Panperissodactyla, making them true ungulates, which has also been supported by some analyses of morphology. [10] [11] However, other morphological analyses have placed Litopterna elsewhere within Laurasiatheria. [12] [13] Didolodontids may be closely related to litopterns, [12] [14] and it has been proposed that they should be classified within Litopterna, [15] but some analyses do not find them to be close relatives. [10] [11]
Molecular sequence data from collagen suggests Notoungulata and Litopterna are more closely related to each other than to Perissodactyla, suggesting that at least part of Meridiungulata is monophyletic. [7] [8] By contrast, morphology-based analyses have found a range of possible positions for notoungulates. They have been found to be elsewhere within Laurasiatheria, [13] within Afrotheria, [12] [14] [16] and as stem-group atlantogenatans. [13] 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. [1]
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. [8] [1]
The clade Sudamericungulata has been proposed to encompass astrapotheres, notoungulates, pyrotheres, and xenoungulates but not litopterns. [14] Such a clade had been found in previous studies, but left unnamed. [16] [11] The study proposing the name Sudamericungulata found them to be afrotheres. [14] The study proposing Sudamericungulata was questioned in a later study, who suggested that the taxon and character sampling in the analysis was poor (including only a single perissodactyl), and that the placement of Sudamericungulata within Afrotheria was not robustly supported, and that a placement within Laurasiatheria was supported for Sudamericungulata and Litopterna when Afrotheria and Laurasiatheria were constrained to be monophyletic by molecular results. [17]
A 2024 study based on morphology supported the monophyly of Meridiungulata as traditionally defined. [18]
The earliest SANUs appeared during the early Paleocene, around 63-65 million years ago. SANU diversity reached its greatest extent during the late Eocene and Oligocene periods. During the Miocene, genus and species diversity was stable, but family diversity declined. During the Pliocene and Pleistocene epochs, SANU diversity substantially declined to the point that there were only a handful of living species by the Late Pleistocene. [1] The causes of the decline are unclear, but may be due to climatic change, [19] or competition/predation from new arrivals from North America as part of the Great American interchange. [1]
Perissodactyla is an order of ungulates. The order includes about 17 living species divided into three families: Equidae, Rhinocerotidae (rhinoceroses), and Tapiridae (tapirs). They typically have reduced the weight-bearing toes to three or one of the five original toes, though tapirs retain four toes on their front feet. The nonweight-bearing toes are either present, absent, vestigial, or positioned posteriorly. By contrast, artiodactyls bear most of their weight equally on four or two of the five toes: their third and fourth toes. Another difference between the two is that perissodactyls digest plant cellulose in their intestines, rather than in one or more stomach chambers as artiodactyls, with the exception of Suina, do.
Ungulates are members of the diverse clade Euungulata, 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 be a polyphyletic and thereby invalid clade based on molecular data. As a result, true ungulates had since been reclassified to the newer clade Euungulata in 2001 within the clade Laurasiatheria while Paenungulata has been reclassified to a distant clade Afrotheria. Living ungulates are divided into two orders: Perissodactyla including equines, rhinoceroses, and tapirs; and Artiodactyla including cattle, antelope, pigs, giraffes, camels, sheep, deer, and hippopotamuses, among others. Cetaceans such as whales, dolphins, and porpoises are also classified as artiodactyls, although they do not have hooves. Most terrestrial ungulates use the hoofed tips of their toes to support their body weight while standing or moving. Two other orders of ungulates, Notoungulata and Litopterna, both native to South America, became extinct at the end of the Pleistocene, around 12,000 years ago.
Afrotheria 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, elephant shrews, otter shrews, tenrecs, aardvarks, hyraxes, elephants, sea cows, and several extinct clades. Most groups of afrotheres share little or no superficial resemblance, and their similarities have only become known in recent times because of genetics and molecular studies. Many afrothere groups are found mostly or exclusively in Africa, reflecting the fact that Africa was an island continent from the Cretaceous until the early Miocene around 20 million years ago, when Afro-Arabia collided with Eurasia.
Litopterna is an extinct order of South American native ungulates that lived from the Paleocene to the end of the Pleistocene-early Holocene around 63 million-12,000 years ago, and were also present in Antarctica during the Eocene. They represent the second most diverse group of South American ungulates after Notoungulata. It is divided into nine families, with Proterotheriidae and Macraucheniidae being the most diverse and last surviving families.
Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene 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 Artiodactyla, being composed thus of several unrelated lineages.
Toxodon is an extinct genus of large ungulate native to South America from the Pliocene to the end of the Late Pleistocene. Toxodon is a member of Notoungulata, an order of extinct South American native ungulates distinct from the two living ungulate orders that had been indigenous to the continent for over 60 million years since the early Cenozoic, prior to the arrival of living ungulates into South America around 2.5 million years ago during the Great American Interchange. Toxodon is a member of the family Toxodontidae, which includes medium to large sized herbivores. Toxodon was one of the largest members of Toxodontidae and Notoungulata, with Toxodon platensis having an estimated body mass of 1,000–1,200 kilograms (2,200–2,600 lb).
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 resembling animals as disparate as rabbits and rhinoceroses. Notoungulata are the largest group of South American native ungulates, with over 150 genera in 14 families having been described, divided into two major subgroupings, Typotheria and Toxodontia. Notoungulates first diversified during the Eocene. Their diversity declined from the late Neogene onwards, with only the large toxodontids persisting until the end of the Pleistocene, perishing as part of the Late Pleistocene megafauna extinctions along with most other large mammals across the Americas. Collagen sequence analysis suggests that notoungulates are closely related to litopterns, another group of South American ungulates, and their closest living relatives being perissodactyls, including rhinoceroses, tapirs and equines as part of the clade Panperissodactyla. However their relationships to other South American ungulates are uncertain. Several groups of notoungulates separately evolved ever-growing cheek teeth.
Macrauchenia is an extinct genus of large ungulate native to South America from the Pliocene or Middle Pleistocene to the end of the Late Pleistocene. It is a member of the extinct order Litopterna, a group of South American native ungulates distinct from the two orders which contain all living ungulates which had been present in South America since the early Cenozoic, over 60 million years ago, prior to the arrival of living ungulates in South America around 2.5 million years ago as part of the Great American Interchange. The bodyform of Macrauchenia has been described as similar to a camel, being one of the largest-known litopterns, with an estimated body mass of around 1 tonne. The genus gives its name to its family, Macraucheniidae, which like Macrauchenia typically had long necks and three-toed feet, as well as a retracted nasal region, which in Macrauchenia manifests as the nasal opening being on the top of the skull behind the eye sockets. This has historically been argued to correspond to the presence of a tapir-like proboscis, though recent authors suggest a moose-like prehensile lip or a saiga antelope-like nose to filter dust are more likely.
Macraucheniidae is a family in the extinct South American ungulate order Litopterna, that resembled camelids. They had three functional digits on the fore and hind feet, as well as elongate necks. The family is generally divided up into two subfamilies, Cramaucheniidae and Macraucheniinae. The family shows retraction of the nasal region, most extremely to the top of the skull in derived macraucheniine taxa like Macrauchenia. which has been interpreted to have supported a probsocis, perhaps like that of a saiga antelope to filter dust, or a moose-like prehensile lip. The earliest unambiguous members of the family date to the late Oligocene around 30 millon years ago. Polymorphis from the Eocene has historically been placed as a macraucheniid, but this has been doubted. Most early representatives had a body masses in the range of 80–120 kilograms (180–260 lb), though some like Llullataruca were as small as 35–55 kilograms (77–121 lb), and the last representatives of the family from the Pleistocene like Macrauchenia were over 1,000 kilograms (2,200 lb). The family reached its apex of diversity during the late Miocene around 10-6 million years ago, before declining to only a few of species belong to the genera Macrauchenia and Xenorhinotherium by the Late Pleistocene.
Pyrotheria is an order of extinct meridiungulate mammals. These elephant-like ungulates include the genera Baguatherium, Carolozittelia, Colombitherium, Griphodon, Propyrotherium, Proticia, and Pyrotherium.
Xenungulata is an order of extinct and primitive South American hoofed mammals that lived from the Late Paleocene to Early Eocene. Fossils of the order are known from deposits in Brazil, Argentina, Peru, and Colombia. The best known member of this enigmatic order is the genus Carodnia, a tapir-like and -sized animal with a gait similar to living African elephants.
Panperissodactyla is a clade of ungulates containing living order Perissodactyla and all extinct ungulates more closely related to Perissodactyla than to Artiodactyla.
Cramauchenia is an extinct genus of litoptern South American ungulate. Cramauchenia was named by Florentino Ameghino. The name has no literal translation. Instead, it is an anagram of the name of a related genus Macrauchenia. This genus was initially discovered in the Sarmiento Formation in the Chubut Province, in Argentina, and later it was found in the Chichinales Formation in the Río Negro Province and the Cerro Bandera Formation in Neuquén, also in Argentina, in sediments assigned to the SALMA Colhuehuapian, as well as the Agua de la Piedra Formation in Mendoza, in sediments dated to the Deseadan. In 1981 Soria made C. insolita a junior synonym of C. normalis. A specimen of C. normalis was described in 2010 from Cabeza Blanca in the Sarmiento Formation, in sediments assigned to the Deseadan SALMA.
Scalabrinitherium is an extinct genus of mammals of the family Macraucheniidae. Fossils of this animal were found among the fossils of prehistoric xenarthrans in the Ituzaingó Formation of Argentina.
Astrapotheria is an extinct order of South American and Antarctic hoofed mammals that existed from the late Paleocene to the Middle Miocene, 59 to 11.8 million years ago. Astrapotheres were large, rhinoceros-like animals and have been called one of the most bizarre orders of mammals with an enigmatic evolutionary history.
Toxodontidae is an extinct family of notoungulate mammals, known from the Oligocene to the Holocene of South America, with one genus, Mixotoxodon, also known from the Pleistocene of Central America and southern North America. Member of the family were medium to large-sized, ranging from around 350–400 kilograms (770–880 lb) in Nesodon to 1,000–1,200 kilograms (2,200–2,600 lb) in Toxodon, and had medium to high-crowned dentition, which in derived members of the group evolved into ever-growing cheek teeth. Isotopic analyses have led to the conclusion that Pleistocene members of the family were flexible mixed feeders.
Proterotheriidae is an extinct family of litoptern ungulates known from the Eocene-Late Pleistocene of South America. Members of the group were small-medium sized cursorial herbivores with brachydont teeth, with their toes showing progressive reduction, with later members of the group bearing weight on a single large toe similar to living horses.
Eoastrapostylops is an extinct genus of astrapothere that lived during the Late Paleocene in what is now Argentina.
Sparnotheriodontidae is an enigmatic extinct family of litopterns known primarily from teeth. Sparnotheriodontids are one of two South American native ungulate clades known to have reached Antarctica, the other being astrapotheres.
The Sarmiento Formation, in older literature described as the Casamayor Formation, is a geological formation in Chubut Province, Argentina, in central Patagonia, which spans around 30 million years from the mid-Eocene to the early Miocene. It predominantly consists of pyroclastic deposits, which were deposited in a semi-arid environment. It is divided up into a number of members. The diverse fauna of the Sarmiento Formation, including a variety of birds, crocodilians, turtles and snakes, also includes many mammals such as South American native ungulates as well as armadillos, and caviomorph rodents.