Toxodon | |
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Skeleton of Toxodon in Buenos Aires | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | †Notoungulata |
Family: | †Toxodontidae |
Subfamily: | †Toxodontinae |
Genus: | †Toxodon Owen, 1837 |
Type species | |
†Toxodon platensis Owen, 1837
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Other species | |
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Synonyms | |
Genus-level
T. platensis
T. burmeisteri
T. chapalmalensis
T. ensenadensis
T. gracilis
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Toxodon (meaning "bow tooth" in reference to the curvature of the teeth) is an extinct genus of large ungulate native to South America from the Late Miocene to early Holocene epochs (Mayoan to Lujanian in the SALMA classification) (about 11.6 million to 11,000 years ago).[1][2] 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.[3] Toxodon is a member of the family Toxodontidae, which includes medium to large sized herbivores.[4] Toxodon was one of the largest members of Toxodontidae and Notoungulata, with Toxodon platensis having an estimated body mass of 1,000–1,200 kilograms (2,200–2,600 lb).[5]
Toxodon has been found across much of South America, excluding southern Patagonia, the Andes and northeastern-most region of the continent.[6] Evidence suggests that Toxodon was ecologically plastic and able to adapt its diet to local conditions.[7]
Toxodon became extinct as part of the end-Pleistocene extinctions around 12,000 years ago, along with most large mammals across the Americas.[3]
Taxonomy
[edit]Charles Darwin was one of the first to collect Toxodon fossils, after paying 18 pence for a T. platensis skull from a farmer in Uruguay.[8] In The Voyage of the Beagle Darwin wrote, "November 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."[9] 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,
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 Mr. Owen states, proves indisputably that it was intimately related to the 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 different Orders, at the present time so well separated, blended together in different points of the structure of the Toxodon!
Toxodon was described in 1837 by Richard Owen based on remains collected by Darwin.[10]
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, prior to the arrival of living ungulates in South America around 2.5 million years ago as part of the Great American Interchange.[3] Notoungulata represents the most diverse group of indigenous South American ungulates, with over 150 described genera in 13 different families.[11] Notoungulates are morphologically diverse, including forms morphologically distant from Toxodon such as rodent and rabbit-like forms. Toxodon was a member of Toxodontidae, a large bodied group including vaguely rhinoceros like forms.[3]
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 perissodactyls (which contains equids, rhinoceroses and tapirs) as part of the clade Panperissodactyla, making them true ungulates.[12][13] 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 with perissodactyls.[14]
Evolution
[edit]The species Toxodon chapalmalensis is known from the Pliocene (Montehermosan-Chapadmalalan) of Argentina,[15] while Toxodon platensis, the type species, is known from the Pleistocene.
In 2014, a study identifying a new species of toxodontid resolved the phylogenetic relations of the toxodontids, including to Toxodon. The below cladogram was found by the study:[16]
†Notoungulata |
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Description
[edit]![](https://upload.wikimedia.org/wikipedia/commons/thumb/5/5a/ToxodonMLP1_%28cropped%29.jpg/220px-ToxodonMLP1_%28cropped%29.jpg)
Toxodon was about 2.7 m (8 ft 10 in) in body length, with an estimated weight up to 1,415 kg (3,120 lb)[17] and about 1.5 m (4 ft 11 in) high at the shoulder and resembled a heavy rhinoceros, with a short and vaguely hippopotamus-like head.[18] Because of the position of its nasal openings, it is believed that Toxodon had a well-developed snout.[citation needed] Toxodon possessed a large, barrel shaped body. It had short stout legs with plantigrade feet with three functional relatively short toes. The hind limbs are longer and raised higher than the front limbs, giving a sloped appearance to the body.[19] Like horses, it had a stay apparatus allowing the knees to be passively locked while standing.[20]
The vertebrae were equipped with high apophyses, which most likely supported the massive weight and muscles as well as its powerful head.[citation needed] Toxodon had broad jaws which were filled with bow shaped teeth and incisors.[3] The molar teeth of Toxodon have no roots and are ever-growing (euhypsodont), like the incisors of rodents and lagomorphs, and often exhibit enamel hypoplasia.[21]
Palaeobiology
[edit]Toxodon is suggested to have been capable of running at considerable speed.[22] Toxodon is believed to have been ecologically plastic and have had a wide niche breadth,[7] with its diet varying according to local conditions,[23] with an almost totally C3 browsing diet in the Amazon rainforest, mixed feeding C3 in Bahia and the Pampas to almost completely C4 dominated grazing diet in the Chaco.[24] Within the Brazilian Intertropical Region, local climate had little impact on the diet of T. platensis.[25]
T. platensis specimens have been found displaying osteological signs of osteomyelitis and spondyloarthropathies.[26]
Extinction
[edit]Toxodon became extinct at the end of the Late Pleistocene around 12,000 years as part of the Quaternary extinction event, alongside almost all other large animals in South America. Previous mid-Holocene dates are now thought to be in error.[27] Remains from the Arroyo Seco 2 site in the Pampas are associated with butchered megafauna, but it is unclear if the Toxodon itself was actually butchered or the remains were naturally transported to the site.[28]
![](https://upload.wikimedia.org/wikipedia/commons/thumb/7/71/Toxodon_platensis.jpg/220px-Toxodon_platensis.jpg)
![](https://upload.wikimedia.org/wikipedia/commons/thumb/3/37/Toxodon_skull.jpg/220px-Toxodon_skull.jpg)
Distribution
[edit]Toxodon had a wide distribution in South America during the Late Pleistocene, extending from the Pampas into the Amazon rainforest.
Fossils of Toxodon have been found in:[2][29]
- Holocene
- Pleistocene
- San José, Fortin Tres Pozos, Chaco and Luján Formations, Argentina
- Tarija and Ñuapua Formations, Bolivia
- Brazil
- Paraguay
- Sopas and Dolores Formations, Uruguay
- Miocene-Pliocene (Montehermosan)
- Miocene
- Ituzaingó Formation, then described as Entrerriana Formation, Argentina
References
[edit]- ^ Baffa O, Brunetti A, Karmann I, Neto CM (May 2000). "ESR dating of a toxodon tooth from a Brazilian karstic cave". Applied Radiation and Isotopes. 52 (5): 1345–9. Bibcode:2000AppRI..52.1345B. doi:10.1016/S0969-8043(00)00093-2. PMID 10836452.
- ^ a b Turvey ST (28 May 2009). Holocene Extinctions. OUP Oxford. ISBN 978-0-19-157998-1.
- ^ a b c d e Croft DA, Gelfo JN, López GM (30 May 2020). "Splendid Innovation: The Extinct South American Native Ungulates". Annual Review of Earth and Planetary Sciences. 48 (1): 259–290. Bibcode:2020AREPS..48..259C. doi:10.1146/annurev-earth-072619-060126. ISSN 0084-6597. S2CID 213737574. Retrieved 2 January 2024.
- ^ Cassini, Guillermo H.; Flores, David A.; Vizcaíno, Sergio F. (July 2012). "Postnatal ontogenetic scaling of Nesodontine (Notoungulata, Toxodontidae) cranial morphology: Nesodontine cranial allometry". Acta Zoologica. 93 (3): 249–259. doi:10.1111/j.1463-6395.2011.00501.x. hdl:11336/81335.
- ^ Nelson, Allison; Engelman, Russell K.; Croft, Darin A. (12 July 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. S2CID 259866522.
- ^ Cione, Alberto Luis; Gasparini, Germán Mariano; Soibelzon, Esteban; Soibelzon, Leopoldo Héctor; Tonni, Eduardo Pedro (2015), "The GABI in Southern South America", The Great American Biotic Interchange, Dordrecht: Springer Netherlands, pp. 71–96, doi:10.1007/978-94-017-9792-4_3, ISBN 978-94-017-9791-7, S2CID 127856682, retrieved 30 January 2024
- ^ a b Dantas, Mário André Trindade; Cherkinsky, Alexander; Bocherens, Hervé; Drefahl, Morgana; Bernardes, Camila; França, Lucas de Melo (15 August 2017). "Isotopic paleoecology of the Pleistocene megamammals from the Brazilian Intertropical Region: Feeding ecology (δ13C), niche breadth and overlap". Quaternary Science Reviews. 170: 152–163. doi:10.1016/j.quascirev.2017.06.030. ISSN 0277-3791. Retrieved 2 January 2024 – via Elsevier Science Direct.
- ^ Quammen, D. (February 2009). "Darwin's first clues". National Geographic. p. 45.
- ^ Darwin, Charles (1997) [1839]. Browne, J.; Neve, M. (eds.). The Voyage of the Beagle. ISBN 978-0-14-043268-8.
Read, 19th April 1837. A detailed account will appear in the first part of the zoology of Voyage of the Beagle.
- ^ 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 https://revista.geologica.org.ar/raga/article/view/1339
- ^ Rezende Castro, Luis Otavio; García-López, Daniel A.; Bergqvist, Lilian Paglarelli; De Araújo-Júnior, Hermínio Ismael (30 June 2021). "A New Basal Notoungulate from the Itaboraí Basin (Paleogene) of Brazil". Ameghiniana. 58 (3). doi:10.5710/AMGH.05.02.2021.3387. ISSN 0002-7014. S2CID 234220780.
- ^ Welker F, Collins MJ, Thomas JA, Wadsley M, Brace S, Cappellini E, et al. (June 2015). "Ancient proteins resolve the evolutionary history of Darwin's South American ungulates". Nature. 522 (7554): 81–4. Bibcode:2015Natur.522...81W. doi:10.1038/nature14249. hdl:11336/14769. PMID 25799987. S2CID 4467386.
- ^ Buckley, M. (7 May 2015). "Ancient collagen reveals evolutionary history of the endemic South American 'ungulates'". Proceedings of the Royal Society B: Biological Sciences. 282 (1806): 20142671. doi:10.1098/rspb.2014.2671. PMC 4426609. PMID 25833851.
- ^ Westbury M, Baleka S, Barlow A, Hartmann S, Paijmans JL, Kramarz A, et al. (June 2017). "A mitogenomic timetree for Darwin's enigmatic South American mammal Macrauchenia patachonica". Nature Communications. 8: 15951. Bibcode:2017NatCo...815951W. doi:10.1038/ncomms15951. PMC 5490259. PMID 28654082.
- ^ Tomassini, Rodrigo L.; Montalvo, Claudia I.; Deschamps, Cecilia M.; Manera, Teresa (December 2013). "Biostratigraphy and biochronology of the Monte Hermoso Formation (early Pliocene) at its type locality, Buenos Aires Province, Argentina". Journal of South American Earth Sciences. 48: 31–42. Bibcode:2013JSAES..48...31T. doi:10.1016/j.jsames.2013.08.002. hdl:11336/21606. ISSN 0895-9811. Retrieved 2 January 2024 – via Elsevier Science Direct.
- ^ Forasiepi AM, Cerdeno E, Bond M, Schmidt GI, Naipauer M, Straehl FR, et al. (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.
- ^ Fariña RA, Czerwonogora A, di Giacomo M (March 2014). "Splendid oddness: revisiting the curious trophic relationships of South American Pleistocene mammals and their abundance". Anais da Academia Brasileira de Ciências. 86 (1): 311–31. doi:10.1590/0001-3765201420120010. PMID 24676170.
- ^ Fariña RA, Vizcaíno SF, de Iuliis G (2012). Megafauna: Giant Beasts of Pleistocene South America. Indiana University Press. ISBN 978-0-253-00230-3.
- ^ Defler T (2019), "The Native Ungulates of South America (Condylarthra and Meridiungulata)", History of Terrestrial Mammals in South America, Topics in Geobiology, vol. 42, Cham: Springer International Publishing, p. 102, doi:10.1007/978-3-319-98449-0_5, ISBN 978-3-319-98448-3, S2CID 91879648, retrieved 10 July 2020
- ^ Shockey BJ. 2001. "Specialized knee joints in some extinct, endemic, South American herbivores" Acta Palaeontologica Polonica 46:277–88
- ^ Braunn PR, Ribeiro AM, Ferigolo J (July 2014). "Microstructural defects and enamel hypoplasia in teeth of Toxodon Owen, 1837 from the Pleistocene of Southern Brazil". Lethaia. 47 (3): 418–431. Bibcode:2014Letha..47..418B. doi:10.1111/let.12063.
- ^ S.F. Vizcaino, R.A. Farina, J.C. Fernicola "Young Darwin and the ecology and extinction of Pleistocene South American fossil mammals" Revista de la Asociacion Geologica Argentina, 64 (2009), pp. 160-169
- ^ Pansani TR, Muniz FP, Cherkinsky A, Pacheco ML, Dantas MA (October 2019). "Isotopic paleoecology (δ13C, δ18O) of Late Quaternary megafauna from Mato Grosso do Sul and Bahia States, Brazil". Quaternary Science Reviews. 221: 105864. Bibcode:2019QSRv..22105864P. doi:10.1016/j.quascirev.2019.105864. S2CID 202200336.
- ^ MacFadden BJ (September 2005). "Diet and habitat of toxodont megaherbivores (Mammalia, Notoungulata) from the late Quaternary of South and Central America". Quaternary Research. 64 (2): 113–124. Bibcode:2005QuRes..64..113M. doi:10.1016/j.yqres.2005.05.003. Retrieved 2 January 2024 – via Elsevier Science Direct.
- ^ Gomes, Verônica Santos; Lessa, Carlos Micael Bonfim; Oliveira, Gustavo Ribeiro de; Bantim, Renan Alfredo Machado; Sayão, Juliana; Bocherens, Hervé; Araújo-Júnior, Hermínio Ismael de; Dantas, Mário André Trindade (January 2023). "Seasonal variations in diet (δ13C) and climate (δ 18O) inferred through toxodonts enamel teeth during the Late Pleistocene in the brazilian intertropical region". Journal of South American Earth Sciences. 121: 104148. doi:10.1016/j.jsames.2022.104148. Retrieved 19 April 2024 – via Elsevier Science Direct.
- ^ Luna, Carlos A.; De S. Barbosa, Fernando H.; Gonzalez, Romina; Miño-Boilini, Ángel R.; Repetto, Carolina; Zurita, Alfredo E. (7 February 2024). "Bone diseases in a Pleistocene South American native ungulate species: the case of Toxodon platensis Owen, 1837 (Mammalia, Notoungulata, Toxodontidae)". Journal of Quaternary Science. doi:10.1002/jqs.3601. ISSN 0267-8179. Retrieved 28 March 2024 – via Wiley Online Library.
- ^ Politis GG, Messineo PG, Stafford TW, Lindsey EL (March 2019). "Campo Laborde: A Late Pleistocene giant ground sloth kill and butchering site in the Pampas". Science Advances. 5 (3): eaau4546. Bibcode:2019SciA....5.4546P. doi:10.1126/sciadv.aau4546. PMC 6402857. PMID 30854426.
- ^ Politis GG, Gutiérrez MA, Rafuse DJ, Blasi A (28 September 2016). Petraglia MD (ed.). "The Arrival of Homo sapiens into the Southern Cone at 14,000 Years Ago". PLOS ONE. 11 (9): e0162870. Bibcode:2016PLoSO..1162870P. doi:10.1371/journal.pone.0162870. PMC 5040268. PMID 27683248.
- ^ Toxodon at Fossilworks.org
Further reading
[edit]- Cox B, Harrison C, Savage R, Gardiner B (October 1999). The Simon & Schuster Encyclopedia of Dinosaurs and Prehistoric Creatures: A Visual Who's Who of Prehistoric Life. Simon & Schuster. ISBN 978-0-684-86411-2.
Well, that’s interesting to know that Psilotum nudum are known as whisk ferns. Psilotum nudum is the commoner species of the two. While the P. flaccidum is a rare species and is found in the tropical islands. Both the species are usually epiphytic in habit and grow upon tree ferns. These species may also be terrestrial and grow in humus or in the crevices of the rocks.
View the detailed Guide of Psilotum nudum: Detailed Study Of Psilotum Nudum (Whisk Fern), Classification, Anatomy, Reproduction