Xenungulata

Xenungulata Temporal range: Late Paleocene-Early Eocene (Riochican-Casamayoran) ~58.6–48.7 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Interpretation of Carodnia vieirai
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Infraclass:	Placentalia
Order:	†Xenungulata Paula Couto 1952
Family and genera[3]
†Carodniidae Paula Couto 1952[1] †Carodnia Simpson 1935 †Etayoa Villarroel 1987 †Rodcania Gelfo, García-López & Bergqvist 2020[2] †Notoetayoa Gelfo, López & Bond 2008

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 from deposits in Brazil, Argentina, Peru,^[4] 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.^[5]

Description

Xenungulates are characterized by bilophodont M^1–2 and M_1–2, similar to pyrotheres, and complex lophate third molars, similar to uintatheres. Though other relationships, to arctocyonids for example, have been suggested, no proofs thereof have been found. The foot bones of xenungulates were short and robust and their digits terminated in broad, flat, and unfissured hoof-like unguals, quite unlike any other meridiungulates. The discovery of Etayoa in Colombia^[6] made it clear that xenungulates are distinct from other groups: Etayoa lacks lophate molar talonid (in contrast to Carodnia) and, since no distinct lophodonty is present in basal pyrotheres, there is reason to assume that bilophodonty evolved separately in xenungulates and pyrotheres. Xenungulates also show some dental similarity to primitive astrapotheres.^[7]

Taxonomy

Cifelli 1983 grouped Carodnia with pyrotheres based on a similarity in astragalus morphology, but later concluded that this observation was incorrect.^[8]

Notoetayoa is most closely related to Etayoa.^[9]

Distribution

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Locations of Xenungulata fossils
Bogotá Formation
Peñas Coloradas Formation
Mogollón Formation
Itaboraí Formation

Xenungulata fossils have been found in:^[10]

• Bogotá Formation, Casamayoran, Colombia
• Peñas Coloradas Formation, Riochican, Argentina
• Mogollón Formation, Itaboraian-Riochican, Peru
• Itaboraí Formation, Itaboraian, Brazil

References

Bibliography

• v
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Genera of South American native ungulates

• Kingdom: Animalia
• Phylum: Chordata
• Class: Mammalia
• Superorder: Laurasiatheria

Xenungulata, Pyrotheria, Astrapotheria, and other minor groups
†Kollpaniidae? Andinodus Molinodus Pucanodus Simoclaenus Tiuclaenus †Didolodontidae Didolodus Enneoconus Ernestokokenia Escribania? Lamegoia Paulogervaisia Raulvaccia Ricardocifellia Saltaodus Umayodus †Xenungulata †Carodniidae Carodnia Etayoa Notoetayoa Rodcania †Pyrotheria †Colombitheriidae Colombitherium Proticia †Pyrotheriidae Baguatherium Berracotherium Carolozittelia Griphodon Propyrotherium Pyrotherium †Astrapotheria Eoastrapostylops †Trigonostylopidae Shecenia Tetragonostylops Trigonostylops †Astrapotheriidae Antarctodon Astrapodon Comahuetherium Liarthrus Maddenia Parastrapotherium †Albertogaudryinae Albertogaudrya Astraponotus †Astrapotheriinae Astrapothericulus Astrapotherium Scaglia †Uruguaytheriinae Granastrapotherium Hilarcotherium Uruguaytherium Xenastrapotherium

Litopterna
†Litopterna †Protolipternidae Asmithwoodwardia Miguelsoria Protolipterna †Amilnedwardsiidae Amilnedwardsia Ernestohaeckelia Rutimeyeria †Sparnotheriodontidae Notiolofos Phoradiadius Sparnotheriodon Victorlemoinea †Indaleciidae Adiantoides Indalecia †Notonychopidae Notonychops Requisia †Lopholipterna †Adianthidae Adianthus Proadiantus Proectocion? Proheptaconus Thadanius Tricoelodus †Macraucheniidae Polymorphis †Cramaucheniinae Caliphrium Coniopternium Cramauchenia Llullataruca Phoenixauchenia Pternoconius Theosodon †Macraucheniinae Cullinia Huayqueriana Macrauchenia Macraucheniopsis Micrauchenia Oxyodontherium Paranauchenia Promacrauchenia Scalabrinitherium Windhausenia Xenorhinotherium †Proterotheriidae Mesolicaphrium Neodolodus Olisanophus Promylophis Pseudobrachytherium †Anisolambdinae Anisolambda Eolicaphrium Guilielmofloweria Heteroglyphis Lambdaconops Paranisolambda Protheosodon †Proterotheriinae Anisolophus Brachytherium Diadiaphorus Diplasiotherium Eoauchenia Epecuenia Epitherium Lambdaconus Neobrachytherium Neolicaphrium Paramacrauchenia Picturotherium Prolicaphrium Proterotherium Tetramerorhinus Thoatheriopsis Thoatherium Uruguayodon Villarroelia †Megadolodinae Bounodus Megadolodus Uncertain placement Kyraodus Proacrodon Salladolodus Xesmodon Wainka

Notoungulata
incertae sedis Acoelodus Archaeogaia Carolodarwinia Perutherium †Notioprogonia Satshatemnus Seudenius †Henricosborniidae Henricosbornia Nanolophodon Othnielmarshia Peripantostylops Simpsonotus †Notostylopidae Boreastylops Chilestylops Edvardotrouessartia Homalostylops Notostylops Otrhonia Parastylops †Toxodontia †Homalodotheriidae Asmodeus Chasicotherium Homalodotherium Trigonolophodon †Isotemnidae Anisotemnus Coelostylodon Distylophorus Hedralophus Isotemnus Lophocoelus Pampatemnus Periphragnis Pleurocoelodon Pleurostylodon Rhyphodon Thomashuxleya †Eutoxodontia †Leontiniidae Anayatherium Ancylocoelus Colpodon Coquenia Elmerriggsia Gualta Henricofilholia Huilatherium Leontinia Martinmiguelia Purperia? Scarrittia Taubatherium Termastherium †Notohippidae Acoelohyrax Argyrohippus Deuterotherium? Edvardocopeia Eomorphippus Eurygenium Mendozahippus Moqueguahippus Morphippus Nesohippus Notohippus Pampahippus Pascualihippus Patagonhippus Perhippidion Plexotemnus Puelia Purperia? Rhynchippus Rosendo Stilhippus Teushentherium Trimerostephanos †Toxodontidae †Nesodontinae Adinotherium Nesodon Proadinotherium †Toxodontinae Abothrodon Andinotoxodon Calchaquitherium Ceratoxodon Charruatoxodon Dinotoxodon Eutomodus Falcontoxodon Gyrinodon Haplodontherium Hemixotodon Hoffstetterius Hyperoxotodon Mesenodon Mesotoxodon Minitoxodon Mixotoxodon Neoadinotherium Neotoxodon Nesodonopsis Nonotherium Ocnerotherium Palyeidodon Paratrigodon Pericotoxodon Piauhytherium Pisanodon Plesiotoxodon Posnanskytherium Prototrigodon Stenotephanos Stereotoxodon Toxodon Toxodontherium Trigodon Trigodonops Xotodon †Typotheria Campanorco Griphotherion †Archaeopithecidae Archaeopithecus Teratopithecus †Oldfieldthomasiidae Allalmeia Brachystephanus Camargomendesia Colbertia Dolichostylodon Itaboraitherium Kibenikhoria Maxschlosseria Oldfieldthomasia Paginula Suniodon Tsamnichoria Ultrapithecus Xenostephanus †Interatheriidae Antofagastia Ignigena Johnbell Munizia Punapithecus †Interatheriinae Archaeophylus Argyrohyrax Boleatherium Brucemacfaddenia Caenophilus Choichephilum Cochilius Eopachyrucos Federicoanaya Interatherium Juchuysillu Miocochilius Neoicochilus Patriarchus Proargyrohyrax Progaleopithecus Protypotherium Santiagorothia †Notopithecinae Antepithecus Guilielmoscottia Notopithecus Transpithecus †Mesotheriidae Fiandraia? Trachytherus †Mesotheriinae Altitypotherium Caraguatypotherium Eotypotherium Eutypotherium Hypsitherium Mesotherium Microtypotherium Plesiotypotherium Pseudotypotherium Rusconitherium Typotheriopsis †Archaeohyracidae Archaeohyrax Archaeotypotherium Bryanpattersonia Eohyrax Pascualhyrax Protarchaeohyrax Pseudhyrax Punahyrax †Hegetotheriidae †Hegetotheriinae Ethegotherium Hegetotheriopsis Hegetotherium Hemihegetotherium Prohegetotherium Pseudohegetotherium Sallatherium †Pachyrukhinae Medistylus Propachyrucos Prosotherium Tremacyllus †Pachyrukhini Pachyrukhos Paedotherium Tremacyllus