Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Cladus: Synapsida
Cladus: Eupelycosauria
Cladus: Sphenacodontia
Cladus: Sphenacodontoidea
Cladus: Therapsida
Cladus: Theriodontia
Cladus: Cynodontia
Cladus: Eucynodontia
Cladus: Probainognathia
Cladus: Prozostrodontia
Cladus: Mammaliaformes
Classis: Mammalia
Subclassis: Trechnotheria
Infraclassis: Zatheria
Supercohors: Theria
Cohors: Eutheria
Infraclassis: Placentalia
Cladus: Boreoeutheria
Superordo: Laurasiatheria
Cladus: †Meridiungulata
Ordo: †Litopterna
Classis: Mammalia
Cladus: †Meridiungulata
Ordo: †Litopterna
Familiae: †Adianthidae – †Macraucheniidae – †Notonychopidae – †Proterotheriidae – †Protolipternidae
Name
Litopterna Ameghino, 1889: XXIII
References
Primary references
Ameghino, F. 1889. Contribucion al conocimiento de los mamiferos fosiles de la República Argentina. Actas de la Academia nacional de ciencias de la República Argentina en Cordoba 6: 1–1027, 98 plates. BHL DOI: 10.5962/bhl.title.121288 Reference page.
Additional references
Bastos, A. C. F., & L. P. Bergqvist. 2007. A postura locomotora de Protolipterna ellipsodontoides Cifelli, 1983 (Mammalia: Litopterna: Protolipternidae) da Bacia de São José de Itaboraí, Rio de Janeiro (Paleoceno superior). Anuário do Instituto de Geociências 30(1): 58–66.
Cifelli, R. L. 1983. Eutherian tarsals from the Late Paleocene of Brazil. American Museum novitates 2761: 1–31.
Cifelli, R. L. 1991. A new adianthid litoptern (Mammalia) from the Miocene of Chile. Revista Chilena de Historia Natural 64: 119–125.
Dozo, M. T. & B. Vera. 2010. First skull and associated postcranial bones of Macraucheniidae (Mammalia, Litopterna) from the Deseadan Salma (late Oligocene) of Cabeza Blanca (Chubut, Argentina). Journal of vertebrate paleontology 30(6): 1818–1826.
Rose, K. D. 2009. The Beginning of the Age of Mammals. JHU Press.
Ubilla, M., D. Perea, M. Bond & A. Rinderknecht. 2011. The first cranial remains of the Pleistocene proterotheriid Neolicaphrium Frenguelli, 1921 (Mammalia, Litopterna): a comparative approach. Journal of vertebrate paleontology 31(1): 193–201.
Vernacular names
català: Litopterns
English: Litopterns
Esperanto: Litopternoj
español: Litopternos
italiano: Litopterni
日本語: 滑距目
polski: Litopterny
português: Litopternos
русский: Литоптерны
Litopterna (from Ancient Greek: λῑτή πτέρνα "smooth heel") 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.[1] It is divided into nine families, with Proterotheriidae and Macraucheniidae being the most diverse and last surviving families.[2]
Diversity
Skeleton of Theosodon (Macraucheniidae)
Historic life restoration of Thoatherium (Proterotheriidae)
The body forms of many litopterns, notably in the limb and skull structure, are broadly similar to those of living ungulates, unlike other South American native ungulate groups, which are often strongly divergent from living ungulates.[3] Paleocene and Eocene litopterns generally had small body masses, with Protolipterna (Protolipternidae) estimated to have had a body mass of 0.5–1.5 kilograms (1.1–3.3 lb), though the Eocene sparnotheriodontids were considerably larger, with estimated body masses of around 400 kilograms (880 lb). Most proterotheriids had body masses of around 15 to 80 kilograms (33 to 176 lb) while many macraucheniids had body masses of around 80–120 kilograms (180–260 lb). Some of the last macraucheniids like Macrauchenia were considerably larger, with body masses around a ton.[1] Adianthidae generally had small body masses, with members of the genus Adianthus estimated to weigh 7.4–20 kilograms (16–44 lb). Members of the proterotheriid subfamily Megadolodinae are noted for having bunodont (rounded cusp) molar teeth, which is largely unique to litopterns among South American native ungulates.[2][1] Litopterns of the mid-late Cenozoic had hinge-like limb joints and hooves similar to those of modern ungulates, with the weight being supported on three toes in macraucheniids and one in proterotheriids, with the protherotheriid Thoatherium developing greater toe reduction than that present in living horses.[1] Macraucheniids had long necks and limbs.[4]
Members of the macraucheniid subfamily Macraucheniinae saw the progressive migration of the nasal opening to the top of the skull,[5] which was often historically suggested to indicate the presence of a trunk, though other authors have suggested that a moose-like prehensile lip,[6] or a saiga-like nose to filter dust[7] are more likely.
(A) Theosodon, (B) Scalabrinitherium, (C) Macrauchenia, portraying how the nasal bones shifted backwards on the skull, with the nasal opening following suit.
Ecology
Litopterns were likely hindgut fermenters.[3] At least some macraucheniids like Macrauchenia are suggested to have been mixed feeders feeding on both browse and grass.[8] Sparnotheriodontids are suggested to have been browsers.[9] Some proterotheriids are suggested to have been browsers,[10] while some members proterotheriid subfamily Megadolodinae like Megadolodus have been suggested to have been omnivorous with at least part of their diet consisting of hard fruit.[2]
Evolutionary history
Litopterna, like other "South American native ungulates" is thought to have originated from groups of archaic "condylarths" that migrated from North America.[1] Sequencing of the collagen proteome and mitochondrial genome of Macrauchenia has revealed that litopterns are true ungulates, sharing a common ancestor with Notoungulata, and with their closest living relatives being Perissodactyla (the group containing living equines, rhinoceros and tapirs) as part of the clade Panperissodactyla, with the split from Perissodactyla being estimated at around 66 million years ago.[11][12] The relationship of Litopterna to other South American native ungulate groups is uncertain, though it may be closely related to the "condylarth" group Didolodontidae. The earliest litopterns appeared during the early Paleocene, around 63 million years ago.[1]
Aside from South America, sparnotheriodontids are also known from the Eocene aged La Meseta Formation in the Antarctic Peninsula, representing the only record of litopterns on the Antarctic continent. Litopterns declined during the Pliocene and Pleistocene, likely as a result of climatic change and competition with recently immigrated North American ungulates who arrived as part of the Great American interchange, following the connection of the previously isolated North and South America via the Isthmus of Panama.[1] Macrauchenia, Macraucheniopsis, Xenorhinotherium (Macraucheniidae) and Neolicaphrium (Proterotheriidae) were the last surviving genera of litopterns. All four became extinct at the end of the Late Pleistocene around 12,000 years ago as part of the Quaternary extinction event, along with most other large mammals in the Americas, co-inciding with the arrival of the first humans to the continent. It is possible that hunting had a causal role in their extinction.[13][14][15][1]
Classification
Order Litopterna[16]
Proacrodon
Family Protolipternidae
Asmithwoodwardia
Miguelsoria
Protolipterna
Family Indaleciidae
Adiantoides leali
Cramauchenia normalis
Theosodon patagonica
Macrauchenia
Xenorhinotherium bahiensis
Adiantoides
Indalecia
Family Sparnotheriodontidae
Phoradiadius
Notiolofos
Sparnotheriodon
Victorlemoinea
Family Amilnedwardsiidae
Amilnedwardsia
Ernestohaeckelia
Rutimeyeria
Family Notonychopidae
Notonychops
Requisia
Superfamily Macrauchenioidea
Family Adianthidae
Proectocion
Adianthinae
Adianthus
Proadiantus
Proheptaconus
Thadanius
Tricoelodus
Family Macraucheniidae
Llullataruca
Subfamily Cramaucheniinae
Coniopternium
Caliphrium
Cramauchenia
Phoenixauchenia
Polymorphis
Pternoconius
Theosodon
Subfamily Macraucheniinae
Cullinia
Huayqueriana
Macrauchenia
Macraucheniopsis
Oxyodontherium
Paranauchenia
Promacrauchenia
Scalabrinitherium
Windhausenia
Xenorhinotherium
Superfamily Proterotherioidea
Family Proterotheriidae
Anisolophus floweri
Diadiaphorus majusculus
Thoatherium crepidatum
Anisolambda
Anisolophus
Brachytherium
Diadiaphorus
Diplasiotherium
Eoauchenia
Eolicaphrium
Epecuenia
Epitherium
Guilielmofloweria
Heteroglyphis
Lambdaconus
Lambdaconops
Mesolicaphrium
Neobrachytherium
Neodolodus
Neolicaphrium
Olisanophus
Paramacrauchenia
Paranisolambda
Picturotherium
Prolicaphrium
Promylophis
Proterotherium
Protheosodon
Pseudobrachytherium
Tetramerorhinus
Thoatherium
Thoatheriopsis
Villarroelia
Uruguayodon
Wainka
Xesmodon
Megadolodinae
Bounodus
Megadolodus
References
Croft, Darin A.; Gelfo, Javier N.; López, Guillermo M. (2020-05-30). "Splendid Innovation: The Extinct South American Native Ungulates". Annual Review of Earth and Planetary Sciences. 48 (1): 259–290. Bibcode:2020AREPS..48..259C. doi:10.1146/annurev-earth-072619-060126. ISSN 0084-6597. S2CID 213737574.
Carrillo, Juan D.; Suarez, Catalina; Benites-Palomino, Aldo Marcelo; Vanegas, Andrés; Link, Andrés; Rincón, Aldo F.; Luque, Javier; Cooke, Siobhán B.; Tallman, Melissa; Billet, Guillaume (2023-08-31). "New remains of Neotropical bunodont litopterns and the systematics of Megadolodinae (Mammalia: Litopterna)". Geodiversitas. 45 (15). doi:10.5252/geodiversitas2023v45a15. ISSN 1280-9659. S2CID 261638835.
Croft, Darin A.; Lorente, Malena (2021-08-17). Smith, Thierry (ed.). "No evidence for parallel evolution of cursorial limb adaptations among Neogene South American native ungulates (SANUs)". PLOS ONE. 16 (8): e0256371. doi:10.1371/journal.pone.0256371. ISSN 1932-6203. PMC 8370646. PMID 34403434.
Schmidt, Gabriela I.; Ferrero, Brenda S. (2014-07-29). "Taxonomic reinterpretation of Theosodon hystatus Cabrera and Kraglievich, 1931 (Litopterna, Macraucheniidae ) and phylogenetic relationships of the family". Journal of Vertebrate Paleontology. 34 (5): 1231–1238. Bibcode:2014JVPal..34.1231S. doi:10.1080/02724634.2014.837393. hdl:11336/18953. ISSN 0272-4634. S2CID 86091386.
Forasiepi, Analía M.; MacPhee, Ross D. E.; Del Pino, Santiago Hernández; Schmidt, Gabriela I.; Amson, Eli; Grohé, Camille (2016-06-22). "Exceptional Skull of Huayqueriana (Mammalia, Litopterna, Macraucheniidae) From the Late Miocene of Argentina: Anatomy, Systematics, and Paleobiological Implications". Bulletin of the American Museum of Natural History. 404: 1–76. doi:10.1206/0003-0090-404.1.1. ISSN 0003-0090. S2CID 89219979.
Moyano, Silvana Rocio; Giannini, Norberto Pedro (November 2018). "Cranial characters associated with the proboscis postnatal-development in Tapirus (Perissodactyla: Tapiridae) and comparisons with other extant and fossil hoofed mammals". Zoologischer Anzeiger. 277: 143–147. doi:10.1016/j.jcz.2018.08.005. hdl:11336/86349. S2CID 92143497.
Blanco, R. Ernesto; Jones, Washington W.; Yorio, Lara; Rinderknecht, Andrés (October 2021). "Macrauchenia patachonica Owen, 1838: Limb bones morphology, locomotory biomechanics, and paleobiological inferences". Geobios. 68: 61–70. Bibcode:2021Geobi..68...61B. doi:10.1016/j.geobios.2021.04.006.
de Oliveira, Karoliny; Asevedo, Lidiane; Calegari, Marcia R.; Gelfo, Javier N.; Mothé, Dimila; Avilla, Leonardo (August 2021). "From oral pathology to feeding ecology: The first dental calculus paleodiet study of a South American native megamammal". Journal of South American Earth Sciences. 109: 103281. Bibcode:2021JSAES.10903281D. doi:10.1016/j.jsames.2021.103281. S2CID 233678648.
Bond, M.; Reguero, M. A.; Vizcaíno, S. F.; Marenssi, S. A. (January 2006). "A new 'South American ungulate' (Mammalia: Litopterna) from the Eocene of the Antarctic Peninsula". Geological Society, London, Special Publications. 258 (1): 163–176. doi:10.1144/GSL.SP.2006.258.01.12. ISSN 0305-8719.
Morosi, Elizabeth; Ubilla, Martin (2019-02-07). "Dietary and palaeoenvironmental inferences in Neolicaphrium recens Frenguelli, 1921 (Litopterna, Proterotheriidae) using carbon and oxygen stable isotopes (Late Pleistocene; Uruguay)". Historical Biology. 31 (2): 196–202. doi:10.1080/08912963.2017.1355914. ISSN 0891-2963. S2CID 133673644.
Welker, Frido; Collins, Matthew J.; Thomas, Jessica A.; Wadsley, Marc; Brace, Selina; Cappellini, Enrico; Turvey, Samuel T.; Reguero, Marcelo; Gelfo, Javier N. (March 18, 2015). "Ancient proteins resolve the evolutionary history of Darwin/'s South American ungulates". Nature. 522 (7554): 81–84. Bibcode:2015Natur.522...81W. doi:10.1038/nature14249. hdl:11336/14769. ISSN 1476-4687. PMID 25799987. S2CID 4467386.
Westbury, Michael; Baleka, Sina; Barlow, Axel; Hartmann, Stefanie; Paijmans, Johanna L.A.; Kramarz, Alejandro; Forasiepi, Analía M; Bond, Mariano; Gelfo, Javier N.; Reguero, Marcelo A.; López-Mendoza, Patricio; Taglioretti, Matias; Scaglia, Fernando; Rinderknecht, Andrés; Jones, Washington (2017-06-27). "A mitogenomic timetree for Darwin's enigmatic South American mammal Macrauchenia patachonica". Nature Communications. 8 (1): 15951. Bibcode:2017NatCo...815951W. doi:10.1038/ncomms15951. ISSN 2041-1723. PMC 5490259. PMID 28654082.
Webb, S. D. (1976). "Mammalian Faunal Dynamics of the Great American Interchange". Paleobiology. 2 (3): 220–234. Bibcode:1976Pbio....2..220W. doi:10.1017/S0094837300004802. JSTOR 2400220. S2CID 251050063.
Marshall, L. G.; Cifelli, R. L. (1990). "Analysis of changing diversity patterns in Cenozoic land mammal age faunas, South America". Palaeovertebrata. 19: 169–210. Retrieved 2018-10-08.
Webb, S. D. (1991). "Ecogeography and the Great American Interchange". Paleobiology. 17 (3): 266–280. Bibcode:1991Pbio...17..266W. doi:10.1017/S0094837300010605. JSTOR 2400869. S2CID 88305955.
"PBDB". paleobiodb.org. Retrieved 2022-04-29.
Further reading
McKenna, Malcolm C; Bell, Susane K (1997). Classification of Mammals Above the Species Level. New York: Columbia University Press. ISBN 978-0-231-11013-6.
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