Paratheria (mammals)

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The term "Paratheria" was proposed as a result of the hypothesis that edentates, such as this brown-throated sloth (Bradypus variegatus), are distinct from other placental mammals. Bradypus.jpg
The term "Paratheria" was proposed as a result of the hypothesis that edentates, such as this brown-throated sloth (Bradypus variegatus), are distinct from other placental mammals.

Paratheria is an obsolete term for a taxonomic group including the xenarthran mammals (sloths, anteaters, and armadillos) and various groups thought to be related to them. It was proposed by Oldfield Thomas in 1887 to set apart the sloths, anteaters, armadillos, and pangolins, usually classified as placentals, from both marsupial and placental mammals, an arrangement that received little support from other workers. When teeth of the extinct gondwanathere mammals were first discovered in Argentina in the 1980s, they were thought to be related to xenarthrans, leading to renewed attention for the hypothesis that xenarthrans are not placentals. However, by the early 1990s, gondwanatheres were shown to be unrelated to xenarthrans, and xenarthrans are still considered to be placentals.

Contents

History

The term "Paratheria" was coined by British mammalogist Oldfield Thomas in 1887 in a review of tooth development in mammals. He found that the "Edentata" were especially distinctive. In this group, he included the sloths, anteaters, and armadillos, which are still placed together as Xenarthra, as well as the pangolins and the aardvark. According to Thomas, edentate teeth would be derived from the very earliest stage of mammalian dental evolution. Consequently, he suggested that they should be given a grouping separate from the other major groupings of mammals, for which terms had been introduced by Thomas Huxley: Eutheria (placentals) and Metatheria (marsupials). For this new grouping, he suggested the name Paratheria "to indicate their position by the side of, but separate from, the other Mammals" (the Greek παρά para means "beside"). [1] Thomas had included one other mammal among the edentates, the aardvark; however, he was unable to provide a satisfactory scenario for the origin of its wholly unique dentition, which he could only compare with that of some fish. [1] Thomas's arrangement was foreshadowed by Henri Marie Ducrotay de Blainville's 1839 classification; he placed edentates (except the sloth Bradypus , which he considered to be a primate) as a major division, the Maldentés ("poorly toothed"). This group was considered to be distinct from the other monodelphes (placentals), the Bien dentés ("well-toothed"). [2] Similarly, Paul Gervais proposed in 1855 that edentates should be placed in a separate subclass of mammals. [3]

Thomas's hypothesis received little support, or even attention, in subsequent years. In 1893, Henry Fairfield Osborn remarked that new studies of edentate teeth indicated that they were not as distinct as Thomas thought. [4] William Berryman Scott did, however, place Paratheria as a separate subclass in 1904, [5] although he apparently did not follow Thomas's theories about the origins of edentate teeth. [6] In 1910, William King Gregory reviewed the interrelationships of mammals and placed edentates among other placentals, [7] though he gave "Paratheria" as an alternative name for his superorder Edentata, which included Xenarthra and tentatively Pholidota (pangolins), Tubulidentata (aardvarks), and the fossil Taeniodonta. [8] In 1976, Eli Minkoff also used "Paratheria" for a placental superorder that included Edentata (for the sloths, armadillos, and anteaters) and Pholidota. [9]

The Paratheria hypothesis enjoyed a brief renaissance when unusual, high-crowned teeth began turning up in the Cretaceous and Paleocene fossil record of Argentina. [10] In 1984, Sudamerica ameghinoi , from the Paleocene of Argentina, was assigned to Xenarthra within Paratheria, ranked as a cohort (a taxonomic rank between infraclass and superorder). [11] Two years later, José Bonaparte named Gondwanatherium patagonicum from the Late Cretaceous of Argentina, which he thought to be related to Sudamerica, and tentatively assigned it to Paratheria, now ranked as an infraclass. [12] Bonaparte described an additional related animal, Vucetichia gracilis , from the Argentinean Late Cretaceous in 1990; by then he classified it in the order Gondwanatheria, which was tentatively assigned to the infraclass Paratheria. [13] Bonaparte argued against George Gaylord Simpson's 1931 view that xenarthrans derive from the Tertiary Palaeanodonta of North America, and instead suggested that xenarthrans, and perhaps pangolins, split from eutherians (placentals and their extinct relatives) as early as the Early Cretaceous and derived from some early "pantothere" [14] (a now-abandoned grouping of early mammals, including dryolestoids among others). [15]

However, Bonaparte himself had abandoned the proposed relationship between xenarthrans and gondwanatherians by 1993. [16] Instead, gondwanatherians were shown to be related to another Late Cretaceous Argentinean animal, Ferugliotherium (which turned out to be undistinguishable from Vucetichia), and through it to multituberculates. [16] The relation between multituberculates and gondwanatheres later became controversial, but they are no longer thought to be related to xenarthrans. [17] By 1996, "few if any systematists would ... doubt the eutherian affinities of xenarthrans" [10] and molecular data have also supported the placement of Xenarthra within placentals as one of four major clades. [18] The name "Paratheria" is no longer in use.

Related Research Articles

<span class="mw-page-title-main">Xenarthra</span> Superorder of mammals including anteaters, sloths, and armadillos

Xenarthra is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. Extinct xenarthrans include the glyptodonts, pampatheres and ground sloths. Xenarthrans originated in South America during the late Paleocene about 60 million years ago. They evolved and diversified extensively in South America during the continent's long period of isolation in the early to mid Cenozoic Era. They spread to the Antilles by the early Miocene and, starting about 3 million years ago, spread to Central and North America as part of the Great American Interchange. Nearly all of the formerly abundant megafaunal xenarthrans became extinct at the end of the Pleistocene.

<span class="mw-page-title-main">Multituberculata</span> Extinct order of mammals

Multituberculata is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, and reached a peak diversity during the Late Cretaceous and Paleocene. They eventually declined from the mid-Paleocene onwards, disappearing from the known fossil record in the late Eocene. They are the most diverse order of Mesozoic mammals with more than 200 species known, ranging from mouse-sized to beaver-sized. These species occupied a diversity of ecological niches, ranging from burrow-dwelling to squirrel-like arborealism to jerboa-like hoppers. Multituberculates are usually placed as crown mammals outside either of the two main groups of living mammals—Theria, including placentals and marsupials, and Monotremata—but usually as closer to Theria than to monotremes. They are considered to be closely related to Euharamiyida and Gondwanatheria as part of Allotheria.

<span class="mw-page-title-main">Placentalia</span> Infraclass of mammals in the clade Eutheria

Placental mammals 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 distinguished from monotremes and marsupials in that the fetus is carried in the uterus of its mother to a relatively late stage of development. The name is something of a misnomer considering that marsupials also nourish their fetuses via a placenta, though for a relatively briefer period, giving birth to less developed young which are then nurtured for a period inside the mother's pouch.

<span class="mw-page-title-main">Gondwanatheria</span> Extinct group of Mammaliaformes that lived during the Upper Cretaceous through the Miocene

Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa and Antarctica during the Upper Cretaceous through the Paleogene. Until recently, they were known only from fragmentary remains. They are generally considered to be closely related to the multituberculates and likely the euharamiyidians, well known from the Northern Hemisphere, with which they form the clade Allotheria.

Ferugliotherium is a genus of fossil mammals in the family Ferugliotheriidae from the Campanian and/or Maastrichtian period of Argentina. It contains a single species, Ferugliotherium windhauseni, which was first described in 1986. Although originally interpreted on the basis of a single brachydont (low-crowned) molar as a member of Multituberculata, an extinct group of small, rodent-like mammals, it was recognized as related to the hypsodont (high-crowned) Sudamericidae following the discovery of additional material in the early 1990s. After a jaw of the sudamericid Sudamerica was described in 1999, these animals were no longer considered to be multituberculates and a few fossils that were previously considered to be Ferugliotherium were assigned to unspecified multituberculates instead. Since 2005, a relationship between gondwanatheres and multituberculates has again received support. A closely related animal, Trapalcotherium, was described in 2009 on the basis of a single tooth.

Lavanify is a mammalian genus from the late Cretaceous of Madagascar. The only species, L. miolaka, is known from two isolated teeth, one of which is damaged. The teeth were collected in 1995–1996 and described in 1997. The animal is classified as a member of Gondwanatheria, an enigmatic extinct group with unclear phylogenetic relationships, and within Gondwanatheria as a member of the family Sudamericidae. Lavanify is most closely related to the Indian Bharattherium; the South American Sudamerica and Gondwanatherium are more distantly related. Gondwanatheres probably ate hard plant material.

<span class="mw-page-title-main">Afrotheria</span> Clade of mammals containing elephants and elephant shrews

Afrotheria 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, 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.

Ferugliotheriidae is one of three known families in the order Gondwanatheria, an enigmatic group of extinct mammals. Gondwanatheres have been classified as a group of uncertain affinities or as members of Multituberculata, a major extinct mammalian order. The best-known representative of Ferugliotheriidae is the genus Ferugliotherium from the Late Cretaceous epoch in Argentina. A second genus, Trapalcotherium, is known from a single tooth, a first lower molariform, from a different Late Cretaceous Argentinean locality. Another genus known from a single tooth, Argentodites, was first described as an unrelated multituberculate, but later identified as possibly related to Ferugliotherium. Finally, a single tooth from the Paleogene of Peru, LACM 149371, perhaps a last upper molariform, and a recent specimen from Mexico, may represent related animals.

<span class="mw-page-title-main">Mammal classification</span> Taxonomy of mammals

Mammalia is a class of animal within the phylum Chordata. Mammal classification has been through several iterations since Carl Linnaeus initially defined the class. No classification system is universally accepted; McKenna & Bell (1997) and Wilson & Reader (2005) provide useful recent compendiums. Many earlier ideas from Linnaeus et al. have been completely abandoned by modern taxonomists, among these are the idea that bats are related to birds or that humans represent a group outside of other living things. Competing ideas about the relationships of mammal orders do persist and are currently in development. Most significantly in recent years, cladistic thinking has led to an effort to ensure that all taxonomic designations represent monophyletic groups. The field has also seen a recent surge in interest and modification due to the results of molecular phylogenetics.

<i>Fruitafossor</i> Extinct family of mammals

Fruitafossor was a termite-eating mammal endemic to North America during the Late Jurassic epoch.

<span class="mw-page-title-main">Pilosa</span> Order of mammals

The order Pilosa is a clade of xenarthran placental mammals, native to the Americas. It includes anteaters and sloths. The name comes from the Latin word for "hairy".

<i>Doedicurus</i> An extinct genus of mammals belonging to the armadillo order, Cingulata

Doedicurus is an extinct genus of glyptodont from South America containing one species, D. clavicaudatus. Glyptodonts are a member of the family Chlamyphoridae, which also includes some modern armadillo species, and they are classified in the superorder Xenarthra alongside sloths and anteaters. Being a glyptodont, it was a rotund animal with heavy armor and a carapace. Averaging at an approximate 1,400 kg (3,100 lb), it was one of the largest glyptodonts to have ever lived. Though glyptodonts were quadrupeds, large ones like Doedicurus may have been able to stand on two legs like other xenarthrans. It notably sported a spiked tail club, which may have weighed 40 or 65 kg in life, and it may have swung this in defense against predators or in fights with other Doedicurus at speeds of perhaps 11 m/s.

<span class="mw-page-title-main">Myrmecophagidae</span> Family of mammals

The Myrmecophagidae are a family of anteaters, the name being derived from the Ancient Greek words for 'ant' and 'eat'. Two genera and three species are in the family, consisting of the giant anteater, and the tamanduas. The fossil Eurotamandua from the Messel Pit in Germany may be an early anteater, but its status is currently debated.

<i>Metacheiromys</i> Extinct genus of mammal

Metacheiromys is an extinct genus of placental mammals from extinct paraphyletic subfamily Metacheiromyinae within extinct paraphyletic family Metacheiromyidae in extinct order Palaeanodonta, that lived in North America from the early to middle Eocene.

<i>Ernanodon</i> Extinct genus of mammal

Ernanodon is an extinct genus of placental mammal from extinct family Ernanodontidae within extinct order Palaeanodonta, that lived from the middle to late Paleocene in China and Mongolia.

<span class="mw-page-title-main">Anteater</span> Mammals known for eating ants and termites

Anteaters are the four extant mammal species in the suborder Vermilingua, commonly known for eating ants and termites. The individual species have other names in English and other languages. Together with the sloths, they are within the order Pilosa. The name "anteater" is also commonly applied to the unrelated aardvark, numbat, echidnas, and pangolins, although they are not closely related to them.

Several mammals are known from the Mesozoic of Madagascar. The Bathonian Ambondro, known from a piece of jaw with three teeth, is the earliest known mammal with molars showing the modern, tribosphenic pattern that is characteristic of marsupial and placental mammals. Interpretations of its affinities have differed; one proposal places it in a group known as Australosphenida with other Mesozoic tribosphenic mammals from the southern continents (Gondwana) as well as the monotremes, while others favor closer affinities with northern (Laurasian) tribosphenic mammals or specifically with placentals. At least five species are known from the Maastrichtian, including a yet undescribed species known from a nearly complete skeleton that may represent a completely new group of mammals. The gondwanathere Lavanify, known from two teeth, is most closely related to other gondwanatheres found in India and Argentina. Two other teeth may represent another gondwanathere or a different kind of mammal. One molar fragment is one of the few known remains of a multituberculate mammal from Gondwana and another has been interpreted as either a marsupial or a placental.

Argentodites is a possible multituberculate mammal from the Cretaceous of Argentina. The single species, Argentodites coloniensis, is known from a single blade-like fourth lower premolar (p4) from the La Colonia Formation, which is mostly or entirely Maastrichtian in age. The p4 is 4.15 mm long and bears eight cusps on its upper margin and long associated ridges on both sides. The enamel consists of prisms that are completely or partly surrounded by a sheath and that are on average 6.57 μm apart. Zofia Kielan-Jaworowska, who described and named the fossil in 2007, regarded it as a multituberculate, perhaps a cimolodontan—and thus, a member of a mostly Laurasian (northern) group and an immigrant to Argentina from North America—on the basis of the shape of the tooth and features of its enamel. In 2009, however, two teams argued that Argentodites may in fact be close to or identical with Ferugliotherium, a member of the small Gondwanan (southern) group Gondwanatheria; although their relationships are disputed, gondwanatheres may themselves be multituberculates.

Groeberiidae is a family of strange non-placental mammals from the Eocene and Oligocene epochs of Patagonia, Argentina and Chile, South America. Originally classified as paucituberculate marsupials, they were suggested to be late representatives of the allothere clade Gondwanatheria. However, the relationship of the type genus, Groeberia, to Gondwanatheria has been firmly rejected by other scholars.

<span class="mw-page-title-main">Pholidotamorpha</span> Clade of mammals

Pholidotamorpha is a clade of placental mammals from mirorder Ferae that includes the order Pholidota and extinct order Palaeanodonta.

References

  1. 1 2 Thomas 1887, p. 459.
  2. Gregory 1910, p. 333.
  3. Gervais 1855, pp. 243–244.
  4. Osborn 1893, p. 500.
  5. Scott 1904, p. 3.
  6. Matthew & Granger 1918, p. 646.
  7. Gregory 1910, p. 342.
  8. Gregory 1910, p. 466.
  9. Minkoff 1976, p. 155.
  10. 1 2 Gaudin et al. 1996, p. 32.
  11. Gurovich 2005, p. 236.
  12. Bonaparte 1986, pp. 264, 269.
  13. Bonaparte 1990, p. 77.
  14. Bonaparte 1990, p. 85–86.
  15. Kielan-Jaworowska, Cifelli & Luo 2004, pp. 371–372.
  16. 1 2 Krause & Bonaparte 1993, p. 9379.
  17. Gurovich & Beck 2009, p. 25.
  18. Hallström & Janke 2008, p. 312.

Literature cited