Theosodon

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Theosodon
Temporal range: Early-Mid Miocene (Colhuehuapian-Laventan)
21.0–11.8  Ma
Theosodon patagonicum 2.jpg
Theosodon patagonicum skull
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Litopterna
Family: Macraucheniidae
Subfamily: Cramaucheniinae
Genus: Theosodon
Ameghino, 1887
Type species
Theosodon lydekkeri
Ameghino, 1887
Species
  • T. arozquetaiMcGrath et al., 2018
  • T. fontanaeAmeghino, 1891
  • T. garretorumScott, 1910
  • T. gracilisAmeghino, 1891
  • T. karaikensisAmeghino, 1904
  • T. lallemantiMercerat, 1891
  • T. lydekkeriAmeghino, 1887
  • T. patagonicum(Ameghino, 1891)
Synonyms
  • Pseudocoelosoma [1]
    Ameghino, 1891

Theosodon is an extinct genus of litoptern mammal from the Early to Middle Miocene of South America.

Contents

Description

Reconstruction Theosodon NT.jpg
Reconstruction

Theosodon was long-legged with a long neck resembling modern llamas or guanacos. It was large for a litoptern, reaching up to 2 metres (6.6 ft) in length and weighing up to 125–170 kilograms (276–375 lb). [2]

It had a long neck and tapir-like, three-toed feet, [2] [3] and like other litopterns and modern horses, tapirs and rhinos, it bore its weight on its middle toes. [2]

Extraordinarily, rather than having nostrils at the front of its head, Theosodon had its nostrils on the top of its snout, halfway between the forehead and the tip of the snout, and its nostrils pointed upwards rather than forwards, possibly as an adaptation for browsing on prickly vegetation. [2]

History and species

Theosodon has been known since the 19th century, and by 1910 seven species had been described within the genus, all from the early Miocene Santa Cruz Formation in Argentina. [4] The name Theosodon means "god tooth". The first fossils were only dental remains, but it is unknown why this name was given. [2] Though seven species had originally been described, the exact number of discovered species remains uncertain as it varies between different authors. There are either ten or seven species, and in 2014 Schmidt and Ferrero put forward that the genus needed a full taxonomic revision due to this issue. [4]

"Theosodon" hystatus

In 1931, fossils found in the late Miocene Arroyo Chasicó Formation were assigned to the genus under the name Theosodon hystatus. This was questioned in 1995, and in 2014 it was moved to the genus Paranauchenia due to similarities to the species Paranauchenia denticulata. This move was supported by phylogenetic analysis that showed the two species formed monophyletic clade. [4]

"Theosodon" arozquetai

In 2018 a new species of macraucheniid litoptern was described based on the partial remains of a skull and two metapodials. A phylogenetic analysis tentatively linked the species to Theosodon as a sister clade and it was named Theosodon arozquetai pending revision. Its body mass was estimated to be somewhere from 80.8 to 116.2 kilograms (178 to 256 lb). It dated to the middle Miocene, about 13 million years ago, somewhat later than most other Theosodon species. [5]

Classification

Reconstructed skeleton Reports of the Princeton University Expeditions to Patagonia, 1896-1899. J. B. Hatcher in charge (1901) (14749571352).jpg
Reconstructed skeleton

Theosodon is in the subfamily Cramaucheniinae within the family Macraucheniidae. A phylogenetic analysis of the family in 2014 found that Cramaucheniinae is a paraphyletic group, and that Theosodon is a sister clade to the subfamily Macraucheniinae, the least primitive of the members of Cramaucheniinae. [4]

Proadiantus

Tricoelodus

Polymorphis

Pternoconius

Coniopternium

Cramauchenia

Theosodon

Scalabrinitherium

Oxyodontherium

Huayqueriana

Paranauchenia denticulata

Paranauchenia hystatus

Cullinia

Promacrauchenia

Windhausenia

Xenorhinotherium

Macraucheniopsis

Macrauchenia

Cramaucheniinae
Macraucheniinae

Paleobiology

Restoration of Theosodon garretorum and Borhyaena tuberata Theosodon.jpg
Restoration of Theosodon garretorum and Borhyaena tuberata

Theosodon was a terrestrial and cursorial animal that may have lived in both forests and more open environments. [2] Due to its size and its long neck, Theosodon was likely a high browser, stripping leaves off of trees and shrubs high off the ground. [2] Theosodon had a slender jaw compared to many other litopterns, indicating its food was softer, [6] such as dicotyledons. [7] As its nostrils are upward-facing and at the centre of its head rather than the front, Theosodon may have used this adaptation to feed on thorny plants. Some modern animals such as giraffes feed on thorny plants, and their nostrils are also further back and face more upwards than forwards. [2]

Other fossils were found in the Sarmiento Formation of Argentina, [8] [9] the Yecua Formation of Bolivia, the Honda Group of Colombia, the Chucal and Río Frías Formations of Chile and the Ipururo Formation of Peru. [10]

Related Research Articles

<span class="mw-page-title-main">Litopterna</span> Extinct order of hoofed mammals

Litopterna 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. It is divided into nine families, with Proterotheriidae and Macraucheniidae being the most diverse and last surviving families.

<i>Macrauchenia</i> Extinct genus of litopterns

Macrauchenia was a large, long-necked and long-limbed, three-toed native South American ungulate in the order Litopterna. The genus gives its name to its family, the Macraucheniidae or "robust litopterns". Like other litopterns, it is most closely related to the odd-toed ungulates (Perissodactyla), from which litopterns diverged approximately 66 million years ago. The oldest fossils in the genus date to the late Miocene, around seven million years ago, and M. patachonica disappears from the fossil record during the late Pleistocene, around 20,000-10,000 years ago. M. patachonica is one of the last and best known member of the family and is known primarily from the Luján Formation in Argentina, but is known from localities across southern South America. Another genus of macraucheniid Xenorhinotherium was present in northeast Brazil and Venezuela during the Late Pleistocene. The type specimen was discovered by Charles Darwin during the voyage of the Beagle. In life, Macrauchenia may have resembled a humpless camel, though the two taxa are not closely related. It fed on plants in a variety of environments across what is now South America. Among the species described, M. patachonica and M. ullomensis are considered valid; M. boliviensis is considered a nomen dubium; and M. antiqua has been moved to the genus Promacrauchenia.

<span class="mw-page-title-main">Macraucheniidae</span> Family in the extinct South American ungulate order Litopterna, that resembled various camelids

Macraucheniidae is a family in the extinct South American ungulate order Litopterna, that resembled various camelids. The reduced nasal bones of their skulls was originally suggested to have housed a small proboscis, similar to that of the saiga antelope. However, one study suggested that they were openings for large moose-like nostrils. Conversely, prehistoric pictographs by indigenous people seems to depict animals interpreted as macraucheniids with trunks. Their hooves were similar to those of rhinoceroses today, with a simple ankle joint and three digits on each foot. Thus, they may have been capable of rapid directional change when running away from predators, such as large phorusrhacid terror birds, sparassodont metatherians, giant short-faced bears (Arctotherium) and saber-toothed cats (Smilodon). Macraucheniids probably lived in large herds to gain protection against these predators, as well as to facilitate finding mates for reproduction.

<i>Protypotherium</i> Extinct genus of notoungulates

Protypotherium is an extinct genus of notoungulate mammals native to South America during the Oligocene and Miocene epochs. A number of closely related animals date back further, to the Eocene. Fossils of Protypotherium have been found in the Deseadan Fray Bentos Formation of Uruguay, Muyu Huasi and Nazareno Formations of Bolivia, Cura-Mallín and Río Frías Formations of Chile, and Santa Cruz, Salicas, Ituzaingó, Aisol, Cerro Azul, Cerro Bandera, Cerro Boleadoras, Chichinales, Sarmiento and Collón Curá Formations of Argentina.

Paranauchenia is an extinct genus of South American litopterns belonging to the family Macraucheniidae. It is known only from fossil finds in Argentina. It possessed three toes and long limbs. The species Paranauchenia denticulata lived in the Miocene epoch in Argentina. Fossils have been found in the Arroyo Chasicó and Ituzaingó Formations of Argentina.

Huayqueriana is an extinct genus of South American litoptern, related to Macrauchenia, and belonging to the same family, Macraucheniidae. It was formerly known as Macrauchenidia latidens, described in 1939 by Cabrera, but redefined as Huayqueriana in 2016 based on the earlier name convention of Rovereto 1914. The genus is named after the Huayquerías Formation and the eponymous Huayquerian South American land mammal age defined at the formation.

<i>Xenorhinotherium</i> Extinct genus of litopterns

Xenorhinotherium is an extinct genus of macraucheniine macraucheniids, closely related to Macrauchenia of Patagonia. The type species is X. bahiense.

<i>Cramauchenia</i> Extinct genus of litoptern South American ungulate

Cramauchenia is an extinct genus of litoptern South American ungulate. Cramauchenia was named by Florentino Ameghino. The name has no literal translation. Instead, it is an anagram of the name of a related genus Macrauchenia. This genus was initially discovered in the Sarmiento Formation in the Chubut Province, in Argentina, and later it was found in the Chichinales Formation in the Río Negro Province and the Cerro Bandera Formation in Neuquén, also in Argentina, in sediments assigned to the SALMA Colhuehuapian, as well as the Agua de la Piedra Formation in Mendoza, in sediments dated to the Deseadan. In 1981 Soria made C. insolita a junior synonym of C. normalis. A specimen of C. normalis was described in 2010 from Cabeza Blanca in the Sarmiento Formation, in sediments assigned to the Deseadan SALMA.

Victorlemoinea is an extinct litoptern genus of the family Sparnotheriodontidae, that lived from the Early to Middle Eocene. Fossils of Victorlemoinea have been found in the Las Flores, Sarmiento and Koluel Kaike Formations of Argentina, the Itaboraí Formation of Brazil and La Meseta Formation, Antarctica.

<i>Scalabrinitherium</i> Extinct genus of litopterns

Scalabrinitherium is an extinct genus of mammals of the family Macraucheniidae. Fossils of this animal were found among the fossils of prehistoric xenarthrans in the Ituzaingó Formation of Argentina.

<span class="mw-page-title-main">Proterotheriidae</span> Extinct family of litopterns

Proterotheriidae is an extinct family of litoptern ungulates known from the Oligocene-Late Pleistocene of South America. Members of the group were small-medium sized cursorial herbivores with brachydont teeth, with their toes showing progressive reduction, with later members of the group bearing weight on a single large toe similar to living horses.

Cullinia is an extinct genus of litoptern, an order of South American native ungulates that included horse-like and camel-like animals such as Macrauchenia. It is only known from fragmentary remains. Cullinia levis is known from Chasicoan remains found in the Arroyo Chasicó Formation of Argentina, and remains from the Brazilian state of Acre and the Huayquerian Ituzaingó Formation have been assigned to Cullinia sp..

<span class="mw-page-title-main">Cramaucheniinae</span> Extinct subfamily of mammals

Cramaucheniinae is a paraphyletic subfamily of macraucheniids that originated in the middle Eocene. The size range of the group ranged from small, basal forms to larger and more derived forms. During their evolution, the cramaucheniines undergone a trend from evolving from small basal forms such as Polymorphis into larger, more derived taxa such as Theosodon.

Proterotherium is an extinct genus of litoptern mammal of the family Proterotheriidae that lived during the Late Miocene of Argentina and Chile. Fossils of this genus have been found in the Ituzaingó Formation of Argentina, and the Galera Formation of Chile.

Promacrauchenia is an extinct genus of macraucheniids that lived during the Late Miocene to Late Pliocene epochs of what is now Argentina and Bolivia. It belongs to the subfamily Macraucheniinae, which also includes Huayqueriana, Macrauchenia, and Xenorhinotherium. Fossils of this genus have been found in the Ituzaingó, Andalhuala, and Cerro Azul Formations of Argentina.

Llullataruca is an extinct genus of macraucheniid litoptern. It lived during the Middle Miocene of what is now Bolivia.

Polymorphis is an extinct genus of litopterns belonging to the family Macraucheniidae. It lived during the Middle Eocene of Argentina.

<i>Tetramerorhinus</i> Extinct genus of litopterns

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

Pternoconius is an extinct genus of macraucheniid litoptern from the Late Oligocene and Early Miocene of Argentina. Fossils of this genus have been found in the Sarmiento Formation of Argentina.

<span class="mw-page-title-main">Santa Cruz Formation</span> Geological formation in Patagonia

The Santa Cruz Formation is a geological formation in the Magallanes/Austral Basin in southern Patagonia in Argentina and in adjacent areas of Chile. It dates to the late Early Miocene epoch, and is contemporaneous with eponymous Santacrucian SALMA. The formation extends from the Andes to the Atlantic coast. In its coastal section it is divided into two members, the lower, fossil rich Estancia La Costa Member, which has a lithology predominantly consisting of tuffaceous deposits and fine grained sedimentary claystone and mudstone, and the upper fossil-poor Estancia La Angelina Member, which consists of sedimentary rock, primarily claystone, mudstone, and sandstone. The environment of deposition is interpreted to have been mostly fluvial, with the lowermost part of the Estancia La Costa Member being transitional between fluvial and marine conditions. The environment of the Estancia La Costa Member is thought to have been relatively warm and humid, but likely became somewhat cooler and drier towards the end of the sequence. The Santa Cruz Formation is known for its abundance of South American native ungulates, as well as an abundance of rodents, xenarthrans, and metatherians.

References

  1. Scott, William B. (1910). The Litopterna of the Santa Cruz Beds. Reports of the Princeton University Expeditions to Patagonia, 1896–1899. Princeton: Princeton University Press.
  2. 1 2 3 4 5 6 7 8 Croft, Darin A. (18 August 2016). "Chucal, Chile". Horned Armadillos and Rafting Monkeys: The Fascinating Fossil Mammals of South America. Indiana University Press. pp. 104–117. ISBN   978-0-253-02084-0.
  3. Palmer, D., ed. (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals. London: Marshall Editions. p. 247. ISBN   1-84028-152-9.
  4. 1 2 3 4 Schmidt, Gabriela I.; Ferrero, Brenda S. (September 2014). "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 .
  5. Fonseca, Lurdes (31 May 2018). "Just out | Two new macraucheniids (Mammalia: Litopterna) from the late middle Miocene (Laventan South American Land Mammal Age) of Quebrada Honda, Bolivia @ Journal of Vertebrate Paleontology | Paleowire" . Retrieved 6 June 2018.
  6. Cassini, Guillermo; Vizcaíno, Sergio (9 December 2011). "An Approach to the Biomechanics of the Masticatory Apparatus of Early Miocene (Santacrucian Age) South American Ungulates (Astrapotheria, Litopterna, and Notounguluata): Moment Arm Estimation Based on 3D Landmarks". Journal of Mammalian Evolution. 19 (1): 9–25. doi:10.1007/s10914-011-9179-5. hdl: 11336/17970 .
  7. Cassini, Guillermo (April 2013). "Skull Geometric Morphometrics and Paleoecology of Santacrucian (Late Early Miocene; Patagonia) Native Ungulates (Astrapotheria, Litopterna, and Notoungulata)". Ameghiniana. 50 (2): 193–216. doi:10.5710/AMGH.7.04.2013.606. hdl: 11336/26393 . S2CID   128999112.
  8. Soria, Miguel Fernando (1981). "Los Litopterna del Colhuehuapense (Oligoceno tardío) de la Argentina". Revista del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia." (3): 1–54.
  9. Kramarz, Alejandro G.; Bond, Mariano (2005). "Los Litopterna (Mammalia) de la Formación Pinturas, Mioceno Temprano-Medio de Patagonia". Ameghiniana (in Spanish). 42 (3): 611–625. ISSN   1851-8044.
  10. Theosodon at Fossilworks.org