View source for Toxodon

{{Short description|Extinct genus of notoungulates}}
{{Use dmy dates|date=August 2021}}
{{Automatic taxobox
| fossil_range = {{fossilrange|Pliocene|Holocene}}
| image = Toxodon skeleton in BA.JPG
| image_upright = 1.1
| image_caption = Skeleton of ''Toxodon'' in Buenos Aires
| taxon = Toxodon
| authority = [[Richard Owen|Owen]], 1837
| type_species = {{extinct}}'''''Toxodon platensis'''''
| type_species_authority = Owen, 1837
| subdivision_ranks = Other species
| subdivision = * {{extinct}}''T. burmeisteri'' <small>Giebel, 1866</small>
* {{extinct}}''T. chapalmalensis'' <small>Ameghino, 1908</small>
* {{extinct}}''T. ensenadensis'' <small>Ameghino, 1887</small>
* {{extinct}}''T. expansidens'' <small>Cope, 1886</small>
* {{extinct}}''T. gracilis'' <small>Gervais and Ameghino, 1880</small>
| synonyms = '''Genus-level'''
*''Dilobodon'' <small>Ameghino, 1886</small>
*''Chapalmalodon'' <small>Pascual, 1957</small>
*''Chapadmalodon'' <small>Tonni ''et al.'', 1992 ([[lapsus#lapsus calami|lapsus calami]])</small>
'''''T. platensis'''''
*''T. angustidens'' <small>Owen, 1846</small>
*''T. owenii'' <small>Burmeister, 1866</small>
*''T. gervaisii'' <small>Gervais & Ameghino, 1880</small>
*''T. aguirrei'' <small>Ameghino, 1917</small>
*''T. gezi'' <small>Ameghino, 1917</small>
'''''T. burmeisteri'''''
*''T. paradoxus'' <small>Ameghino, 1882</small>
*''T. protoburmeisteri'' <small>Ameghino, 1887</small>
*''T. bilobidens'' <small>Ameghino, 1887</small>
'''''T. chapalmalensis'''''
*''Chapalmalodon chapalmalensis'' <small>Pascual, 1957</small>
*''T. chapadmalensis'' <small>Cione & Tonni, 1995 ([[lapsus#lapsus calami|lapsus calami]])</small>
*''T. chapalmalalensis'' <small>Oliva & Cerdeno, 2007 ([[lapsus#lapsus calami|lapsus calami]])</small>
'''''T. ensenadensis'''''
*''T. giganteus'' <small>Moreno, 1888</small>
*''T. elongatus'' <small>Roth, 1898</small>
'''''T. gracilis'''''
*''T. voghti'' <small>Moreno, 1888</small>
}}
'''''Toxodon''''' (meaning "bow tooth" in reference to the curvature of the teeth) is an [[Extinction|extinct]] [[genus]] of large [[ungulate]] native to [[South America]] from the [[Pliocene]] to the end of the [[Late Pleistocene]].<ref>{{cite journal | vauthors = Baffa O, Brunetti A, Karmann I, Neto CM | title = ESR dating of a toxodon tooth from a Brazilian karstic cave | journal = Applied Radiation and Isotopes | volume = 52 | issue = 5 | pages = 1345–9 | date = May 2000 | pmid = 10836452 | doi = 10.1016/S0969-8043(00)00093-2 | bibcode = 2000AppRI..52.1345B }}</ref><ref name=":0">{{cite book |url= https://books.google.com/books?id=mbU-F42JU1AC&q=Propaopus&pg=PA349|title=Holocene Extinctions |last=Turvey |first=Samuel T. | name-list-style = vanc |date=2009-05-28 |publisher=OUP Oxford |isbn=978-0-19-157998-1 }}</ref> ''Toxodon'' is a member of [[Notoungulata]], an order of extinct [[South American native ungulates]] distinct from the two living ungulate orders that had been indigenous to the continent for over 60 million years since the early Cenozoic, prior to the arrival of living ungulates into South America around 2.5 million years ago during the [[Great American Interchange]].<ref name=":2">{{Cite journal |last1=Croft |first1=Darin A. |last2=Gelfo |first2=Javier N. |last3=López |first3=Guillermo M. |name-list-style=vanc |date=30 May 2020 |title=Splendid Innovation: The Extinct South American Native Ungulates |url=https://www.annualreviews.org/doi/10.1146/annurev-earth-072619-060126 |journal=[[Annual Review of Earth and Planetary Sciences]] |language=en |volume=48 |issue=1 |pages=259–290 |bibcode=2020AREPS..48..259C |doi=10.1146/annurev-earth-072619-060126 |issn=0084-6597 |s2cid=213737574 |access-date=2 January 2024}}</ref> ''Toxodon'' is a member of the family [[Toxodontidae]], which includes medium to large sized herbivores.<ref>{{Cite journal |last1=Cassini |first1=Guillermo H. |last2=Flores |first2=David A. |last3=Vizcaíno |first3=Sergio F. |date=July 2012 |title=Postnatal ontogenetic scaling of Nesodontine (Notoungulata, Toxodontidae) cranial morphology: Nesodontine cranial allometry |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1463-6395.2011.00501.x |journal=Acta Zoologica |language=en |volume=93 |issue=3 |pages=249–259 |doi=10.1111/j.1463-6395.2011.00501.x |hdl-access=free |hdl=11336/81335}}</ref> ''Toxodon'' was one of the largest members of Toxodontidae and Notoungulata, with ''Toxodon platensis'' having an estimated body mass of {{Convert|1000-1200|kg|lb}}.
Remains of ''Toxodon'' were first collected by [[Charles Darwin]] during the [[Second voyage of HMS Beagle|voyage of the Beagle]] in 1832-33, and later scientifically named by [[Richard Owen]] in 1837. Both Darwin and Owen were puzzled by ''Toxodon''<nowiki/>'s unusual anatomical features, including its long, ever-growing cheek teeth.
''Toxodon'' has been found across much of South America, excluding southern Patagonia, the Andes and northeastern-most region of the continent.<ref>{{Citation |last1=Cione |first1=Alberto Luis |title=The GABI in Southern South America |date=2015 |work=The Great American Biotic Interchange |pages=71–96 |url=https://link.springer.com/10.1007/978-94-017-9792-4_3 |access-date=2024-01-30 |place=Dordrecht |publisher=Springer Netherlands |language=en |doi=10.1007/978-94-017-9792-4_3 |isbn=978-94-017-9791-7 |last2=Gasparini |first2=Germán Mariano |last3=Soibelzon |first3=Esteban |last4=Soibelzon |first4=Leopoldo Héctor |last5=Tonni |first5=Eduardo Pedro|s2cid=127856682 }}</ref> Evidence suggests that ''Toxodon'' was ecologically plastic and able to adapt its diet to local conditions.<ref name=":3" />
''Toxodon'' became extinct as part of the [[Late Pleistocene extinctions|end-Pleistocene extinctions]] around 12,000 years ago, along with most large mammals across the Americas. The extinctions followed the [[Peopling of the Americas|arrival of humans to South America]], who may have been a contributory factor in the extinctions.<ref name=":2" /> Several sites have been found suggesting human interaction with ''Toxodon''.
== Taxonomy and evolution ==
[[File:Toxodon platensis.jpg|thumb|Historical restoration of ''Toxodon platensis'' from 1913]]
[[Charles Darwin]], who was in South America as part of the [[Second voyage of HMS Beagle|second voyaging expedition of the HMS Beagle]], was one of the first to collect ''Toxodon'' fossils.<ref name=":8" /> In September-October 1832 and October 1833, Darwin collected several isolated teeth as well as a mandible from various localities in northern Argentina.<ref name=":8" /> On November 26, 1833, Darwin paid 18&nbsp;pence (equivalent to £6.40 pound sterling in 2018<ref name=":1" />) for a ''T.&nbsp;platensis'' skull from a farmer in Uruguay.<ref>{{cite magazine |last=Quammen |first=D. |date=February 2009 |title=Darwin's first clues |magazine=[[National Geographic (magazine)|National Geographic]] |page=45}}</ref><ref>{{Cite web |title=Darwin's Fossil Mammals - Toxodon platensis - cranium - Data Portal |url=https://data.nhm.ac.uk/dataset/darwins-fossil-mammals/resource/dadf577f-fa52-4f48-8cdb-d11c98448009 |access-date=2024-11-28 |website=data.nhm.ac.uk |language=en-GB}}</ref> In his book covering the expedition, ''[[The Voyage of the Beagle]].'' Darwin wrote, "November&nbsp;26th – I set out on my return in a direct line for Montevideo. Having heard of some giant's bones at a neighbouring farm-house on the Sarandis, a small stream entering the Rio Negro, I rode there accompanied by my host, and purchased for the value of eighteen pence the head of the ''Toxodon''." The skull had been propped up against a fence and been used as target practice for throwing stones by local children, who had knocked out its teeth.<ref>{{cite book |last=Darwin |first=Charles |title=[[The Voyage of the Beagle]] |date=1997 |publisher=Penguin |isbn=978-0-14-043268-8 |editor1-last=Browne |editor1-first=J. |quote=Read, 19th&nbsp;April 1837. A detailed account will appear in the first part of the zoology of ''[[The Voyage of the Beagle|Voyage of the Beagle]]''. |editor2-last=Neve |editor2-first=M. |orig-year=1839}}</ref><ref name=":1">{{Cite news |last=Kennedy |first=Maev |date=2018-04-06 |title=Darwin's lost fossils – including a sloth the size of a car – to be made public |url=https://www.theguardian.com/science/2018/apr/06/charles-darwin-lost-fossils-including-a-sloth-the-size-of-a-car-to-be-made-public |access-date=2024-11-28 |work=The Guardian |language=en-GB |issn=0261-3077}}</ref> Since Darwin discovered that the fossils of similar mammals of South America were different from those in Europe, he invoked many debates about the evolution and natural selection of animals.
In his own words, Darwin wrote down in his journal,
{{blockquote| Lastly, the ''Toxodon'', perhaps one of the strangest animals ever discovered: In size it equaled an [[elephant]] or [[megatherium]], but the structure of its teeth, as [[Richard Owen|Mr.&nbsp;Owen]] states, proves indisputably that it was intimately related to the [[Rodent|Gnawers]], the order which, at the present day, includes most of the smallest quadrupeds: In many details it is allied to the [[Pachydermata]]: Judging from the position of its eyes, ears, and nostrils, it was probably aquatic, like the [[Dugong]] and [[Manatee]], to which it is also allied. How wonderfully are the [[order (biology)|different Orders]], at the present time so well separated, blended together in different points of the structure of the Toxodon!}}
''Toxodon'' and its [[type species]], ''T. platensis'', were described in 1837 by [[Richard Owen]] based on remains collected by Darwin, in a paper titled "''A description of the cranium of the Toxodon platensis, a gigantic extinct mammiferous species, referrible by its dentition to the Rodentia, but with affinities to the [[Pachydermata]] and the herbivorous [[Cetacea]]''", reflecting the many unusual characterstics of its anatomy.<ref name=":8">Fernicola, J. C., Vizcaino, S. F., & De Iuliis, G. (2009). The fossil mammals collected by Charles Darwin in South America during his travels on board the HMS Beagle. ''Revista De La Asociación Geológica Argentina'', ''64''(1), 147-159. Retrieved from <nowiki>https://revista.geologica.org.ar/raga/article/view/1339</nowiki></ref>
=== Evolution ===
''Toxodon'' is a member of [[Notoungulata]], a group of [[South American native ungulates]] that had been part of the fauna of South America since the [[Paleocene]], over 60 million years ago, and had evolved in isolation in South America, prior to the arrival of living ungulates in South America around 2.5 million years ago as part of the [[Great American Interchange]].<ref name=":2" /> Notoungulata represents the most diverse group of indigenous South American ungulates, with over 150 described genera in 13 different families.<ref>{{Cite journal |last1=Rezende Castro |first1=Luis Otavio |last2=García-López |first2=Daniel A. |last3=Bergqvist |first3=Lilian Paglarelli |last4=De Araújo-Júnior |first4=Hermínio Ismael |date=2021-06-30 |title=A New Basal Notoungulate from the Itaboraí Basin (Paleogene) of Brazil |url=https://bioone.org/journals/ameghiniana/volume-58/issue-3/AMGH.05.02.2021.3387/A-New-Basal-Notoungulate-from-the-Itabora%c3%ad-Basin-Paleogene-of/10.5710/AMGH.05.02.2021.3387.full |journal=Ameghiniana |volume=58 |issue=3 |doi=10.5710/AMGH.05.02.2021.3387 |issn=0002-7014 |s2cid=234220780}}</ref> Notoungulates are morphologically diverse, including forms morphologically distant from ''Toxodon'' such as rodent and rabbit-like forms.<ref name=":2" />
Analysis of [[collagen]] sequences obtained from ''Toxodon'' as well as from the [[litoptern]] (another group of indigenous South American ungulates) ''[[Macrauchenia]]'' found that notoungulates and litopterns were closely related to each other, and form a [[sister group]] to [[perissodactyl]]s (which contains equids, rhinoceroses and tapirs) as part of the clade [[Panperissodactyla]], making them true [[ungulate]]s.<ref name="WelkerCollins2015">{{cite journal |display-authors=6 |vauthors=Welker F, Collins MJ, Thomas JA, Wadsley M, Brace S, Cappellini E, Turvey ST, Reguero M, Gelfo JN, Kramarz A, Burger J, Thomas-Oates J, Ashford DA, Ashton PD, Rowsell K, Porter DM, Kessler B, Fischer R, Baessmann C, Kaspar S, Olsen JV, Kiley P, Elliott JA, Kelstrup CD, Mullin V, Hofreiter M, Willerslev E, Hublin JJ, Orlando L, Barnes I, MacPhee RD |date=June 2015 |title=Ancient proteins resolve the evolutionary history of Darwin's South American ungulates |url=http://eprints.whiterose.ac.uk/91438/1/Welker_postprint.docx |journal=Nature |volume=522 |issue=7554 |pages=81–4 |bibcode=2015Natur.522...81W |doi=10.1038/nature14249 |pmid=25799987 |s2cid=4467386 |hdl-access=free |hdl=11336/14769}}</ref><ref name="Buckley2015">{{cite journal |last=Buckley |first=M. |date=7 May 2015 |title=Ancient collagen reveals evolutionary history of the endemic South American 'ungulates' |journal=Proceedings of the Royal Society&nbsp;B: Biological Sciences |volume=282 |issue=1806 |pages=20142671 |doi=10.1098/rspb.2014.2671 |pmc=4426609 |pmid=25833851}}</ref> This finding has been corroborated by an analysis of mitochondrial DNA extracted from a ''Macrauchenia'' fossil, which yielded a date of 66 million years ago for the time of the split from perissodactyls.<ref name="Westbury2017">{{cite journal |display-authors=6 |vauthors=Westbury M, Baleka S, Barlow A, Hartmann S, Paijmans JL, Kramarz A, Forasiepi AM, Bond M, Gelfo JN, Reguero MA, López-Mendoza P, Taglioretti M, Scaglia F, Rinderknecht A, Jones W, Mena F, Billet G, de Muizon C, Aguilar JL, MacPhee RD, Hofreiter M |date=June 2017 |title=A mitogenomic timetree for Darwin's enigmatic South American mammal ''Macrauchenia patachonica'' |journal=Nature Communications |volume=8 |page=15951 |bibcode=2017NatCo...815951W |doi=10.1038/ncomms15951 |pmc=5490259 |pmid=28654082}}</ref>
''Toxodon'' belongs to [[Toxodontidae]], a large bodied group of notoungulates which first appeared in the [[Late Oligocene]] ([[Deseadan]]), ~28-23 million years ago,<ref>{{Cite journal |last1=Bonini |first1=Ricardo A. |last2=Schmidt |first2=Gabriela I. |last3=Reguero |first3=Marcelo A. |last4=Cerdeño |first4=Esperanza |last5=Candela |first5=Adriana M. |last6=Solís |first6=Natalia |date=May 2017 |title=First record of Toxodontidae (Mammalia, Notoungulata) from the late Miocene–early Pliocene of the southern central Andes, NW Argentina |url=https://www.cambridge.org/core/product/identifier/S0022336016001608/type/journal_article |journal=Journal of Paleontology |language=en |volume=91 |issue=3 |pages=566–576 |doi=10.1017/jpa.2016.160 |bibcode=2017JPal...91..566B |issn=0022-3360|hdl=11336/49805 |hdl-access=free }}</ref> and underwent a great radiation during the [[Miocene]] epoch (~23-5.3 million years ago), when they reached their apex of diversity.<ref>{{Cite journal |last1=Ferrero |first1=Brenda S. |last2=Schmidt |first2=Gabriela I. |last3=Costamagna |first3=Donato |last4=Miño-Boilini |first4=Ángel R. |last5=Zurita |first5=Alfredo E. |last6=Quiñones |first6=Sofía I. |last7=Cuadrelli |first7=Francisco |last8=Luna |first8=Carlos A. |last9=Solís |first9=Natalia |last10=Candela |first10=Adriana M. |date=March 2024 |title=First record of Posnanskytherium (Notoungulata, Toxodontidae) in the late Neogene of eastern Puna, Argentina |url=https://link.springer.com/10.1007/s10914-023-09700-5 |journal=Journal of Mammalian Evolution |language=en |volume=31 |issue=1 |doi=10.1007/s10914-023-09700-5 |issn=1064-7554}}</ref> The diversity of toxodontids, along with other notoungulates began to decline from around the [[Pliocene]] onwards,<ref name=":2" /> possibly as a result of climate change, as well as the arrival of competitors and predators from North America during the Great American Interchange following formation of the [[Isthmus of Panama]].<ref>{{Cite journal |last1=Solórzano |first1=Andrés |last2=Núñez-Flores |first2=Mónica |last3=Rodríguez-Serrano |first3=Enrique |date=November 2024 |title=The rise and fall of notoungulates: How Andean uplift, available land area, competition, and depredation driven its diversification dynamics |url=https://linkinghub.elsevier.com/retrieve/pii/S1342937X24002193 |journal=Gondwana Research |language=en |volume=135 |pages=116–132 |doi=10.1016/j.gr.2024.08.002|bibcode=2024GondR.135..116S }}</ref> By the [[Late Pleistocene]] ([[Lujanian]]), the once great diversity of notoungulates had declined to only a few of species of toxodontids, with all other notoungulate families having become extinct.<ref name=":2" />
[[Cladogram]] of Toxodontidae, showing the position of ''Toxodon'' relative to other toxodontids, after Forasiepi et al, 2014:<ref name="AMFetal2014">{{cite journal |display-authors=6 |vauthors=Forasiepi AM, Cerdeno E, Bond M, Schmidt GI, Naipauer M, Straehl FR, Martinelli AG, Garrido AC, Schmitz MD, Crowley JL |year=2014 |title=New toxodontid (Notoungulata) from the Early Miocene of Mendoza, Argentina |url=http://sedici.unlp.edu.ar/handle/10915/127531 |journal=Paläontologische Zeitschrift |volume=89 |issue=3 |pages=611–634 |doi=10.1007/s12542-014-0233-5 |s2cid=129293436 |hdl-access=free |hdl=11336/20443}}</ref>
{{clade| style=font-size:85%; line-height:85%
|label1=&nbsp;  †'''[[Notoungulata]]'''  &nbsp;
|1={{clade
|1=†''[[Pampahippus|Pampahippus arenalesi]]''
|2={{clade
|1={{clade
|1=†''[[Rhynchippus]]'' spp.
|2={{clade
|1=†[[Scarrittia|''Scarrittia canquelensis'']]
|2=†[[Leontinia|''Leontinia gaudri'']] }} }}
|label2=&nbsp; †'''[[Toxodontidae]]''' &nbsp;
|2={{clade
|1=†[[Proadinotherium|''Proadinotherium leptognathum'']]
|2={{clade
|1={{clade
|1=†''[[Adinotherium]]'' spp.
|2={{clade
|1=†''[[Nesodon|Nesodon taweretus]]''
|2=†''[[Nesodon|Nesodon imbricatus]]'' }} }}
|2={{clade
|1=†''[[Palyeidodon obtusum]]''
|2={{clade
|1=†''[[Hyperoxotodon|Hyperoxotodon speciosus]]''
|2={{clade
|1={{clade
|1=†''[[Nonotherium|Nonotherium henningi]]''
|2=†''[[Xotodon]]'' spp. }}
|2={{clade
|1={{clade
|1=†''[[Andinotoxodon|Andinotoxodon bolivariensis]]''
|2={{clade
|1={{clade
|1=†''[[Dinotoxodon|Dinotoxodon paranensis]]''
|2=†'''''Toxodon platensis''''' }}
|2={{clade
|1=†''[[Gyrinodon|Gyrinodon quassus]]''
|2={{clade
|1=†''[[Ocnerotherium|Ocnerotherium intermedium]]''
|2=†''[[Hoffstetterius|Hoffstetterius imperator]]'' }} }} }} }}
|2={{clade
|1=†''[[Posnanskytherium|Posnanskytherium desaguaderoi]]''
|2={{clade
|1=†''[[Pisanodon|Pisanodon nazari]]''
|2={{clade
|1=†''[[Pericotoxodon|Pericotoxodon platignathus]]''
|2={{clade
|1={{clade
|1=†''[[Calchaquitherium|Calchaquitherium mixtum]]''
|2=†''[[Mixotoxodon|Mixotoxodon larensis]]'' }}
|2={{clade
|1=†''[[Paratrigodon|Paratrigodon euguii]]''
|2=†''[[Trigodon|Trigodon gaudri]]'' }} }}
}} }} }} }} }} }} }} }} }} }} }}
}}
=== Species ===
There has not been a recent taxonomic revision of the genus ''Toxodon'', leaving the number of valid species uncertain.<ref name=":4">{{Cite journal |last1=Carrillo |first1=Juan D. |last2=Püschel |first2=Hans P. |date=December 2023 |title=Pleistocene South American native ungulates (Notoungulata and Litopterna) of the historical Roth collections in Switzerland, from the Pampean Region of Argentina |journal=[[Swiss Journal of Palaeontology]] |language=en |volume=142 |issue=1 |page=28 |bibcode=2023SwJP..142...28C |doi=10.1186/s13358-023-00291-5 |issn=1664-2376 |pmc=10558389 |pmid=37810207 |doi-access=free}}</ref>
The species ''Toxodon chapalmalensis'' is known from the [[Pliocene]] ([[Montehermosan]]-[[Chapadmalalan]]) of Argentina,<ref>{{Cite journal |last1=Tomassini |first1=Rodrigo L. |last2=Montalvo |first2=Claudia I. |last3=Deschamps |first3=Cecilia M. |last4=Manera |first4=Teresa |date=December 2013 |title=Biostratigraphy and biochronology of the Monte Hermoso Formation (early Pliocene) at its type locality, Buenos Aires Province, Argentina |url=http://dx.doi.org/10.1016/j.jsames.2013.08.002 |journal=[[Journal of South American Earth Sciences]] |volume=48 |pages=31–42 |bibcode=2013JSAES..48...31T |doi=10.1016/j.jsames.2013.08.002 |issn=0895-9811 |access-date=2 January 2024 |via=Elsevier Science Direct |hdl-access=free |hdl=11336/21606}}</ref> while ''Toxodon platensis'', the type species, is known from the [[Pleistocene wolf|Pleistocene]]. The validity of other potential species like ''Toxodon darwini'' Burmeister, 1866, and ''Toxodon ensenadensis'' Ameghino, 1887 from the Early Pleistocene of Argentina is uncertain, and the species ''Toxodon gezi'' C. Ameghino, 1917 and ''Toxodon aguirrei'' Ameghino, 1917 have been considered junior synonyms of ''Toxodon platensis'' by recent authors.<ref>{{Cite journal |last1=Guérin |first1=Claude |last2=Faure |first2=Martine |date=March 2013 |title=Un nouveau Toxodontidae (Mammalia, Notoungulata) du Pléistocène supérieur du Nordeste du Brésil |url=http://dx.doi.org/10.5252/g2013n1a7 |journal=Geodiversitas |volume=35 |issue=1 |pages=155–205 |doi=10.5252/g2013n1a7 |issn=1280-9659}}</ref> Some recent authors have argued that that ''Toxodon gracilis'' Gervais and Ameghino, 1880, should be recognised as a distinct species from the Pleistocene of the Pampas significantly smaller than ''T. platensis,'' with these authors suggesting that ''T. platensis'' and ''T. gracilis'' represent the only valid species of ''Toxodon'' in the Pleistocene of the Pampas region.<ref name=":4" /> Other authors have argued that all Pleistocene ''Toxodon'' species should be considered synonymous with ''T. platensis''.<ref>{{Cite journal |last1=Costamagna |first1=Donato |last2=Ferrero |first2=Brenda S. |last3=Giri |first3=Federico |last4=Ribeiro |first4=Ana María |date=August 2024 |title=Study of the shape and size in lower molars of Toxodon platensis (Mammalia: Toxodontidae) of the Late Pleistocene of South America |url=https://linkinghub.elsevier.com/retrieve/pii/S0016699524000627 |journal=Geobios |language=en |doi=10.1016/j.geobios.2024.05.005}}</ref>
== Description ==
The bodyform of ''Toxodon'' and other toxodontids have been compared to those of hippopotamuses and rhinoceroses.<ref name=":42">{{Citation |last1=Cassini |first1=Guillermo H. |title=Paleobiology of Santacrucian native ungulates (Meridiungulata: Astrapotheria, Litopterna and Notoungulata) |date=2012-10-11 |work=Early Miocene Paleobiology in Patagonia |pages=243–286 |editor-last=Vizcaíno |editor-first=Sergio F. |url=https://www.cambridge.org/core/product/identifier/9780511667381%23c19461-14-1/type/book_part |access-date=2023-06-29 |edition=1 |publisher=Cambridge University Press |doi=10.1017/cbo9780511667381.015 |isbn=978-0-511-66738-1 |last2=Cerdeño |first2=Esperanza |last3=Villafañe |first3=Amalia L. |last4=Muñoz |first4=Nahuel A. |editor2-last=Kay |editor2-first=Richard F. |editor3-last=Bargo |editor3-first=M. Susana}}</ref> ''Toxodon platensis'' is one of the largest known toxodontids and notoungulates, with an estimated body mass of approximately {{Convert|1000-1200|kg|lb}},<ref name=":02">{{Cite journal |last1=Nelson |first1=Allison |last2=Engelman |first2=Russell K. |last3=Croft |first3=Darin A. |date=September 2023 |title=How to weigh a fossil mammal? South American notoungulates as a case study for estimating body mass in extinct clades |url=https://link.springer.com/10.1007/s10914-023-09669-1 |journal=Journal of Mammalian Evolution |language=en |volume=30 |issue=3 |pages=773–809 |doi=10.1007/s10914-023-09669-1 |issn=1064-7554}}</ref> and a body length of around {{Convert|2.7|m}}.<ref name=":12">Fariña RA, Vizcaíno SF& De Iuliis G. 2012 ''Megafauna: giant beasts of Pleistocene South America''. Bloomington, IN: Indiana University Press. p. 264-265</ref>
The skull of ''Toxodon'' is proportionally large,<ref name=":02" /> and triangular in shape when viewed from above.<ref>{{Cite journal |last1=Ferrero |first1=Brenda S. |last2=Schmidt |first2=Gabriela I. |last3=Pérez-García |first3=María I. |last4=Perea |first4=Daniel |last5=Ribeiro |first5=Ana M. |date=2021-08-27 |title=A new Toxodontidae (Mammalia, Notoungulata) from the upper Pliocene–lower Pleistocene of Uruguay |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2021.2023167 |journal=Journal of Vertebrate Paleontology |language=en |volume=41 |issue=5 |doi=10.1080/02724634.2021.2023167 |bibcode=2021JVPal..41E3167F |issn=0272-4634}}</ref> All of the teeth in the jaws are high-crowned ([[hypsodont]]).<ref name=":22">Scott WB. [https://www.biodiversitylibrary.org/item/212227#page/175/mode/1up Mammalia of the Santa Cruz Beds. Volume VI, Paleontology. Part II, Toxodontia]. In: Scott WB, editor. Reports of the Princeton University Expeditions to Patagonia, 1896–1899. Stuttgart: Princeton University, E. Schweizerbart’sche Verlagshandlung (E. Nägele); 1912. pp. 211-216</ref> Like other toxodontids, the upper and lower first [[Incisor|incisors]] (I1 and i1) are large and protrude, with the second upper incisors (I2) and lower third incisors (i3) being modified into evergrowing tusks.<ref name=":9">{{Cite journal |last1=Braunn |first1=Patrícia |last2=Ferigolo |first2=Jorge |last3=Ribeiro |first3=Ana |date=2021 |title=Enamel microstructure of permanent and deciduous teeth of notoungulate Toxodon: development, functional and evolutionary implications |url=http://www.app.pan.pl/article/item/app007722020.html |journal=Acta Palaeontologica Polonica |volume=66 |doi=10.4202/app.00772.2020|doi-access=free }}</ref> The upper incisors display an arched shape,<ref name=":52">S.F. Vizcaino, R.A. Farina, J.C. Fernicola "[https://www.researchgate.net/publication/288069897_Young_Darwin_and_the_ecology_and_extinction_of_Pleistocene_South_American_fossil_mammals#pf4 Young Darwin and the ecology and extinction of Pleistocene South American fossil mammals]" Revista de la Asociacion Geologica Argentina, 64 (2009), pp. 160-169</ref> while the lower incisors project horizontally forwards at the front of the lower jaw.<ref name=":22" /><ref name=":52" /> The wide front of the lower jaw with the horizontally-arranged incisors has been described as "spade-like".<ref name=":52" /> There is a gap ([[diastema]]) between the incisors and the cheek teeth.<ref name=":7">E. Anderson Who's who in the Pleistocene: a mammalian bestiary P.S. Martin, R.G. Klein (Eds.), Quaternary Extinctions: a Prehistoric Revolution, The University of Arizona Press, Tucson, AZ (1989), pp. 64</ref> Like other derived toxodontids, ''Toxodon'' had long, ever-growing (hypselodont) cheek (premolar and molar) teeth,<ref name=":32">{{Cite journal |last1=Gomes Rodrigues |first1=Helder |last2=Herrel |first2=Anthony |last3=Billet |first3=Guillaume |date=2017-01-31 |title=Ontogenetic and life history trait changes associated with convergent ecological specializations in extinct ungulate mammals |journal=Proceedings of the National Academy of Sciences |language=en |volume=114 |issue=5 |pages=1069–1074 |bibcode=2017PNAS..114.1069G |doi=10.1073/pnas.1614029114 |issn=0027-8424 |pmc=5293108 |pmid=28096389 |doi-access=free}}</ref> with the name ''Toxodon'' deriving from the curved shape of the upper molars, which are bowed inwards towards the midline of the skull to fit in the upper jaw. Evergrowing cheek teeth are unknown in any living ungulates, but are present in some other mammal groups like [[Wombat|wombats]] and [[Rodent|rodents]]. The surface of the cheek teeth is primarily composed of [[Dentin|dentine]].<ref name=":2" />
The [[thoracic vertebrae]] of ''Toxodon'' have elongate [[neural spines]], which likely anchored muscles which supported the large head.<ref name=":12" /> The legs of ''Toxodon'' are relatively short, with their bones being robust.<ref name=":6">{{Cite journal |last1=Oliveira |first1=Édison V. |last2=Bélo |first2=Pétrius S. |last3=Fambrini |first3=Gelson L. |last4=Sial |first4=Alcides N. |last5=Silva |first5=Ana K.B. |last6=Barreto |first6=Alcina M.F. |date=October 2019 |title=A new late Pleistocene ichnological site with mammal footprints from Brazil |url=https://linkinghub.elsevier.com/retrieve/pii/S089598111930135X |journal=Journal of South American Earth Sciences |language=en |volume=94 |pages=102216 |doi=10.1016/j.jsames.2019.102216|bibcode=2019JSAES..9402216O }}</ref> The hindlimb is considerably longer than the forelimb, resulting in the back being elevated and the shoulder, neck and head being relatively low.<ref name=":22" /> The ulna has a strongly backwardly projecting [[olecranon]] process similar to that of rhinos, suggesting that the front leg was held extended when standing.<ref name=":12" /> The (distal) part of the femur closest to the foot shows a pronounced medial trochlear ridge, which likely served along with the [[patella]] (kneecap) to allow the knees to be locked when standing akin to the [[stay apparatus]] of living horses as an energy saving mechanism.<ref>Shockey BJ. 2001. "[https://www.app.pan.pl/article/item/app46-277.html Specialized knee joints in some extinct, endemic, South American herbivores]" ''[[Acta Palaeontologica Polonica]]'' 46:277–88</ref> There are three functional digits on each foot,<ref name=":6" /><ref name=":7" /> which are tipped with [[hoof]]-like [[Phalanx bone|phalanges]].<ref>{{Cite journal |last1=Lorente |first1=Malena |last2=Gelfo |first2=Javier N. |last3=López |first3=Guillermo M. |date=April 2019 |title=First skeleton of the notoungulate mammal Notostylops murinus and palaeobiology of Eocene Notostylopidae |url=https://www.idunn.no/doi/10.1111/let.12310 |journal=Lethaia |language=en |volume=52 |issue=2 |pages=244–259 |doi=10.1111/let.12310 |bibcode=2019Letha..52..244L |issn=0024-1164}}</ref>
{{gallery|ToxodonMLP1 (cropped).jpg|Mount at Museo de La Plata, Argentina|Toxodon platensis skull 3.JPG|Skull in front-on view|File:Crani de Toxodon, Museu de Ciències Naturals de València.JPG|Skull in oblique view|Em - Toxodon platensis - 1.jpg|Skeleton in side-on view|Cráneo de Toxodon.jpg|Skull in side-on view|||||||width=200|height=180|align=center}}
== Distribution ==
''Toxodon'' had a widespread distribution in South America east of the Andes, ranging from northern Argentina and Bolivia to the western Amazon on the Peru-Brazil border, to [[Northeast Region, Brazil|Northeast Brazil]].<ref name=":5" /> Although some authors suggest that the distribution of ''Toxodon'' extended into Venezuela,<ref name=":9" /> other authors suggest that the related ''[[Mixotoxodon]]'' (which ranged as far north as the southern United States) was the only toxodontid present in the region during the Pleistocene.<ref>{{Cite journal |last1=Carrillo-Briceño |first1=Jorge Domingo |last2=Amson |first2=Eli |last3=Zurita |first3=Alfredo |last4=Sánchez-Villagra |first4=Marcelo Ricardo |date=2016-12-12 |title=Hermann Karsten (1817–1908): a German naturalist in the Neotropics and the significance of his paleovertebrate collection |url=https://fr.copernicus.org/articles/20/21/2016/ |journal=Fossil Record |language=en |volume=20 |issue=1 |pages=21–36 |doi=10.5194/fr-20-21-2016 |doi-access=free |bibcode=2016FossR..20...21C |issn=2193-0074|hdl=11336/81200 |hdl-access=free }}</ref>
== Palaeobiology ==
[[File:A-Pleistocene-meso-megamammals-from-Sergipe-Brazil.jpg|thumb|353x353px|Life restoration of ''Toxodon platensis'' (background centre-left) in a Pleistocene Brazilian landscape, alongside the giant ground sloth [[Eremotherium laurillardi|''Eremotherium laurillardi'']] (foreground right), the [[glyptodonts]] ''[[Glyptotherium]]'' and ''[[Panochthus]]'', the [[pampathere]] [[Holmesina|''Holmesina paulacoutoi'']] (midground centre-left)  and the armadillo [[Pachyarmatherium|''Pachyarmatherium brasiliense'']] (foreground left)]]
Although some authors have suggested that ''Toxodon'' was semiaquatic based on the similarity of some aspects of its anatomy to hippopotamuses, this has been disputed by other authors, and analysis of oxygen isotope ratios (which differ between terrestrial and aquatic animals) suggests a terrestrial lifestyle for ''Toxodon''.<ref>{{Cite journal |last1=Lopes |first1=Renato P. |last2=Ribeiro |first2=Ana Maria |last3=Dillenburg |first3=Sérgio Rebello |last4=Schultz |first4=Cesar Leandro |date=January 2013 |title=Late middle to late Pleistocene paleoecology and paleoenvironments in the coastal plain of Rio Grande do Sul State, Southern Brazil, from stable isotopes in fossils of Toxodon and Stegomastodon |url=https://linkinghub.elsevier.com/retrieve/pii/S0031018212006293 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=369 |pages=385–394 |doi=10.1016/j.palaeo.2012.10.042|bibcode=2013PPP...369..385L }}</ref><ref name=":3" /> As such, it has been suggested that ''Toxodon'' was probably more ecologically comparable to rhinoceroses.<ref name=":3" />
''Toxodon'' is suggested to have been capable of moving at considerable speed.<ref name=":52" /> ''Toxodon'' is believed to have been ecologically plastic and have had a wide niche breadth,<ref name=":3">{{Cite journal |last1=Dantas |first1=Mário André Trindade |last2=Cherkinsky |first2=Alexander |last3=Bocherens |first3=Hervé |last4=Drefahl |first4=Morgana |last5=Bernardes |first5=Camila |last6=França |first6=Lucas de Melo |date=15 August 2017 |title=Isotopic paleoecology of the Pleistocene megamammals from the Brazilian Intertropical Region: Feeding ecology (δ13C), niche breadth and overlap |url=https://www.sciencedirect.com/science/article/pii/S0277379117300616 |journal=[[Quaternary Science Reviews]] |volume=170 |pages=152–163 |doi=10.1016/j.quascirev.2017.06.030 |issn=0277-3791 |access-date=2 January 2024 |via=Elsevier Science Direct}}</ref> with its diet varying according to local conditions,<ref>{{Cite journal |last1=Pansani |first1=Thaís Rabito |last2=Muniz |first2=Fellipe Pereira |last3=Cherkinsky |first3=Alexander |last4=Pacheco |first4=Mírian Liza Alves Forancelli |last5=Dantas |first5=Mário André Trindade | name-list-style = vanc |date=October 2019|title=Isotopic paleoecology (δ13C, δ18O) of Late Quaternary megafauna from Mato Grosso do Sul and Bahia States, Brazil |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119302926 |journal=[[Quaternary Science Reviews]] |language=en |volume=221 |pages=105864 |doi=10.1016/j.quascirev.2019.105864 |bibcode=2019QSRv..22105864P |s2cid=202200336 }}</ref> with an almost totally [[C3 carbon fixation|C<sub>3</sub>]] [[Browsing (herbivory)|browsing]] diet in the Amazon rainforest, mixed feeding C<sub>3</sub> in [[Bahia]] and the [[Pampas]] to almost completely [[C4 carbon fixation|C<sub>4</sub>]] dominated [[Grazing (behaviour)|grazing]] diet in the [[Gran Chaco|Chaco]].<ref name=":5">{{Cite journal |last=MacFadden |first=Bruce J. |name-list-style=vanc |date=September 2005 |title=Diet and habitat of toxodont megaherbivores (Mammalia, Notoungulata) from the late Quaternary of South and Central America |url=http://linkinghub.elsevier.com/retrieve/pii/S0033589405000712 |journal=[[Quaternary Research]] |language=en |volume=64 |issue=2 |pages=113–124 |bibcode=2005QuRes..64..113M |doi=10.1016/j.yqres.2005.05.003 |access-date=2 January 2024 |via=Elsevier Science Direct}}</ref> Within the Brazilian Intertropical Region, local climate had little impact on the diet of ''T. platensis''.<ref>{{Cite journal |last1=Gomes |first1=Verônica Santos |last2=Lessa |first2=Carlos Micael Bonfim |last3=Oliveira |first3=Gustavo Ribeiro de |last4=Bantim |first4=Renan Alfredo Machado |last5=Sayão |first5=Juliana |last6=Bocherens |first6=Hervé |last7=Araújo-Júnior |first7=Hermínio Ismael de |last8=Dantas |first8=Mário André Trindade |date=January 2023 |title=Seasonal variations in diet (δ13C) and climate (δ 18O) inferred through toxodonts enamel teeth during the Late Pleistocene in the brazilian intertropical region |url=https://linkinghub.elsevier.com/retrieve/pii/S0895981122004345 |journal=[[Journal of South American Earth Sciences]] |language=en |volume=121 |pages=104148 |doi=10.1016/j.jsames.2022.104148 |access-date=19 April 2024 |via=Elsevier Science Direct}}</ref> Although ''Toxodon'' is thought to have inhabited open landscapes like [[steppe]] and [[Savanna|savannah]],<ref>{{Cite journal |last1=Dantas |first1=Mário André Trindade |last2=Dutra |first2=Rodrigo Parisi |last3=Cherkinsky |first3=Alexander |last4=Fortier |first4=Daniel Costa |last5=Kamino |first5=Luciana Hiromi Yoshino |last6=Cozzuol |first6=Mario Alberto |last7=Ribeiro |first7=Adauto de Souza |last8=Vieira |first8=Fabiana Silva |date=January 2013 |title=Paleoecology and radiocarbon dating of the Pleistocene megafauna of the Brazilian Intertropical Region |url=https://www.cambridge.org/core/product/identifier/S0033589400002945/type/journal_article |journal=Quaternary Research |language=en |volume=79 |issue=1 |pages=61–65 |doi=10.1016/j.yqres.2012.09.006 |bibcode=2013QuRes..79...61D |issn=0033-5894}}</ref><ref>{{Cite journal |last1=Lopes |first1=Renato Pereira |last2=Scherer |first2=Carolina Saldanha |last3=Pereira |first3=Jamil Corrêa |last4=Dillenburg |first4=Sérgio Rebello |date=July 2023 |title=Paleoenvironmental changes in the Brazilian Pampa based on carbon and oxygen stable isotope analysis of Pleistocene camelid tooth enamel |url=https://onlinelibrary.wiley.com/doi/10.1002/jqs.3502 |journal=Journal of Quaternary Science |language=en |volume=38 |issue=5 |pages=702–718 |doi=10.1002/jqs.3502 |bibcode=2023JQS....38..702L |issn=0267-8179}}</ref> in some areas like the southwestern Brazilian Amazon, it is suggested to have inhabited [[woodland]].<ref>{{Cite journal |last1=Asevedo |first1=Lidiane |last2=Ranzi |first2=Alceu |last3=Kalliola |first3=Risto |last4=Pärssinen |first4=Martti |last5=Ruokolainen |first5=Kalle |last6=Cozzuol |first6=Mário Alberto |last7=Nascimento |first7=Ednair Rodrigues do |last8=Negri |first8=Francisco Ricardo |last9=Souza-Filho |first9=Jonas P. |last10=Cherkinsky |first10=Alexander |last11=Trindade Dantas |first11=Mário André |date=January 2021 |title=Isotopic paleoecology (δ13C, δ18O) of late Quaternary herbivorous mammal assemblages from southwestern Amazon |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379120306624 |journal=Quaternary Science Reviews |language=en |volume=251 |pages=106700 |doi=10.1016/j.quascirev.2020.106700}}</ref><ref>{{Cite journal |last1=Chahud |first1=Artur |last2=de Oliveira Costa |first2=Paulo Ricardo |last3=Ferreira Figueiredo |first3=Gisele |last4=Okumura |first4=Mercedes |date=January 2023 |title=Quaternary ungulates of the Abismo Ponta de Flecha Cave, Ribeira of Iguape Valley, Southeast Brazil: Zooarchaeological and Paleoenvironmental aspects |url=https://linkinghub.elsevier.com/retrieve/pii/S0895981122003935 |journal=Journal of South American Earth Sciences |language=en |volume=121 |pages=104107 |doi=10.1016/j.jsames.2022.104107|bibcode=2023JSAES.12104107C }}</ref>
Like living animals of similar size, it has been suggested that ''Toxodon'' probably only gave birth to a single offspring at a time.<ref>{{Cite journal |last=Andrea |first=Elissamburu |date=July 2016 |title=Prediction of offspring in extant and extinct mammals to add light on paleoecology and evolution |url=https://linkinghub.elsevier.com/retrieve/pii/S0031018216300542 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=453 |pages=73–79 |doi=10.1016/j.palaeo.2016.03.033|bibcode=2016PPP...453...73A |hdl=11336/54314 |hdl-access=free }}</ref>
''T. platensis'' bones have been found displaying signs of disease like [[osteomyelitis]] and [[Spondyloarthropathy|spondyloarthropathies]].<ref>{{Cite journal |last1=Luna |first1=Carlos A. |last2=De S. Barbosa |first2=Fernando H. |last3=Gonzalez |first3=Romina |last4=Miño-Boilini |first4=Ángel R. |last5=Repetto |first5=Carolina |last6=Zurita |first6=Alfredo E. |date=7 February 2024 |title=Bone diseases in a Pleistocene South American native ungulate species: the case of Toxodon platensis Owen, 1837 (Mammalia, Notoungulata, Toxodontidae) |url=https://onlinelibrary.wiley.com/doi/10.1002/jqs.3601 |journal=[[Journal of Quaternary Science]] |volume=39 |issue=8 |pages=1206–1215 |language=en |doi=10.1002/jqs.3601 |issn=0267-8179 |access-date=28 March 2024 |via=Wiley Online Library}}</ref>
Tracks probably attributable to ''Toxodon'' have been reported from eastern [[Pernambuco]] in Northeast Brazil.<ref name=":6" />
Isotopic analysis suggests that ''Toxodon'' may have been predated upon by the large sabertooth cat ''[[Smilodon populator]]'', the apex predator of South American ecosystems during much of the Pleistocene.<ref>{{Cite journal |last1=Dantas |first1=Mário André Trindade |last2=Cherkinsky |first2=Alexander |last3=Lessa |first3=Carlos Micael Bonfim |last4=Santos |first4=Luciano Vilaboim |last5=Cozzuol |first5=Mario Alberto |last6=Omena |first6=Érica Cavalcante |last7=Silva |first7=Jorge Luiz Lopes |last8=Sial |first8=Alcides Nóbrega |last9=Bocherens |first9=Hervé |date=2020-07-14 |title=Isotopic paleoecology (δ13C, δ18O) of a late Pleistocene vertebrate community from the Brazilian Intertropical Region |journal=Revista Brasileira de Paleontologia |language=en |volume=23 |issue=2 |pages=138–152 |doi=10.4072/rbp.2020.2.05 |issn=2236-1715 |doi-access=free}}</ref>
== Extinction ==
''Toxodon'' became extinct at the end of the Late Pleistocene around 12,000 years as part of the [[Late Pleistocene extinctions|end-Pleistocene extinction event]] alongside almost all other large animals in South America. Previous mid-Holocene dates are now thought to be in error.<ref>{{cite journal |vauthors=Politis GG, Messineo PG, Stafford TW, Lindsey EL |date=March 2019 |title=Campo Laborde: A Late Pleistocene giant ground sloth kill and butchering site in the Pampas |journal=[[Science Advances]] |volume=5 |issue=3 |pages=eaau4546 |bibcode=2019SciA....5.4546P |doi=10.1126/sciadv.aau4546 |pmc=6402857 |pmid=30854426}}</ref> These extinctions followed the [[Peopling of the Americas|first arrival of humans in the Americas]], and it has been suggested human hunting may have been a casual factor in the extinctions.<ref name=":2" /> Several sites record apparent interactions between ''Toxodon'' and humans. Remains of ''Toxodon'' from the Arroyo Seco 2 site in the Pampas are associated with unambiguously butchered megafauna, but it is unclear if the ''Toxodon'' itself was actually butchered or the remains were naturally transported to the site.<ref>{{cite journal |vauthors=Politis GG, Gutiérrez MA, Rafuse DJ, Blasi A |date=28 September 2016 |title=The Arrival of Homo sapiens into the Southern Cone at 14,000 Years Ago |journal=[[PLOS ONE]] |volume=11 |issue=9 |pages=e0162870 |bibcode=2016PLoSO..1162870P |doi=10.1371/journal.pone.0162870 |pmc=5040268 |pmid=27683248 |doi-access=free |veditors=Petraglia MD}}</ref> At the Paso Otero 5 site in the Pampas of northeast Argentina, burned bones of ''Toxodon'' alongside those of numerous other extinct megafauna species are associated with [[Fishtail projectile point|Fishtail points]] (a type of knapped stone spear point common across South America at the end of the Pleistocene, suggested to be used to hunt large mammals<ref name=":23">{{Cite journal |last1=Prates |first1=Luciano |last2=Rivero |first2=Diego |last3=Perez |first3=S. Ivan |date=2022-10-25 |title=Changes in projectile design and size of prey reveal the central role of Fishtail points in megafauna hunting in South America |journal=Scientific Reports |language=en |volume=12 |issue=1 |page=16964 |bibcode=2022NatSR..1216964P |doi=10.1038/s41598-022-21287-0 |issn=2045-2322 |pmc=9596454 |pmid=36284118}}</ref>). The bones of the megafauna were probably deliberately burned as fuel. No cut marks are visible on the vast majority of bones at the site (with only one bone of a llama possibly displaying any butchery marks), which may be due to the burning degrading the bones.<ref>G. Martínez, M. A. Gutiérrez, [https://www.researchgate.net/publication/255171250_Paso_Otero_5_a_summary_of_the_interdisciplinary_lines_of_evidence_for_reconstructing_early_human_occupation_and_paleoenvironment_in_the_Pampean_region_Argentina Paso Otero 5: A summary of the interdisciplinary lines of evidence for reconstructing early human occupation and paleoenvironment in the Pampean region, Argentina], in ''Peuplements et Préhistoire de l’Amérique'', D. Vialou, Ed. (Muséum National d’ Histoire Naturelle. Departement de Prehistoire, U.M.R, Paris, 2011), pp. 271–284.</ref> Various remains of ''Toxodon platensis'' in the collection of the [[Museum national d'Histoire naturelle|''Museum national d'Histoire naturelle'']] collected from the Pampas region in the 19th century including a femur, an [[Pelvis|iliac]] fragment, a tibia, as well as a mandible (the latter of which has been radiocarbon dated to around 13,000 years ago), have been found to display cut marks indicative of butchery.<ref>{{Cite journal |last=Toledo |first=Marcelo Javier |date=April 2023 |title=Anthropic modifications on megafauna bones in the paleontological collections of the Museum national d'Histoire naturelle de Paris: Historical aspects and implications for the Pampean Pleistocene peopling |url=https://linkinghub.elsevier.com/retrieve/pii/S0003552123000195 |journal=L'Anthropologie |language=en |volume=127 |issue=2 |pages=103134 |doi=10.1016/j.anthro.2023.103134}}</ref>
== References ==
{{Reflist}}
== Further reading ==
{{commons category|Toxodon|lcfirst=yes}}
{{wikispecies|toxodon}}
{{refbegin}}
* {{cite book | first1 = Barry | last1 = Cox | first2 = Colin | last2 = Harrison | first3 = R.J.G. | last3 = Savage | first4 = Brian | last4 = Gardiner | name-list-style = vanc | title = The Simon & Schuster Encyclopedia of Dinosaurs and Prehistoric Creatures: A Visual Who's Who of Prehistoric Life | url = https://archive.org/details/simonschusterenc00coxb | url-access = registration | publisher = Simon & Schuster | date = October 1999 | isbn = 978-0-684-86411-2 }}
{{refend}}
{{Meridiungulata|N.|state=collapsed}}
{{Taxonbar|from=Q131467}}
[[Category:Toxodonts]]
[[Category:Miocene genus first appearances]]
[[Category:Holocene extinctions]]
[[Category:Miocene mammals of South America]]
[[Category:Pliocene mammals of South America]]
[[Category:Pleistocene mammals of South America]]
[[Category:Lujanian]]
[[Category:Ensenadan]]
[[Category:Uquian]]
[[Category:Chapadmalalan]]
[[Category:Montehermosan]]
[[Category:Huayquerian]]
[[Category:Chasicoan]]
[[Category:Mayoan]]
[[Category:Neogene Argentina]]
[[Category:Ituzaingó Formation]]
[[Category:Sopas Formation]]
[[Category:Pleistocene Argentina]]
[[Category:Pleistocene Bolivia]]
[[Category:Pleistocene Brazil]]
[[Category:Pleistocene Paraguay]]
[[Category:Dolores Formation, Uruguay]]
[[Category:Fossils of Argentina]]
[[Category:Fossils of Bolivia]]
[[Category:Fossils of Brazil]]
[[Category:Fossils of Paraguay]]
[[Category:Fossils of Uruguay]]
[[Category:Fossil taxa described in 1837]]
[[Category:Taxa named by Richard Owen]]
[[Category:Prehistoric placental genera]]