Theosodon

{{Automatic taxobox

| fossil_range = Early-Mid Miocene (Colhuehuapian-Laventan)
{{fossil range|21.0|11.8}}

| image = Theosodon patagonica skull (cropped).jpg

| image_caption = Theosodon patagonicum skull cast of a specimen from Patagonia, Argentina.

| image2 = Reports of the Princeton University Expeditions to Patagonia, 1896-1899. J. B. Hatcher in charge (1901) (14749571352).jpg

| image2_caption = T. garretorum, 1901 skeleton illustration

| taxon = Theosodon

| authority = Ameghino, 1887

| type_species = †Theosodon lydekkeri

| type_species_authority = Ameghino, 1887

| subdivision_ranks = Species

| subdivision = * T. arozquetai McGrath et al., 2018

T. fontanae Ameghino, 1891
T. garretorum Scott, 1910
T. gracilis Ameghino, 1891
T. karaikensis Ameghino, 1904
T. lallemanti Mercerat, 1891
T. lydekkeri Ameghino, 1887
T. patagonicum (Ameghino, 1891)

| synonyms = * Pseudocoelosoma{{Cite book| publisher = Princeton University Press| last = Scott| first = William B.| title = The Litopterna of the Santa Cruz Beds| location = Princeton| series = Reports of the Princeton University Expeditions to Patagonia, 1896–1899| date = 1910}}
Ameghino, 1891

}}

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

Description

File:Theosodon NT.jpg

Theosodon was long-legged with a long neck resembling modern llamas or guanacos. It was large for a litoptern, reaching up to {{convert|2|m|ft}} in length and weighing up to {{convert|125|-|170|kg|lb}}.{{cite book|last=Croft|first=Darin A.|date=18 August 2016|title=Horned Armadillos and Rafting Monkeys: The Fascinating Fossil Mammals of South America|publisher=Indiana University Press|chapter=Chucal, Chile|pages=104–117|isbn=978-0-253-02084-0}}

It had a long neck and tapir-like, three-toed feet,{{cite book |editor=Palmer, D.|year=1999 |title= The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|publisher= Marshall Editions|location=London|page= 247|isbn= 1-84028-152-9}} and like other litopterns and modern horses, tapirs and rhinos, it bore its weight on its middle toes.

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.

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.{{cite journal|last1=Schmidt|first1=Gabriela I.|last2=Ferrero|first2=Brenda S.|date=September 2014|title=Taxonomic Reinterpretation of Theosodon hystatus Cabrera and Kraglievich, 1931 (Litopterna, Macraucheniidae) and Phylogenetic Relationships of the Family|journal=Journal of Vertebrate Paleontology|volume=34|issue=5|pages=1231–1238|doi=10.1080/02724634.2014.837393|bibcode=2014JVPal..34.1231S |hdl=11336/18953|hdl-access=free}} The name Theosodon means "god tooth". The first fossils were only dental remains, but it is unknown why this name was given. 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.

File:Reports of the Princeton University Expeditions to Patagonia, 1896-1899. J. B. Hatcher in charge (1901) (14749875265).jpg

= ''"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.

= ''"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 {{convert|80.8|to|116.2|kg|lb}}. It dated to the middle Miocene, about 13 million years ago, somewhat later than most other Theosodon species.{{cite web|url=https://www.paleowire.com/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/|title=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|last=Fonseca|first=Lurdes|date=31 May 2018|access-date=6 June 2018}}

Classification

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.

{{Multiple image

| direction = horizontal

| image1 = Macraucheniidae skulls.jpg

| image2 = Macraucheniidae skulls 2.jpg

| footer = Skulls of Macraucheniidae. A: Theosodon. B: Scalabrinitherium. C: Macrauchenia. n: nasal bones.

| total_width = 480

}}

{{Barlabel

|size=18

|at1=4|label1=Cramaucheniinae|bar1=green

|at2=13|label2=Macraucheniinae|bar2=purple

|cladogram=

{{clade| style=font-size:100%; width:auto;

|1={{cladex

|1=Proadiantus

|2={{cladex

|1=Tricoelodus

|2={{cladex

|1=Polymorphis|barbegin1=green

|2={{cladex

|1=Pternoconius|bar1=green

|2=Coniopternium|bar2=green

|3=Cramauchenia|bar3=green

|4={{cladex

|1=Theosodon|barend1=green

|2={{cladex

|1=Scalabrinitherium|barbegin1=purple

|2={{cladex

|1=Oxyodontherium|bar1=purple

|2={{cladex

|1=Huayqueriana|bar1=purple

|2={{cladex

|1={{cladex

|1=Paranauchenia denticulata|bar1=purple

|2=Paranauchenia hystatus|bar2=purple

}}

|2={{cladex

|1=Cullinia|bar1=purple

|2={{cladex

|1=Promacrauchenia|bar1=purple

|2={{cladex

|1=Windhausenia|bar1=purple

|2=Xenorhinotherium|bar2=purple

|3={{cladex

|1=Macraucheniopsis|bar1=purple

|2=Macrauchenia|barend2=purple

}}

Paleobiology

File:Theosodon.jpg tuberata]]

Theosodon was a terrestrial and cursorial animal that may have lived in both forests and more open environments. 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. Theosodon had a slender jaw compared to many other litopterns, indicating its food was softer,{{cite journal|last1=Cassini|first1=Guillermo|last2=Vizcaíno|first2=Sergio|date=9 December 2011|title=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=Journal of Mammalian Evolution|volume=19|issue=1|pages=9–25|doi=10.1007/s10914-011-9179-5|hdl=11336/17970|hdl-access=free}} such as dicotyledons.{{cite journal|last=Cassini|first=Guillermo|date=April 2013|title=Skull Geometric Morphometrics and Paleoecology of Santacrucian (Late Early Miocene; Patagonia) Native Ungulates (Astrapotheria, Litopterna, and Notoungulata)|journal=Ameghiniana|volume=50|issue=2|pages=193–216|doi=10.5710/AMGH.7.04.2013.606 |hdl=11336/26393 |s2cid=128999112 |hdl-access=free}} 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.

Other fossils were found in the Sarmiento Formation of Argentina,{{Cite journal |last=Soria |first=Miguel Fernando |date=1981 |title=Los Litopterna del Colhuehuapense (Oligoceno tardío) de la Argentina. |journal=Revista del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia." |issue=3 |pages=1–54}}{{Cite journal |last1=Kramarz |first1=Alejandro G. |last2=Bond |first2=Mariano |date=2005 |title=Los Litopterna (Mammalia) de la Formación Pinturas, Mioceno Temprano-Medio de Patagonia |journal=Ameghiniana |url=https://www.ameghiniana.org.ar/index.php/ameghiniana/article/view/928 |language=es |volume=42 |issue=3 |pages=611–625 |issn=1851-8044}} the Yecua Formation of Bolivia, the Honda Group of Colombia, the Chucal and Río Frías Formations of Chile and the Ipururo Formation of Peru.[https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=43503 Theosodon] at Fossilworks.org