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Mixotoxodon Temporal range: Mid-Late Pleistocene (Ensenadan-Lujanian & Irvingtonian-Rancholabrean) ~1.8–0.012 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Life restoration of Mixotoxodon larensis
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	†Notoungulata
Family:	†Toxodontidae
Subfamily:	†Toxodontinae
Genus:	†Mixotoxodon Van Frank 1957
Species:	†M. larensis
Binomial name
†Mixotoxodon larensis Van Frank 1957
Some locations where Mixotoxodon fossils have been found

Mixotoxodon ("mixture Toxodon") is an extinct genus of notoungulate of the family Toxodontidae inhabiting South America, Central America and parts of southern North America during the Pleistocene epoch, from 1,800,000—12,000 years ago.

Description

Mixotoxodon is known by fragmentary remains, usually mandible fragments and teeth. Although the general appearance probably was very similar to another toxodontid from the Pleistocene, the better known Toxodon, their fossils shown that the outer borders of the symphysis in the lower jaw don't diverge anteriorly, and the incisors form a semicircular structure that protrude less than the incisors of Toxodon; the snout was cylindrical, instead of the broad hippo-like muzzle of Toxodon. The straight snout and the narrow lower incisors closely packed, suggest that this animal had a different feeding strategy compared to their southern relative, although the teeth of both genera were adapted to deal with abrasive food. It was a rhino-sized animal, a 2012 study estimated a weight of up to 3.8 tonnes (4.2 short tons), which makes it the largest member of Notoungulata, though this may be an overestimate.

Mixotoxodon is known from a single species, M. larensis. Mixotoxodon is the only notoungulate known to have migrated out of South America during the Great American Interchange. Its fossils have been found in northern South America, in Central America, in Veracruz and Michoacán, Mexico (with a possible find in Tamaulipas), and eastern Texas, USA. The genus was also one of the last surviving notoungulates, along with related genera such as the better-known Toxodon. The name refers to the fact that Mixotoxodon combines characteristics typical of different toxodontid subfamilies.

Phylogeny

The cladogram below is based in the study published by Analía Forasiepi and colleagues (2014), showing the position of Mixotoxodon inside Toxodontidae:

Fossil distribution

This list indicates the countries and places where Mixotoxodon fossils have been found. The list follows Rincón, 2011, unless otherwise indicated:

North America

• United States
  • Harris County, Texas
• Mexico
  • Hihuitlán, Michoacán
  • La Estribera, Veracruz (Polaco et al., 2004)
• Guatemala
  • Santa Amelia River, Petén Department (Woodburne, 1969)
• Honduras
  • Yeroconte, Lempira department (Webb and Perrigo, 1984)
  • Orillas del Humuya, Comayagua department
• El Salvador
  • Tomayate, San Salvador department (Cisneros, 2005)
  • Cuscatlán Formation, Barranca del Sisimico, San Vicente department
  • Hormiguero, San Miguel department (Webb and Perrigo, 1984)
• Nicaragua
  • El Bosque, Estelí department (Leidy, 1886)
• Costa Rica
  • Bajo de los Barrantes, Alajuela province (Laurito, 1993; Valerio, 1939; Spencer et al., 1997)
• Panama
  • Ocú, Herrera Province (Gazin, 1956)

South America

• Colombia
  • Rotinet Formation, Chívolo, Magdalena (De Porta 1959; Villarroel and Clavijo, 2005)
• Venezuela
  • Mene de Inciarte, Zulia (Rincón, 2011)
  • Quebrada Ocando, Falcón State (Bocquentin-Villanueva, 1984)
  • Muaco, Falcón State (Royo y Gómez, 1960, Bocquentin-Villanueva, 1979)
  • Cerro Misión, Falcón State (Rincón, 2004)
  • Zumbador Cave, Falcón State
  • Agua Viva del Totumo, Lara State (Karsten, 1886)
  • San Miguel, Lara Sate (Van Frank, 1957)
  • El Tocuyo, Lara State
  • El Breal de Orocual, Monagas State (Rincón et al., 2009)
• Brazil
  • Juruá River, Acre State (Paula Couto, 1982; Rancy, 1981)
  • Araras/Periquitos, Rondônia
• Bolivia
  • Cara Cara, Beni Department (Hoffstetter, 1968)
• Argentina
  • Dique Los Quiroga, Santiago del Estero

References

1. Mixotoxodon at Fossilworks.org
2. "NEW REMAINS OF Mixotoxodon larensis Van Frank 1957 (Mammalia: Notoungulata) FROM MENE DE INCIARTE TAR PIT, NORTH-WESTERN VENEZUELA Ascanio D. Rincó" (PDF).
3. Paula-Couto, 1979, p. 404.
4. Elissamburu, 2012, p. 108.
5. Nelson, Allison; Engelman, Russell K.; Croft, Darin A. (September 2023). "How to weigh a fossil mammal? South American notoungulates as a case study for estimating body mass in extinct clades". Journal of Mammalian Evolution. 30 (3): 773–809. doi:10.1007/s10914-023-09669-1. ISSN 1064-7554.
6. McKenna & Bell, 1997, p. 461; Cisneros, 2005, p. 246.
7. "NEW REMAINS OF Mixotoxodon larensis Van Frank 1957 (Mammalia: Notoungulata) FROM MENE DE INCIARTE TAR PIT, NORTH-WESTERN VENEZUELA Ascanio D. Rincó" (PDF).
8. Arroyo-Cabrales et al., 2010, pp. 193-194
9. "A New Occurrence of Toxodonts in the Pleistocene of México". ResearchGate. Retrieved 2016-01-23.
10. Lucas, Spencer G.; Morgan, Gary S.; Spielmann, Justin A.; Prothero, Donald R. (2008-01-01). Neogene Mammals: Bulletin 44. New Mexico Museum of Natural History and Science.
11. Lundelius et al., p. 229.
12. van Frank, 1957, p. 6.
13. Forasiepi, A. A. M.; Cerdeño, E.; Bond, M.; Schmidt, G. I.; Naipauer, M.; Straehl, F. R.; Martinelli, A. N. G.; Garrido, A. C.; Schmitz, M. D.; Crowley, J. L. (2014). "New toxodontid (Notoungulata) from the Early Miocene of Mendoza, Argentina". Paläontologische Zeitschrift. 89 (3): 611–634. doi:10.1007/s12542-014-0233-5. hdl:11336/20443. S2CID 129293436.
14. Rincón 2011, p. 896.
15. Lundelius et al., 2013, p.229
16. Chimento & Agnolin, 2011, p. 90

Bibliography and further reading

• Arroyo-Cabrales, Joaquín; Polaco, Oscar J.; Johnson, Eileen; Ferrusquía-Villafranca, Ismael (2010-02-01). "A perspective on mammal biodiversity and zoogeography in the Late Pleistocene of México". Quaternary International. 212 (2). Elsevier: 187–197. Bibcode:2010QuInt.212..187A. doi:10.1016/j.quaint.2009.05.012.
• Chimento, Nicolás R., and Federico L. Agnolin. Mamíferos del Pleistoceno Superior de Santiago del Estero (Argentina) y sus afinidades paleobiogeográficas. Papéis Avulsos de Zoologia 51.6 (2011): 83-100.
• Cisneros, J.C. 2005. New Pleistocene vertebrate fauna from El Salvador. Revista Brasileira de Paleontologia, 8(3):239-255.
• De Porta, Jaime., 1959: Nueva subespecie de Toxodóntido del Cuaternario de Colombia.- Boletín de Geolología, Universidad Industrial de Santander, 3: 55-61.
• Elissamburu A., 2012. Estimación de la masa corporal en géneros del Orden Notoungulata. Estudios Geológicos, Vol 68, No 1, doi:10.3989/egeol.40336.133
• Laurito, César Alberto. Análisis topológico y sistemático del Toxodonte de Bajo de los Barrantes, provincia de Alajuela, Costa Rica. Revista Geológica de América Central 16 (1993).
• Lucas, Spencer G., Guillermo E. Alvarado, and Eduardo Vega. The pleistocene mammals of Costa Rica. Journal of Vertebrate Paleontology 17.2 (1997): 413-427.
• Lundelius, E. L.; Bryant, V. M.; Mandel, R.; Thies, K. J.; Thoms, A. (January 2013). "The first occurrence of a toxodont (Mammalia, Notoungulata) in the United States". Journal of Vertebrate Paleontology. 33 (1): 229–232. Bibcode:2013JVPal..33..229L. doi:10.1080/02724634.2012.711405. hdl:1808/13587. S2CID 53601518.
• MacFadden, Bruce J. (2005). Diet and habitat of toxodont megaherbivores (Mammalia, Notoungulata) from the late Quaternary of South and Central America. Quaternary Research 64 (2005) 113 – 124.
• McKenna, Malcolm C., and Bell, Susan K. 1997. Classification of Mammals Above the Species Level. Columbia University Press, New York, 631 pp. ISBN 0-231-11013-8
• Paula-Couto, C. (1979). Capítulo XXI, Ordem Notoungulata Roth, 1903. In Tratado de Paleomastozoologia. Academia Brasileira de Ciências, 590 p, Rio de Janeiro.
• Rincón, Ascanio D. Los mamíferos fósiles del Pleistoceno de la cueva del Zumbador (fa. 116), Estado Falcón, Venezuela. Boletín de la Sociedad Venezolana de Espeleología 37 (2003): 18-26.
• Rincón, Ascanio D. New remains of Mixotoxodon larensis Van Frank 1957 (Mammalia: Notoungulata) from Mene de Inciarte tar pit, north-western Venezuela. Interciencia 36.12 (2011): 894-899.
• Van Frank, R. 1957. A fossil collection from northern Venezuela. 1, Toxodontidae (Mammalia, Notoungulata). American Museum Novitates, 1850:1-38.
• Villarroel A., C. & J. Clavijo ( 2005). Los mamíferos fósiles y las edades de las sedimentitas continentales del Neógeno de la Costa Caribe Colombiana. Revista de la Academia Colombiana de Ciencias 29 (112): 345-356. ISSN 0370-3908.

v t e Genera of South American native ungulates
Kingdom: Animalia Phylum: Chordata Class: Mammalia Superorder: Laurasiatheria
Xenungulata, Pyrotheria, Astrapotheria, and other minor groups †Kollpaniidae? Andinodus Molinodus Pascualodus? Pucanodus Simoclaenus Tiuclaenus †Didolodontidae Didolodus Enneoconus Ernestokokenia Escribania? Lamegoia Paulogervaisia Raulvaccia Ricardocifellia Saltaodus Umayodus Xesmodon †Protolipternidae Asmithwoodwardia Miguelsoria Protolipterna †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 †Eulitopterna †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 †Proterotherioidea †Anisolambdidae †Anisolambdinae Anisolambda Eolicaphrium Guilielmofloweria Lambdaconops Paranisolambda Polyacrodon Protheosodon Wainka †Sparnotheriodontinae Heteroglyphis Notiolofos Phoradiadius Sparnotheriodon Victorlemoinea †Proterotheriidae Mesolicaphrium Neodolodus Olisanophus Promylophis Prothoatherium Pseudobrachytherium Salladolodus †Megadolodinae Bounodus Megadolodus †Proterotheriinae Anisolophus Brachytherium Diadiaphorus Diplasiotherium Eoauchenia Epecuenia Epitherium Lambdaconus Neobrachytherium Neolicaphrium Paramacrauchenia Picturotherium Prolicaphrium Proterotherium Tetramerorhinus Thoatheriopsis Thoatherium Uruguayodon Villarroelia †Notopterna †Amilnedwardsiidae Amilnedwardsia Ernestohaeckelia Rutimeyeria †Indaleciidae Adiantoides Indalecia †Notonychopidae Notonychops Requisia Incertae sedis Kyraodus Proacrodon
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

• Toxodonts
• Pleistocene first appearances
• Holocene extinctions
• Pleistocene mammals of North America
• Irvingtonian
• Pleistocene Costa Rica
• Pleistocene El Salvador
• Pleistocene Guatemala
• Pleistocene Honduras
• Pleistocene Mexico
• Pleistocene Nicaragua
• Pleistocene Panama
• Pleistocene United States
• Pleistocene mammals of South America
• Uquian
• Ensenadan
• Lujanian
• Pleistocene Argentina
• Pleistocene Bolivia
• Pleistocene Brazil
• Pleistocene Colombia
• Pleistocene Venezuela
• Fossils of Costa Rica
• Fossils of El Salvador
• Fossils of Guatemala
• Fossils of Honduras
• Fossils of Mexico
• Fossils of Nicaragua
• Fossils of Panama
• Fossils of Argentina
• Fossils of Bolivia
• Fossils of Brazil
• Fossils of Colombia
• Fossils of Venezuela
• Fossil taxa described in 1957
• Prehistoric placental genera