Tetramerorhinus

Tetramerorhinus Temporal range: Early Miocene-Middle Miocene (Colhuehuapian-Laventan) ~21–11.8 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Left lower jaw of Tetramerorhinus mixtum
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	†Litopterna
Family:	†Proterotheriidae
Subfamily:	†Proterotheriinae
Genus:	†Tetramerorhinus Ameghino 1894
Type species
†Tetramerorhinus fortis Ameghino, 1894
Species
T. cingulatum Ameghino 1891 T. fleaglei Kramarz & Bond 2005 T. mixtum Ameghino 1894 T. lucarius Ameghino 1894 T. prosistens Ameghino 1899
Synonyms[1]
Genus synonymy Tichodon Ameghino, 1894 T. cingulatum Proterotherium cingulatum Ameghino, 1891 Tetramerorhinus fortis Ameghino, 1891 Licaphrium cavum Zittel, 1893 Proterotherium principale Ameghino, 1894 Tichodon quadrilobus Ameghino, 1894 T. lucarius Diadiaphorus santaecrucis Lydekker, 1894 Licaphrium proclivum Ameghino, 1894 Proterotherium brachygnathum Ameghino, 1894 Proterotherium nitens Ameghino, 1894 Proterotherium perpolitum Ameghino, 1894 Proterotherium karaikense Ameghino, 1904 Proterotherium politum Ameghino, 1904 Proterotherium brachygnathus Scott, 1910 T. prosistens Proterotherium prosistens Ameghino, 1899 Diadiaphorus zamius Ameghino, 1899 Heptaconus obcallatus Ameghino, 1899 T. mixtum Proterotherium mixtum Ameghino, 1894 Proterotherium pyramidatum Ameghino, 1894 Proterotherium browni Scott, 1910 Proterotherium dodgei Scott, 1910

Tetramerorhinus is an extinct genus of proterotheriid litoptern that lived during the Early and Middle Miocene in what is now Argentina and Peru.^[2]

Description

This animal resembled a small horse, with slender, elongated legs. The body was approximately a meter long, and it is supposed to have weighed between 30 and 50 kilograms. The skull had a relatively elongated snout, with nasal bones longer than Diadiaphorus and with less developed upper incisors. The orbits were positioned near the middle of the skull. The molars had lower crowns than those of more derived proterotheriids, such as Proterotherium.^[3][4]

The endocranial cast of a specimen of Tetramerorhinus lucarius has been preserved, which allowed to reconstruct certain structures of the animal's brain. The brain was quite large relatively to the size of the skull, due to the development, especially in the anterior part, of the neocortex, and to the complexity of the cerebellum. The olfactory lobes were small and separated by a deep notch, while the cerebellum was elongated and separated from the two cerebral hemispheres by a deep and narrow depression. The rhinencephalon was highly developed, and the piriform lobes were in a much less lateral position than those of archaic ungulates such as Phenacodus, and presented a sort of circular depression of unknown significance. Numerous longitudinal grooves covered the surface of the neocortex. The cranial nerves were unusually arranged relatively to those of other ungulates, and the foramen ovale was posterior to the tentorium osseum.^[5]

Classification

The genus Tetramerorhinus was first described in 1894 by Florentino Ameghino, based on fossil remains found in Early Miocene deposits in Patagonia. Numerous species have been ascribed to this genus, such as Tetramerorhinus cingulatum, T. fleaglei, T. mixtum, T. lucarius, T. prosisttens.^[6][7]

Tetramerorhinus is a proterotheriid, a group of litopterns with characteristics similar to those of equids, although they were not closely related. Tetramerorhinus seems to have been a derived form, closely related to Anisolophus and Proterotherium.

Below is a phylogenetic tree of the Proterotheriidae, based on the work of McGrath et al. 2020.^[8]

References

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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