notoungulata

Notoungulata is divided into two major suborders, Typotheria and Toxodontia, alongside some basal groups (Notostylopidae and Henricosborniidae) which are potentially paraphyletic.{{Cite journal|last1=Croft|first1=Darin A.|last2=Gelfo|first2=Javier N.|last3=López|first3=Guillermo M.|date=2020-05-30|title=Splendid Innovation: The Extinct South American Native Ungulates|url=https://www.annualreviews.org/doi/10.1146/annurev-earth-072619-060126|journal=Annual Review of Earth and Planetary Sciences|language=en|volume=48|issue=1|pages=259–290|doi=10.1146/annurev-earth-072619-060126|bibcode=2020AREPS..48..259C|s2cid=213737574 |issn=0084-6597|url-access=subscription}} Notoungulates make up over half the described diversity of indigenous South American ungulates, with over 150 genera in 14 different families.{{Cite journal |last1=Rezende Castro |first1=Luis Otavio |last2=García-López |first2=Daniel A. |last3=Bergqvist |first3=Lilian Paglarelli |last4=De Araújo-Júnior |first4=Hermínio Ismael |date=2021-06-30 |title=A New Basal Notoungulate from the Itaboraí Basin (Paleogene) of Brazil |url=https://bioone.org/journals/ameghiniana/volume-58/issue-3/AMGH.05.02.2021.3387/A-New-Basal-Notoungulate-from-the-Itabora%c3%ad-Basin-Paleogene-of/10.5710/AMGH.05.02.2021.3387.full |journal=Ameghiniana |volume=58 |issue=3 |doi=10.5710/AMGH.05.02.2021.3387 |issn=0002-7014|url-access=subscription }}

This order is proposed to be united with other South American native ungulates in the super-order Meridiungulata. The notoungulate and litoptern native ungulates of South America have been shown by studies of collagen and mitochondrial DNA sequences to be a sister group to the perissodactyls, making them true ungulates.{{Harvnb|Welker|Collins|Thomas|Wadsley|2015}}{{Harvnb|Buckley|2015}}{{Harvnb|Westbury|Baleka|Barlow|Hartmann|2017}} The estimated divergence date is 66 million years ago. This conflicts with the results of some morphological analyses which posited them as afrotherians. It is in line with some more recent morphological analyses which suggested they were basal euungulates. Panperissodactyla has been proposed as the name of an unranked clade to include perissodactyls and their extinct South American ungulate relatives.

Cifelli has argued that Notioprogonia is paraphyletic, as it would include the ancestors of the remaining suborders. Similarly, Cifelli indicated that Typotheria would be paraphyletic if it excluded Hegetotheria and he advocated inclusion of Archaeohyracidae and Hegetotheriidae in Typotheria.{{Harvnb|Cifelli|1993}}

Notoungulata were for many years taken to include the order Arctostylopida, whose fossils are found mainly in China. Recent studies, however, have concluded that Arctostylopida are more properly classified as gliriforms, and that the notoungulates were therefore never found outside South and Central America.{{Harvnb|Missiaen|Smith|Guo|Bloch|2006}}

Notoungulates are united by a number of morphological characters of the skull, particularly the inner ear and teeth.{{Cite journal |last=Macrini |first=Thomas E. |last2=Flynn |first2=John J. |last3=Ni |first3=Xijun |last4=Croft |first4=Darin A. |last5=Wyss |first5=André R. |date=November 2013

|title=Comparative study of notoungulate ( P lacentalia, M ammalia) bony labyrinths and new phylogenetically informative inner ear characters |url=https://onlinelibrary.wiley.com/doi/10.1111/joa.12108 |journal=Journal of Anatomy |language=en |volume=223 |issue=5 |pages=442–461 |doi=10.1111/joa.12108 |issn=0021-8782 |pmc=4399357 |pmid=24102069}}

Based on an analysis of 133 morphological characters in 50 notoungulate genera, Billet in 2011 concluded that Homalodotheriidae, Leontiniidae, Toxodontidae, Interatheriidae, Mesotheriidae, and Hegetotheriidae are the only monophyletic families of notoungulates. Some studies have suggested that Pyrotheria, often ranked as an independent order, should also be included within Notoungulata.{{Cite journal |last=Billet |first=Guillaume |date=December 2011 |title=Phylogeny of the Notoungulata (Mammalia) based on cranial and dental characters |url=http://www.tandfonline.com/doi/abs/10.1080/14772019.2010.528456 |journal=Journal of Systematic Palaeontology |language=en |volume=9 |issue=4 |pages=481–497 |doi=10.1080/14772019.2010.528456 |bibcode=2011JSPal...9..481B |issn=1477-2019|url-access=subscription }}

{{Clade |style=font-size:85%;line-height:90%;

|label1=Notoungulata

|1={{Clade

|1=Henricosbornia

|2={{Clade

|1=Simpsonotus

|2={{Clade

|1=Notostylops

|2=Pyrotherium

}}

|3={{Clade

|label1=Toxodontia

|1={{Clade

|1={{Clade

|1={{Clade

|1=Pampahippus

|2=Rhyphodon

}}

|2={{Clade

|1=Thomashuxleya

|2=Periphragnis

}}

}}

|2={{Clade

|1=Pleurostylodon

|2={{Clade

|label1=Homalodotheriidae

|1={{Clade

|1=Asmodeus

|2=Homalodotherium

}}

|label2=Leontiniidae

|2={{Clade

|1=Colpodon

|2=Ancylocoelus

|3={{Clade

|1=Leontinia

|2=Scarrittia

}}

}}

|3=Eomorphippus

|4=Eurygenium

|5=Rhynchippus

|6=Morphippus

|7={{Clade

|1=Pascualihippus

|2={{Clade

|1=Argyrohippus

|label2=Toxodontidae

|2={{Clade

|1=Nesodon

|2=Adinotherium

|3={{Clade

|1=Posnanskytherium

|2=Hoffstetterius

|3=Toxodon

}}

}}

}}

}}

}}

}}

}}

|label2=Typotheria

|2={{Clade

|1=Colbertia

|2=Oldfieldthomasia

|3=Campanorco

|4=Acropithecus

|5=Ultrapithecus

|label6=Interatheriidae

|6={{Clade

|1=Notopithecus

|2={{Clade

|1=Federicoanaya

|2=Protypotherium

|3=Miocochilius

|4={{Clade

|1=Archaeophylus

|2={{Clade

|1=Plagiarthrus

|2={{Clade

|1=Interatherium

|2=Cochilius

}}

}}

}}

}}

}}

|7={{Clade

|1=Eohyrax

|2={{Clade

|1=Pseudhyrax

|2={{Clade

|label1=Mesotheriidae

|1={{Clade

|1={{Clade

|1=Plesiotypotherium

|2=Mesotherium

}}

|2=Trachytherus

}}

|2=Archaeotypotherium

|3=Archaeohyrax

|label4=Hegetotheriidae

|4={{Clade

|1={{Clade

|1=Prohegetotherium

|2=Hegetotherium

}}

|2={{Clade

|1=Prosotherium

|2=Paedotherium

}}

}}

}}

}}

}}

}}

}}

}}

}}

}}

• Suborder Notioprogonia (probably paraphyletic)
• Family Henricosborniidae
• Family Notostylopidae
• Suborder Toxodontia
• Family Isotemnidae
• Family Leontiniidae
• Family Notohippidae (paraphyletic)
• Family Toxodontidae
• Family Homalodotheriidae
• Suborder Typotheria
• Family Archaeohyracidae
• Family Archaeopithecidae
• Family Campanorcidae
• Family Hegetotheriidae
• Family Interatheriidae
• Family Mesotheriidae
• Family Oldfieldthomasiidae

Notoungulates varied widely in body size, with early diverging notoungulates like Simpsonotus, and some hegetotheriid and interatheriid typotherians having a body mass of approximately {{Convert|1-2|kg|lb}}, while the toxodontid Toxodon is suggested to have had a body mass exceeding {{Convert|1000|kg|lb}}. Typotheres generally occupied small-medium body size niches, while toxodontians were generally medium-large sized animals. The families Interatheriidae, Hegetotheriidae, Mesotheriidae and Toxodontidae separately evolved high crowned (hypsodont) ever-growing (hypeselodont) cheek teeth,{{Cite journal |last1=Gomes Rodrigues |first1=Helder |last2=Herrel |first2=Anthony |last3=Billet |first3=Guillaume |date=2017-01-31 |title=Ontogenetic and life history trait changes associated with convergent ecological specializations in extinct ungulate mammals |journal=Proceedings of the National Academy of Sciences |language=en |volume=114 |issue=5 |pages=1069–1074 |doi=10.1073/pnas.1614029114 |issn=0027-8424 |pmc=5293108 |pmid=28096389 |doi-access=free|bibcode=2017PNAS..114.1069G }} with high crowned species constituting the majority of notoungulates from the Late Oligocene onward. This adaptation was historically suggested to be the result of a diet increasingly incorporating grass, but this has been questioned, and other authors suggesting that it may have been due to the increasing intake of abrasive particles from volcanic sources. Many typotheres have bodyforms convergent on rodents, hyraxes and rabbits,{{Citation |last1=Cassini |first1=Guillermo H. |title=Paleobiology of Santacrucian native ungulates (Meridiungulata: Astrapotheria, Litopterna and Notoungulata) |date=2012-10-11 |url=https://www.cambridge.org/core/product/identifier/9780511667381%23c19461-14-1/type/book_part |work=Early Miocene Paleobiology in Patagonia |pages=243–286 |editor-last=Vizcaíno |editor-first=Sergio F. |access-date=2023-06-29 |edition=1 |publisher=Cambridge University Press |doi=10.1017/cbo9780511667381.015 |isbn=978-0-511-66738-1 |last2=Cerdeño |first2=Esperanza |last3=Villafañe |first3=Amalia L. |last4=Muñoz |first4=Nahuel A. |editor2-last=Kay |editor2-first=Richard F. |editor3-last=Bargo |editor3-first=M. Susana|url-access=subscription }} with some rabbit-like hegetotheriids suggested to have developed a rabbit-like bounding locomotion.{{Cite journal |last1=Seckel |first1=Lauren |last2=Janis |first2=Christine |date=December 2008 |title=Convergences in Scapula Morphology among Small Cursorial Mammals: An Osteological Correlate for Locomotory Specialization |url=http://link.springer.com/10.1007/s10914-008-9085-7 |journal=Journal of Mammalian Evolution |language=en |volume=15 |issue=4 |pages=261–279 |doi=10.1007/s10914-008-9085-7 |issn=1064-7554|url-access=subscription }} The basal notungulate Notostylops and the mesotheriids are suggested to have engaged in digging, with mesotheriids suggested to have had an ecology similar to wombats. Toxodontids have sometimes been compared to rhinoceroses and hippopotamuses in overall bodyform and tooth morphology. The Miocene toxodontian Homalodotherium had claws on its forelimbs and is thought to have had an ecology similar to the extinct chalicotheres, rearing on its hindlegs to feed. Like perissodactyls, notoungulates were likely primitively hindgut fermenters,{{Cite journal |last1=Croft |first1=Darin A. |last2=Lorente |first2=Malena |date=2021-08-17 |editor-last=Smith |editor-first=Thierry |title=No evidence for parallel evolution of cursorial limb adaptations among Neogene South American native ungulates (SANUs) |journal=PLOS ONE |language=en |volume=16 |issue=8 |pages=e0256371 |doi=10.1371/journal.pone.0256371 |issn=1932-6203 |pmc=8370646 |pmid=34403434 |doi-access=free |bibcode=2021PLoSO..1656371C }} but it has also been proposed that some of them may have had fermentation more similar to ruminants based on their skeletal anatomy, though this is uncertain.

{{gallery|Thomashuxleya.jpg|Restoration of Thomashuxleya, an early toxodontian notoungulate from the Eocene (Lutetian) of Argentina|Hegetotherium.svg|Skull of Hegetotherium (Typotheria, Hegetotheriidae)|Mesotherium cristatum.png|Skull of Mesotherium (Typotheria, Mesotheriidae)|Miocochilius anomopodus - skeleton - Honda Group - Colombia.jpg|Skeleton of Miocochilius (Typotheria, Interatheriidae)|Nesodon imbricatus skeleton reconstruction.jpg|Skeleton of Nesodon (Toxodontia, Toxodontidae)|ProtypotheriumSinclair 1.jpg|Skeleton of Protypotherium (Typotheria, Interatheriidae)|Skeleton-of-Pachyrukhos-from-Sinclair-1909.png|Skeleton of Pachyrukhos (Typotheria, Hegetotheriidae)|Homalodotherium FMNH.jpg|Skeleton of Homalodotherium (Toxodontia, Homalodotheriidae)|width=200|height=180|Rhynchippus equinus.jpg

|Rhynchippus (Toxodontia, Notohippidae)|Scarrittia canquelensis 43.jpg|Scarrittia (Toxodontia, Leontiniidae)}}

The oldest notoungulates appeared during the Paleocene, probably originating from "condylarth" ancestors that had migrated from North America. Notoungulates and other South American native ungulates reached their apex of diversity during the Eocene and Oligocene. Notoungulate species diversity was stable during the Miocene, though 45% of the family diversity of the group became extinct during the interval, including Homalodotheriidae, Leontiniidae, and Interatheriidae. The diversity of the group declined during the Pliocene and Pleistocene, which is coeval in time with the Great American Interchange, which allowed ungulates and other mammals from North America to enter South America. This decline has historically been attributed to competition with the new North American arrivals, though earlier views had probably overstated the importance of this, with climatic change also likely being an important factor.{{Cite journal |last1=Seoane |first1=Federico D. |last2=Roig Juñent |first2=Sergio |last3=Cerdeño |first3=Esperanza |date=2017-01-02 |title=Phylogeny and paleobiogeography of Hegetotheriidae (Mammalia, Notoungulata) |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2017.1278547 |journal=Journal of Vertebrate Paleontology |language=en |volume=37 |issue=1 |pages=e1278547 |doi=10.1080/02724634.2017.1278547 |bibcode=2017JVPal..37E8547S |issn=0272-4634|hdl=11336/45231 |hdl-access=free }} As part of the Great American interchange, the toxodontid Mixotoxodon migrated into Central and North America, with its furthest northern record being in Texas.{{Cite journal |last1=Lundelius |first1=Ernest L. |last2=Bryant |first2=Vaughn M. |last3=Mandel |first3=Rolfe |last4=Thies |first4=Kenneth J. |last5=Thoms |first5=Alston |date=January 2013 |title=The first occurrence of a toxodont (Mammalia, Notoungulata) in the United States |url=http://www.tandfonline.com/doi/abs/10.1080/02724634.2012.711405 |journal=Journal of Vertebrate Paleontology |language=en |volume=33 |issue=1 |pages=229–232 |doi=10.1080/02724634.2012.711405 |bibcode=2013JVPal..33..229L |issn=0272-4634|hdl=1808/13587 |hdl-access=free }} The last hegetotheriids are known from the Early Pleistocene (with a supposed Middle Pleistocene record being considered questionable). The youngest known member of Typotheria, the mesotheriid Mesotherium, has its last records in the late Middle Pleistocene, around 220,000 years ago.{{Cite journal |last1=Fernández-Monescillo |first1=Marcos |last2=Martínez |first2=Gastón |last3=García López |first3=Daniel |last4=Frechen |first4=Manfred |last5=Romero-Lebrón |first5=Eugenia |last6=Krapovickas |first6=Jerónimo M. |last7=Haro |first7=J. Augusto |last8=Rodríguez |first8=Pablo E. |last9=Rouzaut |first9=Sabrina |last10=Tauber |first10=Adan A. |date=February 2023 |title=The last record of the last typotherid (Notoungulata, Mesotheriidae, Mesotherium cristatum) for the middle Pleistocene of the western Pampean region, Córdoba Province, Argentina, and its biostratigraphic implications |url=https://linkinghub.elsevier.com/retrieve/pii/S027737912200556X |journal=Quaternary Science Reviews |language=en |volume=301 |pages=107925 |doi=10.1016/j.quascirev.2022.107925|bibcode=2023QSRv..30107925F |url-access=subscription }} The last notoungulates, the toxodontids Toxodon, Mixotoxodon and Piauhytherium became extinct at the end of the Late Pleistocene around 12,000 years ago as part of the Late Pleistocene megafauna extinctions, along with most other large mammals in the Americas. The extinction coincides with the arrival of the first humans to the Americas and they are suggested to have been a causal factor in the extinction.

{{Portal|Paleontology|Prehistoric mammals}}

{{Reflist}}

{{Refbegin}}

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| last = Billet | first = Guillaume

| title = Phylogeny of the Notoungulata (Mammalia) based on cranial and dental characters

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| volume = 9 | issue = 4 | pages = 481–97

| doi = 10.1080/14772019.2010.528456 | bibcode = 2011JSPal...9..481B

| s2cid = 84159942

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| s2cid = 23315598

| url = https://biblio.ugent.be/publication/353125

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{{Refend}}

{{Commons category|Notoungulata}}

• {{Cite book

| last = Carroll | first = Robert Lynn

| title = Vertebrate Paleontology and Evolution

| year = 1988 | publisher = W.H. Freeman and Company | location = New York

| isbn = 9780716718222 | oclc = 14967288 }}

• {{Cite book

| last = McKenna | first = M.C.

| chapter = Toward a phylogenetic classification of the Mammalia

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• {{Cite book

| last1 = McKenna | first1 = Malcolm C.

| last2 = Bell | first2 = Susan K.

| title = Classification of Mammals Above the Species Level

| year = 1997 | publisher = Columbia University Press | location = New York

| isbn = 0231110138 | oclc = 37345734 }}

{{Meridiungulata|N.}}

{{Taxonbar|from=Q131322}}

{{Authority control}}

Category:Paleocene first appearances

Category:Eocene mammals

Category:Pleistocene extinctions

Category:Mammal orders