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{{Short description|Genus of large, heavily armored mammals}}
{{good article}}
{{Automatic taxobox
| fossil_range = [[Pliocene]]?-[[Pleistocene]] ([[Montehermosan]]?–[[Lujanian]])<br />~{{fossil range|3.200|0.011}}
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'''''Glyptodon''''' (
The [[type species]], ''G. clavipes'', was described in 1839 by notable British [[Paleontology|paleontologist]] [[Richard Owen|Sir Richard Owen]]. Later in the 19th century, dozens of complete skeletons were unearthed from localities and described by paleontologists such as [[Florentino Ameghino]] and [[Hermann Burmeister]]. During this era, many species of ''Glyptodon'' were dubbed, some of them based on fragmentary or isolated remains. Fossils from North America were also assigned to ''Glyptodon'', but all of them have since been placed in the closely related genus ''[[Glyptotherium]]''. It was not until the later end of the 1900s and 21st century that full review of the genus came about, restricting ''Glyptodon'' to just five species under one genus.
Glyptodonts were typically large,
Glyptodonts existed for millions of years, though ''Glyptodon'' itself was one its last surviving members. ''Glyptodon'' was one of many South American [[megafauna]], with many native groups such as [[Notoungulata|notoungulates]] and ground sloths reaching immense sizes. ''Glyptodon'' had a mixed diet of grasses and other plants, instead living at the edge forests and grasslands where the shrubbery was lower. ''Glyptodon'' had a wide muzzle, an adaptation for bulk feeding. The armor could have protected the animal from predators, of which many coexisted with ''Glyptodon'', including the "saber-tooth cat" ''[[Smilodon]]'', the large [[Canidae|canid]] ''[[Protocyon]]'', and the giant bear ''[[Arctotherium]]''.
''Glyptodon'', along with all other glyptodonts, became extinct at the end of the Late Pleistocene, around 12,000 as part of the [[Late Pleistocene extinctions]], along with most large mammals in the Americas. Evidence of hunting of glyptodonts by recently arrived [[Paleo-Indians|Paleoindians]] suggests that humans may have been a causal factor in the extinctions.
== History ==
=== Confusion with ''Megatherium'' ===
[[File:Georges_Cuvier.jpg|left|thumb|224x224px|[[Georges Cuvier]] (
The history and taxonomy of ''Glyptodon'' is storied and convoluted, as it involved confusion with other genera and [[Nomen dubium|dubious]] species, as well as a lack of detailed data. The first recorded discovery of ''Glyptodon'' was as early as 1814 when Uruguayan priest, scientist, soldier, and later politician [[Dámaso Antonio Larrañaga]] (
Larrañaga wrote to French scientist [[Augustin Saint-Hilaire|Auguste Saint Hilaire]] about the discovery, and the letter was reproduced by Cuvier in 1823 in the second volume of his landmark book ''Recherches sur les ossemens fossiles''.<ref name="Cuvier-1823">{{Cite book |last=Cuvier |first=Georges |url=https://books.google.com/books?id=xqZAAAAAcAAJ&dq=Cuvier,+G.+1824.+Recherches+sur+les+Ossemens+Fossiles,+nouvelle+edition.+G.+Dufour+et+E.+D'Ocagne,+Paris,+547+pp.&pg=PA1 |title=Recherches sur les ossemens fossiles |date=1823 |publisher=chez G. Dufour et E. d'Ocagne |language=fr}}</ref> Larrañaga also noted that similar fossils had been found in "analogous strata near Lake Merrim, on the frontier of the Portuguese colonies ([[South Region, Brazil|southern Brazil]])."<ref name="Cuvier-1823" /><ref name="Owen-1855" /> These fossils were also likely those of glyptodontines, possibly the closely related ''[[Hoplophorus]]''.<ref name="Porpino-2010">{{Cite journal |last1=Porpino |first1=Kleberson de O. |last2=Fernicola |first2=Juan C. |last3=Bergqvist |first3=Lílian P. |date=2010-05-18 |title=Revisiting the intertropical Brazilian species Hoplophorus euphractus (Cingulata, Glyptodontoidea) and the phylogenetic affinities of Hoplophorus
Another work on the armored ''Megatherium'' hypothesis was published in 1833 by Berlin scientist E. D'Alton, who described more of the material sent by Sellow, including portions of the limbs, [[Manus (anatomy)|manus]], and [[shoulder girdle]]. D'Alton recognized the great similarities of the fossils to ''Dasypus'' and speculated that it was a giant armadillo, contrary to the notion that they were from ''Megatherium''. Despite this, D'Alton did not erect a new name for the fossils and instead wrote that additional material was necessary to distinguish it from other armadillos. D'Alton did not mention ''Megatherium'' or its osteoderms in the paper, but he implied that all of the "''Megatherium'' armor" was instead from his armadillo. This hypothesis was supported by Laurillard in 1836, who mentioned that a plaster cast of a large armadillo carapace represented a distinct taxon from ''Megatherium'' and that the armor referred to the sloth was instead from an armadillo.<ref name="Huxley-1865" />
1837 saw the naming of the first glyptodontine, ''Hoplophorus euphractus'', when Danish paleontologist [[Peter Wilhelm Lund]] published a series of memoirs on the fossils of [[Lagoa Santa, Minas Gerais|Lagoa Santa]] in Brazil, dating to the Pleistocene.<ref>Lund, P. W. (1837). ''Blik paa Brasiliens dyreverden foÈr sidste jordomvaeltning''. Popp.</ref><ref name="Porpino-2010" /> The fossils included osteoderms comparable to those described earlier by Larrañaga, as well as teeth, skull fragments, limb bones, and other elements.<ref name="Huxley-1865" /><ref name="Porpino-2010" /> After 1837, several new genera and species of glyptodontines were named in quick succession by European paleontologists: ''Chlamydotherium'' based on Sellow's carapace and ''Orycterotherium'' based on Sellow's femur were named by German scientist H. G. Bronn 1838,<ref>Bronn, H. G. (1858). ''Untersuchungen über die Entwickelungs-Gesetze der organischen Welt während der Bildungs-Zeit unserer Erd-Oberfläche''. E. Schweizerbart.</ref> ''Pachypus'' by Eduard D'Alton in 1839 based on Sellow's 1833 material,<ref name="Huxley-1865" /> ''Neothoracophorus'' (originally ''Thoracophorus'' but the name was preoccupied by a [[Thoracophorus|beetle]]) in 1889 by Argentine paleontologist [[Florentino Ameghino]]<ref name="Ameghino">Ameghino, F. (1889). ''Contribucion al conocimiento de los mamiferos fosiles de la República Argentina: Obra escrita bajo los auspicios de la Academia nacional de ciencias de la República Argentina para ser presentada á la Exposicion universal de Paris de 1889'' (Vol. 6). PE Coni é hijos.</ref> based on isolated osteoderms now identified as those of a juvenile ''Glyptodon'' from Patagonia,<ref name="Zurita">Zurita, A. E., Miño-Boilini, Á. R., Soibelzon, E., Carlini, A. A., & Paredes Rios, F. (2009). The diversity of Glyptodontidae (Xenarthra, Cingulata) in the Tarjia Valley (Bolivia): Systematic, biostratigraphic and paleobiogeographic aspects of a particular assemblage.(With 3 figures and 1 table). ''Neues Jahrbuch fur Geologie und Palaontologie-Abhandlungen'', ''251''(2), 225.</ref> and ''Lepitherium'' in 1839 by [[Isidore Geoffroy Saint-Hilaire|Geoffroy Saint-Hilaire]] based on Sellow's osteoderms.<ref name="Podgorny-2013">{{Cite journal |last=Podgorny |first=Irina |date=2013 |title=Fossil dealers, the practices of comparative anatomy and British diplomacy in Latin America, 1820–1840 |url=https://www.cambridge.org/core/journals/british-journal-for-the-history-of-science/article/abs/fossil-dealers-the-practices-of-comparative-anatomy-and-british-diplomacy-in-latin-america-18201840/19DC25F459111A23072A858372618262 |journal=The British Journal for the History of Science |language=en |volume=46 |issue=4 |pages=647–674 |doi=10.1017/S0007087412000702 |s2cid=147258227 |issn=0007-0874}}</ref><ref name="Saint">Saint-Hilaire, E. G. (1831). ''Recherches sur de grands sauriens: trouvés à l'état fossile vers les confins maritimes de la basse normandie, attribués d'abord au crocodile, puis déterminés sous les noms de téléosaurus et sténéosaurus''. Firmin Didot.</ref> Saint-Hillaire considered the osteoderms found by Sellow to not even be mammal, but instead of a relative of ''[[Teleosaurus]]'', a crocodile-like reptile known from [[Jurassic]] deposits in France.<ref name="Saint" /><ref name="Podgorny-2013" />
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=== Richard Owen and referred species ===
[[File:Glyptodon_Owen_1839.png|left|thumb|[[Richard Owen]]'s 1838 reconstruction of a ''Glyptodon'' skeleton; with the namesake tooth on the right|alt=Richard Owen's reconstruction of Glyptodon from 1838.|250x250px]]
In 1838, British diplomat [[Woodbine Parish|Sir Woodbine Parish]] (1796–1882) was sent an isolated [[Molar (tooth)|molariform]] and a letter about the discovery of several large fossils from the [[Matanza River]] in [[Buenos Aires Province|Buenos Aires]], Argentina that dated to the Pleistocene.<ref name="Owen">Owen, R. (1841). VI.—Description of a Tooth and Part of the Skeleton of the Glyptodon clavipes, a large Quadruped of the Edentate Order, to which belongs the Tesselated Bony Armour described and figured by Mr. Clift in the former Volume of the Transactions of the Geological Society; with a consideration of the question whether the Megatherium possessed an analogous Dermal Armour. ''Transactions of the Geological Society of London'', ''6''(1), 81-106.</ref><ref name="Cuadrelli-2018">{{Cite journal |last1=Cuadrelli |first1=Francisco |last2=Zurita |first2=Alfredo E. |last3=Toriño |first3=Pablo |last4=Miño-Boilini |first4=Ángel R. |last5=Rodríguez-Bualó |first5=Santiago |last6=Perea |first6=Daniel |last7=Acuña Suárez |first7=Gabriel E. |date=2018-09-03 |title=Late Pleistocene Glyptodontinae (Mammalia, Xenarthra, Glyptodontidae) from southern South America: a comprehensive review
Cuadrelli ''et al'' (2018) designated the species a ''[[species inquirenda]]'' due to this issue and commented that more analyses are necessary.<ref name="Cuadrelli-2018" /> In 1860, Signor Maximo Terrero collected a partial skeleton, including a skull and carapace, of ''G. clavipes'' from the [[Salado River (Argentina)|River Salado]] in southern Buenos Aires and dated to the Pleistocene. These fossils were also sent to the Royal College of Surgeons, where they were described in detail by British paleontologist [[Thomas Henry Huxley]] (1825–1895) in 1865 during a comprehensive review of the taxon.<ref name="Huxley-1865" /> This skeleton was also destroyed during WWII, but Huxley published several illustrations that presented great amounts of new information on the taxon.<ref name="Huxley-1865" /><ref name=":1">{{Cite journal |last=Huxley |first=Thomas Henry |date=1862-01-01 |title=Description of a New Specimen of Glyptodon, Recently Acquired by the Royal College of Surgeons of England |journal=Proceedings of the Royal Society of London |volume=12 |pages=316–326 |doi=10.1098/rspl.1862.0071 |jstor=112260 |doi-access=free}}</ref>
Later in 1845, many more fossils found by Parish from Pleistocene layers in Argentina were named as new species of ''Glyptodon'' by Owen: ''G. ornatus, G. reticulatus, G. tuberculatus'', and ''G. clavicaudatus'' in 1847. Of these additional species, only ''G. reticulatus'' is still considered a valid species of ''Glyptodon'' as ''G. ornatus'' was reassigned to the genus ''[[Neosclerocalyptus]],''<ref>Paula Couto, C. D. (1957). Sôbre um gliptodonte do Brasil. ''Boletim Divisão de Geologia e Mineralogia'', ''165'', 1-37.</ref> ''G. tuberculatus'' to ''Panochthus'',<ref name="Burmeister-2">Burmeister, G. (1866). Lista de los mamíferos fósiles del terreno diluviano. In ''Anales del Museo Público de Buenos Aires'' (Vol. 1, No. 3, pp. 121-232).</ref> and ''G. clavicaudatus'' to ''[[Doedicurus]]''.<ref>Burmeister, G. Burmeister 1870–1874. ''Monografia de los glyptodontes en el Museo Público de Buenos Aires. Anales del Museo Público de Buenos Aires'', ''2'', 1-412.</ref> ''G. reticulatus'' was named on the basis of several carapace fragments that had also been recovered from the Matanza River, but they lack detailed locality information and the fossils too were destroyed during WWII. The fragments were cast by the NHMUK as well, being used to diagnose the species.<ref name="Zurita-2018">{{Cite journal |last1=Zurita |first1=Alfredo Eduardo |last2=Gillette |first2=David D. |last3=Cuadrelli |first3=Francisco |last4=Carlini |first4=Alfredo Armando |date=2018-06-01 |title=A tale of two clades: Comparative study of Glyptodon Owen and Glyptotherium Osborn (Xenarthra, Cingulata, Glyptodontidae) |url=https://www.sciencedirect.com/science/article/pii/S0016699517301870 |journal=Geobios |language=en |volume=51 |issue=3 |pages=247–258 |doi=10.1016/j.geobios.2018.04.004 |bibcode=2018Geobi..51..247Z |hdl=11336/83593 |s2cid=134450624 |issn=0016-6995}}</ref>
Other paleontologists also started erecting names for ''Glyptodon'' species after the 1840s, but many of them are now seen as dubious, species inquirenda, or synonymous with previously named species.<ref name="Cuadrelli-2020">{{Cite journal |last1=Cuadrelli |first1=Francisco |last2=Zurita |first2=Alfredo E. |last3=Toriño |first3=Pablo |last4=Miño-Boilini |first4=Ángel R. |last5=Perea |first5=Daniel |last6=Luna |first6=Carlos A. |last7=Gillette |first7=David D. |last8=Medina |first8=Omar |date=2020-09-16 |title=A new species of glyptodontine (Mammalia, Xenarthra, Glyptodontidae) from the Quaternary of the Eastern Cordillera, Bolivia: phylogeny and palaeobiogeography
▲Other paleontologists also started erecting names for ''Glyptodon'' species after the 1840s, but many of them are now seen as dubious, species inquirenda, or synonymous with previously named species.<ref name="Cuadrelli-2020">{{Cite journal |last1=Cuadrelli |first1=Francisco |last2=Zurita |first2=Alfredo E. |last3=Toriño |first3=Pablo |last4=Miño-Boilini |first4=Ángel R. |last5=Perea |first5=Daniel |last6=Luna |first6=Carlos A. |last7=Gillette |first7=David D. |last8=Medina |first8=Omar |date=2020-09-16 |title=A new species of glyptodontine (Mammalia, Xenarthra, Glyptodontidae) from the Quaternary of the Eastern Cordillera, Bolivia: phylogeny and palaeobiogeography |url=https://doi.org/10.1080/14772019.2020.1784300 |journal=Journal of Systematic Palaeontology |volume=18 |issue=18 |pages=1543–1566 |doi=10.1080/14772019.2020.1784300 |s2cid=221064742 |issn=1477-2019}}</ref><ref name="Cuadrelli-2018" /> Par L. Nodot described a new genus and species of glyptodontine in 1857, ''Schistopleurum typus'', on the basis of a caudal tube found in the [[Pampas]] of Argentina, but it has since been synonymized with ''G. reticulatus''.<ref name="Nodot">Nodot, L. (1856). ''Description d'un nouveau genre d'édenté fossile renfermant plusieurs espèces voisines du glyptodon: suivie d'une nouvelle méthode de classification applicable à toute l'histoire naturelle, et spécialement à ces animaux, avec un atlas de douze planches lithographiées; ouvrage publié par l'Academie des sciences, arts et belles-lettres de Dijon et l'atlas avec le concours du Conseil municipal de même ville''. Imprimerie Loireau-Feuchot.</ref> Another species now seen as valid, ''G. munizi'', was described in 1881 by Argentine paleontologist Florentino Ameghino (1853-1911) on the basis of several osteoderms found in the Ensenadan of [[Arroyo del Medio]], [[San Nicolás de los Arroyos|San Nicolás, Argentina]].<ref name="Soibelzon-2006">{{Cite journal |last1=Soibelzon |first1=Esteban |last2=Zurita |first2=Alfredo Eduardo |last3=Carlini |first3=Alfredo A. |date=2006 |title=Glyptodon munizi Ameghino (Mammalia, Cingulata, Glyptodontidae): redescripción y anatomía |url=http://www.scielo.org.ar/scielo.php?script=sci_abstract&pid=S0002-70142006000200008&lng=es&nrm=iso&tlng=es |journal=Ameghiniana |volume=43 |issue=2 |pages=377–384 |issn=0002-7014}}</ref><ref>Ameghino, F. 1882. Catálogo de las colecciones de Antropología prehistórica y paleontología de Florentino Ameghino, Partido de Mercedes. En: ''Catálogo de la Sección de la Provincia de Buenos'' ''Aires (República Argentina). Exposición Continental Sudamericana. Anexo A'': 35-42.</ref> For many years the taxon was only known from the fragmentary holotype, but skull and complete carapace material of the species was later described in detail in 2006 that cemented its validity.<ref name="Soibelzon-2006" /><ref name="Cuadrelli-2018" /> German zoologist [[Hermann Burmeister]] described several ''Glyptodon'' fossils in the 1860s, many of them he named as new species of ''Glyptodon'' itself or the synonym ''Schistopleurum'', all of which are now synonyms of ''Glyptodon'' and its species.<ref name="Burmeister">Burmeister, H. (1866). XXXIX.—On Glyptodon and its allies. ''Annals and Magazine of Natural History'', ''18''(106), 299-304.</ref><ref name="Cuadrelli-2018" /> In 1908, Florentino Ameghino named another species of ''Glyptodon, G. chapalmalensis'', based on a carapace fragment that he had collected from the Atlantic Coast of Buenos Aires Province that dated to the [[Chapadmalalan]]. In 1932, A. Castellanos made a new genus for ''G. chapalmalensis'', ''Paraglyptodon'', which later included another species, ''P. uquiensis'', that was based on more complete specimens that had been collected from [[Uquía]], Argentina between 1909 and 1912.<ref>Castellanos, A. (1932). ''Nuevos géneros de gliptodontes en relación con su filogenia''.</ref><ref>Castellanos, A. (1953). ''Descripción de restos de" Paraglyptodon uquiensis" n. sp. de Uquía (Senador Pérez) de Jujuy'' (No. 32). la Provincia.</ref> The former species is dubious, but likely not ''Glyptodon'' based on its age.<ref name="Cruz-2016">{{Cite journal |last1=Cruz |first1=Laura E. |last2=Fernicola |first2=Juan C. |last3=Taglioretti |first3=Matias |last4=Toledo |first4=Nestor |date=2016-03-01 |title=A reassessment of the taxonomic status of Paraglyptodon Castellanos, 1932 (Mammalia, Cingulata, Glyptodontia) |url=https://www.sciencedirect.com/science/article/pii/S0895981115300924 |journal=Journal of South American Earth Sciences |language=en |volume=66 |pages=32–40 |doi=10.1016/j.jsames.2015.11.012 |bibcode=2016JSAES..66...32C |issn=0895-9811}}</ref> ''P. uquiensis'' has been synonymized with ''Glyptodon'' and is possibly a valid species, though further analysis is necessary to settle its status.<ref name="Zurita-2018" /><ref name="Cuadrelli-2018" />[[File:Hunterian_Museum_London_1842.png|thumb|The [[Hunterian Museum (London)|Hunterian Museum]] at the [[Royal College of Surgeons]] {{Circa|1842}} with a mounted skeleton of ''Glyptodon'' in the bottom right|alt=Illustration of the Hunterian Museum featuring a mounted skeleton of Glyptodon.|304x304px]]
=== Reassessment and ''Glyptotherium'' ===
In the 1950s, Argentine paleontologist [[Alfredo Castellanos]] (
Another ''Glyptodon'' species was described in 2020 called ''G. jatunkhirkhi'' by several authors led by Argentine zoologist [[Francisco Cuadrelli]] on the basis of an individual preserving a nearly complete carapace, several caudal rings, and a pelvis that had been collected from [[Yamparáez Province|Yamparaez]], {{Convert|24|km|mi}} southeast of the Bolivian city of [[Sucre]]. The strata they were found in was made up of fluvial, sandy sediments that dated to the [[Late Pleistocene]] from elevations as high as {{Convert|4100-2500|m|ft}} above sea level.<ref name="Cuadrelli-2020" /> Several additional paratypes were referred to the species from other Late Pleistocene sites in [[Cordillera Oriental (Bolivia)|Eastern Cordillera]], Bolivia including a nearly complete skull and several osteoderms.<ref name="Cuadrelli-2020" /> In a phylogenetic analysis conducted by Cuadrelli ''et al''., 2020, ''G. jatunkhirki'' was recovered as the most basal ''Glyptodon'' species, despite being the same age as the more derived species ''G. clavipes.''
Fossils from North America were also described and referred to ''Glyptodon'' starting in 1875, when civil engineers J. N. Cuatáparo and Santiago Ramírez collected a skull, nearly complete carapace, and associated postcranial skeleton of a glyptodontine from a drainage canal near [[Tequixquiac]], [[Mexico]], the fossils coming from the [[Rancholabrean|Rancholabrean stage]] of the Pleistocene.<ref>Cuatáparo, J. N., & Ramírez, S. (1875). ''Descripción de un mamífero fósil de especie desconocida perteneciente al género" Glyptodon": encontrado entre las capas post-terciarias de Tequisquiac, en el Distrito de Zumpango''. F. Diaz de Leon.</ref><ref name="Zurita-2018" /> These fossils were the first found of glyptodontines in North America and were named as a new species of ''Glyptodon'', ''G. mexicanum'', but the fossils have since been lost and the species was synonymized with ''Glyptotherium cylindricum''.<ref name="Zurita-2018" /><ref name="Gillette">{{cite journal |last1=Gillette |first1=David D. |last2=Ray |first2=Clayton E. |title=Glyptodonts of North America |journal=Smithsonian Contributions to Paleobiology |date=1981 |issue=40 |pages=1–255 |url=https://repository.si.edu/bitstream/handle/10088/1966/SCtP-0040-Lo_res.pdf?sequence=2&isAllowed=y |doi=10.5479/si.00810266.40.1}}</ref> Several other North American glyptodontine species were named throughout the late 19th-early 20th century, typically based on fragmentary osteoderms. All North American and Central American fossils of glyptodontines have since been referred to the closely related genus ''Glyptotherium'', which was named in 1903 by American paleontologist [[Henry Fairfield Osborn]].<ref>{{Cite book |last=Osborn |first=Henry Fairfield |url=https://books.google.com/books?id=BMUrAAAAYAAJ&dq=glyptotherium+texanum&pg=PA489 |title=Glyptotherium Texanum, a New Glyptodont, from the Lower Pleistocene of Texas |date=1903 |publisher=order of the Trustees, American Museum of Natural History |language=en}}</ref>
== Taxonomy ==
''Glyptodon'' is the type genus of Glyptodontinae, an extinct [[subfamily]] of large, heavily armored armadillos that first evolved in the [[Eocene|Late Eocene]] (ca. 33.5 mya) and went extinct in the [[Greenlandian|Early Holocene]] during the [[
Glyptodontinae was classified in its own family or even superfamily until in 2016, when [[ancient DNA]] was extracted from the carapace of a 12,000 year old ''Doedicurus'' specimen, and a nearly complete [[mitochondrial genome]] was reconstructed (76x [[Coverage (genetics)|coverage]]). Comparisons with those of modern armadillos revealed that glyptodonts diverged from [[Tolypeutinae|tolypeutine]] and [[Chlamyphorinae|chlamyphorine]] armadillos approximately 34 million years ago in the [[Priabonian|late Eocene]].<ref name="Mitchell2016">{{cite journal |last1=Mitchell |first1=K.J. |last2=Scanferla |first2=A. |last3=Soibelzon |first3=E. |last4=Bonini |first4=R. |last5=Ochoa |first5=J. |last6=Cooper |first6=A. |year=2016 |title=Ancient DNA from the extinct South American giant glyptodont ''Doedicurus'' sp. (Xenarthra: Glyptodontidae) reveals that glyptodonts evolved from Eocene armadillos |url=http://sedici.unlp.edu.ar/handle/10915/101557 |journal=Molecular Ecology |volume=25 |issue=14 |pages=3499–3508 |doi=10.1111/mec.13695 |pmid=27158910|bibcode=2016MolEc..25.3499M |s2cid=3720645 }}</ref><ref name="Delsuc20162" /> This prompted moving them from their own family, Glyptodontidae, to the subfamily Glyptodontinae within the [[Extant taxon|extant]] [[Chlamyphoridae]].<ref name="Delsuc20162" /> Based on this and the fossil record, glyptodonts would have evolved their characteristic shape and large size ([[gigantism]]) quite rapidly, possibly in response to the cooling, drying climate and expansion of open savannas.<ref name="Mitchell2016" /> Chylamyphoridae is a group in the order Cingulata, which includes all extant armadillos in addition to other fossil groups like [[Pachyarmatheriidae]] and [[Pampatheriidae|Pampatheridae]]. Cingulata is itself within the basal mammal group Xenarthra, which includes an array of American mammal groups like [[Anteater|Vermilingua]] (anteaters) and [[Sloth|Folivora]] (sloths and ground sloths) in the order [[Pilosa]]. The following phylogenetic analysis was conducted by Frédéric Delsuc and colleagues in 2016 and represents the phylogeny of Cingulata using ancient DNA from ''Doedicurus'' to determine the position of it and other Glyptodonts:<ref name="Mitchell2016" /><ref name="Delsuc20162" />
{{clade|{{clade
|1=[[Dasypodidae]]
Line 128 ⟶ 132:
}}
}}|style=font-size:90%;line-height:100%;|label1= [[Cingulata]] }}
[[File:Richard-owen2.jpg|thumb|Portrait of [[Richard Owen|Sir Richard Owen]] (
The internal phylogeny of Glyptodontinae is convoluted and in flux, with many species and families erected based on fragmentary or undiagnostic material that lacks comprehensive review.<ref name="Zurita-2011-2" /><ref name="Porpino-2010" /> Glyptodontinae's tribes were long-considered subfamilies before the 2016 analysis.<ref name="Mitchell-2016">{{Cite journal |last1=Mitchell |first1=Kieren J. |last2=Scanferla |first2=Agustin |last3=Soibelzon |first3=Esteban |last4=Bonini |first4=Ricardo |last5=Ochoa |first5=Javier |last6=Cooper |first6=Alan |date=2016 |title=Ancient DNA from the extinct South American giant glyptodont Doedicurus sp. (Xenarthra: Glyptodontidae) reveals that glyptodonts evolved from Eocene armadillos |url=https://onlinelibrary.wiley.com/doi/10.1111/mec.13695 |journal=Molecular Ecology |language=en |volume=25 |issue=14 |pages=3499–3508 |doi=10.1111/mec.13695|pmid=27158910 |bibcode=2016MolEc..25.3499M |s2cid=3720645 }}</ref> One tribe, Glyptodontini (typically labeled Glyptodontinae) is a group of younger, larger glyptodontines that evolved in the [[Middle Miocene]] (ca. 13 mya) with ''[[Boreostemma]]'',<ref>{{Cite journal |last1=Zurita |first1=Alfredo E. |last2=González Ruiz |first2=Laureano R. |last3=Gómez-Cruz |first3=Arley J. |last4=Arenas-Mosquera |first4=Jose E. |date=2013-05-01 |title=The most complete known Neogene Glyptodontidae (Mammalia, Xenarthra, Cingulata) from northern South America: taxonomic, paleobiogeographic, and phylogenetic implications
Below is the phylogenetic analysis conducted by Cuadrelli ''et al''., 2020 of Glyptodontinae, with Glyptodontidae as a family instead of subfamily, that focuses on advanced glyptodonts:<ref name="Cuadrelli-2020" />
Line 142 ⟶ 146:
|1={{clade
|1={{clade
|1=''[[
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|2={{clade
|1={{clade
|1=''[[Glyptotherium|Glyptotherium cylindricum]]''
|2=''[[
|2={{clade
|1='''''Glyptodon jatunkhirkhi'''''
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|1={{clade
|1=''[[Eleutherocercus|Eleutherocercus antiquus]]''
|2=''[[
|2={{clade
|label1=Neosclerocalyptini
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|2={{clade
|1=''[[Panochthus|Panochthus intermedius]]''
|2=''[[
}} }} }} }} }} }} }} }} }} }} }} }} }}|style=font-size:85%;line-height:80%|label1='''Chlamyphoridae'''}}
== Description ==
[[File:Pleistocene SA.jpg|thumb|Life restoration of ''Glyptodon'' with ''[[Megatherium]]''|243x243px]]
Like the extant armadillos and all other glyptodontines, ''Glyptodon'' had a large, bony carapace that covered much of its torso, as well as smaller [[Head|cephalic]] armor covering the roof of its head, akin to that in turtles. The carapace was composed of hundreds of small, hexagonal osteoderms (armored structures made of bone), with ''Glyptodon'' carapaces preserving a total of 1,800 osteoderms each. The anatomy of different ''Glyptodon'' species varies greatly, mostly in the species ''G. jatunkhirkhi'' which is more similar to ''Glyptotherium'' in certain aspects.<ref name="Cuadrelli-2020" />
In the [[axial skeleton]], glyptodontines had strongly fused [[vertebra]]e and [[Pelvis|pelves]] completely connected to the carapace, traits convergently evolved in turtles.<ref name="Huxley-1865" /><ref name="Gillette" /> The large tails of glyptodontines likely served as a counterbalance to the rest of the body and ''Glyptodon''<nowiki/>'s caudal armor ended in a blunt tube that was composed of two concentric tubes fused together, in contrast to those of mace-tailed glyptodontines like ''Neosclerocalyptus'' and ''Doedicurus''.<ref name="Gillette" /> ''Glyptodon'' had graviportal (weight-bearing), short limbs that are very similar to those in other glyptodontines, being indistinguishable from those of some other taxa.<ref name="Zurita-2018" /> The digits of ''Glyptotherium'' are very stout and adapted for weight-bearing, though some preserve large claw sheaths that had an intermediate morphology between claws and hooves.<ref name="Gillette" />
During the Pleistocene, the diversity of glyptodontines diminished but body size increased, with the largest known glyptodont, ''Doedicurus,'' evolving in the Pleistocene.<ref name="Carlini-2022">{{Cite journal |last1=Carlini |first1=Alfredo A. |last2=Carrillo-Briceño |first2=Jorge D. |last3=Jaimes |first3=Arturo |last4=Aguilera |first4=Orangel |last5=Zurita |first5=Alfredo E. |last6=Iriarte |first6=José |last7=Sánchez-Villagra |first7=Marcelo R. |date=2022-06-16 |title=Damaged glyptodontid skulls from Late Pleistocene sites of northwestern Venezuela: evidence of hunting by humans?
=== Skull, mandible, and dentition ===
[[File:Craneo de Glyptodon.jpg|alt=Glyptodon skull|left|thumb|235x235px|Skull of ''Glyptodon'' in side view.]]
Glyptodont dentition contains entirely [[hypsodont]] molariforms, which have one of the most extreme examples of hypsodonty known from terrestrial mammals.{{Sfn|Gillette|Ray|1981|p=200}}
The nasal passage was reduced with heavy [[muscle]] attachments for some unknown purpose. Some have speculated that the muscle attachments were for a [[proboscis]], or [[Torso|trunk]], much like that of a [[tapir]] or [[elephant]]. The lower jaws were very deep and helped support massive chewing muscles to help chew coarse fibrous plants. Some paleontologists have proposed that ''Glyptodon'' and some glyptodontines also had a [[proboscis]] or large snout similar to those in [[proboscidea]]ns and [[tapir]]s,{{Sfn|Gillette|Ray|1981|p=202}} but few have accepted this hypothesis.<ref name="Zurita-2011-2" /><ref name="Sergio F. Vizcaíno-2012" /> Another suggestion, made by A.E. Zurita and colleagues, is that the large nasal sinuses could be correlated with the cold arid climate of [[Pleistocene]] South America.<ref name="Sergio F. Vizcaíno-2012">{{cite journal |author1=Fernicola, Juan Carlos |author2=Néstor Toledo |author3=M. Susana Bargo |author4=Sergio F. Vizcaíno |date=October 2012 |title=A neomorphic ossification of the nasal cartilages and the structure of paranasal sinus system of the glyptodont Neosclerocalyptus Paula Couto 1957 (Mammalia, Xenarthra) |journal=Palaeontologia Electronica |volume=15 |issue=3 |pages=1–22 |doi=10.26879/333 |doi-access=free}}</ref><ref>{{cite web |author=Gillette, David D. |date=2010 |title=Glyptodonts in
=== Vertebrae and pelvis ===
[[File:Catalogue of casts of fossils (Plate (II)) BHL39943960.jpg|left|thumb|A mounted skeleton of ''G. reticulatus (Schistopleurum asperus''), demonstrating its size.|alt=A mounted skeleton of Glyptodon reticulatus.]]
''Glyptodon'' has 7 [[cervical vertebrae]], of which the first 3 cervicals were fused together while the rest of the cervicals were free except for the 7th.<ref name="Huxley-1865" /> The 7th cervical and the first 2 dorsal vertebrae were fused together into a trivertebral, a broad, flat bone with very small [[spinous processes]] (projections from a vertebra) and large articular surfaces that held ribs.<ref name="Huxley-1865" /> All of the other 13 vertebrae in the dorsal column were fused into one long continuous tunnel that is not seen in mammals outside of glyptodontines, some of these vertebrae were so tightly fused that the segments of them cannot be discerned. The centra of these vertebrae were curved, thin bony plates that created a cylinder to support the carapace and the shape of the animal.<ref name="Huxley-1865" /> [[Vertebra|Spinous processes]] in these vertebrae are also heavily reduced, with some being only a thin blade of bone ankylosed with other vertebrae.<ref name="Huxley-1865" /> [[Sacrum|Sacral vertebrae]] in ''Glyptodon'' are also fused and 13 in number, which preserve very unusual oval-shaped, thin, and slightly concave ends on the centra.<ref name="Huxley-1865" /> The pelves are also unusual, as they preserve giant [[Ilium (bone)|ilia]] and are fused to the rest of the skeleton.<ref name="Huxley-1865" />
=== Carapace and osteoderms ===
[[File:Glyptodon asper armor.JPG|left|thumb|Close-up view of carapace|alt=Close-up view of carapace.]][[File:Armour_of_glyptodon.JPG|thumb|''Glyptodon'' carapace in [[Hungarian Natural History Museum]]|alt=A Glyptodon carapace.]]''Glyptodon''{{'}}s osteoderms were attached by [[Synostosis|synotoses]] (bony connections) and were found in double or triple rows on the front and sides of the carapace's edges, as well as in the tail armor and cephalic shield. The carapace's osteoderms were conical with a rounded point, while the ones on the tail were just [[Cone|conical]]. The sulci between these raised structures were deep and wide with parallel lines.<ref name="Zurita2010">{{cite journal |last1=Zurita |first1=A. E. |last2=Soibelzon |first2=L. H. |last3=Soibelzon |first3=E. |last4=Gasparini |first4=G. M. |last5=Cenizo |first5=M. M. |last6=Arzani |first6=H. |year=2010 |title=Accessory protection structures in ''Glyptodon'' Owen (Xenarthra, Cingulata, Glyptodontidae) |journal=Annales de Paléontologie |volume=96 |issue=1 |pages=1–11 |doi=10.1016/j.annpal.2010.01.001 |bibcode=2010AnPal..96....1Z |hdl-access=free |hdl=10915/5356}}</ref> The carapace of ''Glyptodon'' was strongly elongated compared to those of ''Boreostemma'' and ''Glyptotherium,'' with the carapace being relatively 65% longer than the former and 14% than the latter. In ''Glyptodon'', the top-bottom height of the carapace represents 60% of its total length, whereas in ''Glyptotherium'' it is taller at circa 70%. The antero-posterior dorsal profile of the carapace was convex and its posterior half was higher than the anterior. The apex of the carapace was slightly displaced posteriorly in most ''Glyptodon'' species, while in ''Glyptotherium'' and ''Glyptodon jatunkhirkhi'' it was at the center of the midline. The carapace of most species of ''Glyptodon'' is arched subtly, while ''Glyptotherium'' and ''Glyptodon jatunkhirkhi''{{'}}s has a very arched back and convex pre-iliac and concave post-iliac, giving it a saddle-like overhang over the tail. ''Glyptodon'' osteoderms in the antero-lateral regions of the carapace are strongly ankylosed, giving them little flexibility, while in ''Glyptotherium'' they are less ankylosed and more flexible.<ref name="Zurita-2018" /> The osteoderms of the caudal aperture (
Osteoderms on the ventral side of the body were first mentioned by paleontologist Hermann Burmeister in 1866, postulating that there was a ventral plastron like in turtles based on evidence of small armor in the dermis.<ref name="Burmeister" /> This hypothesis has since been disproven, but in the early 2000s, the presence of osteoderms on ''Glyptodon''{{'}}s face, hind legs, and underside was confirmed in several species. The fossils with these characteristics were from the Pleistocene, evolving in younger species like ''G. reticulatus'' . These small to medium-sized ossicles were actually embedded in the dermis and did not connect in a pattern.<ref name="Zurita2010" />
=== Tail ===
[[File:Cola de ejemplar de género Glyptodon.jpg|left|thumb|Armor at end of tail|alt=Tail armor.|231x231px]]
''Glyptodon'' had very primitive tail anatomy for a glyptodont, possessing eight or nine mobile caudal rings of fused, large, conical osteoderms. These enclosed the base of the tail, which terminated in a short caudal tube composed of two fused caudal rings. Caudal rings were composed of two or three rows of pentagonal osteoderms that transitioned from flat, slightly convex in the posterior rings to conical [[
== Paleobiology ==
=== Digging abilities ===
[[File:Glyptodon (4).jpg|thumb|''Glyptodon'' head restoration, [[Museum of Man and Nature|Munich]]|alt=Reconstruction of head.]]
Many armadillo species have digging capabilities, with large claws adapted for scraping dirt in order to make burrows or forage for food underground.<ref>Vizcaíno, S. F., Fariña, R. A., & Mazzetta, G. V. (1999). Ulnar dimensions and fossoriality in armadillos. ''Acta Theriologica'', ''44''.</ref><ref>Carter, T. S., & Encarnaçao, C. D. (1983). Characteristics and use of burrows by four species of armadillos in Brazil. ''Journal of Mammalogy'', ''64''(1), 103-108.</ref> Much of armadillo diets consist of insects and other invertebrates that live underground,<ref>{{Cite journal |last1=Vizcaíno |first1=Sergio F. |last2=Fariña |first2=Richard A. |date=2007-03-29 |title=Diet and locomotion of the armadillo Peltephilus: a new view |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1502-3931.1997.tb00446.x |journal=Lethaia |language=en |volume=30 |issue=1 |pages=79–86 |doi=10.1111/j.1502-3931.1997.tb00446.x}}</ref> in contrast to the herbivorous diets of ''Glyptodon'' and related genera.<ref name="Saarinen-2017">{{Cite journal |last1=Saarinen |first1=Juha |last2=Karme |first2=Aleksis |date=2017-06-15 |title=Tooth wear and diets of extant and fossil xenarthrans (Mammalia, Xenarthra) – Applying a new mesowear approach |url=https://www.sciencedirect.com/science/article/pii/S0031018216306630 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=476 |pages=42–54 |doi=10.1016/j.palaeo.2017.03.027 |bibcode=2017PPP...476...42S |issn=0031-0182}}</ref> Being a large armadillo, ''Glyptodon''<nowiki/>'s fossorial capabilities have been researched on several occasions. Owen (1841) opposed this idea,<ref name="Owen" /> though pushback came from Nodot (1856) and Sénéchal (1865) who believed digging was possible for the genus.<ref name="Nodot" /><ref name="Sénéchal" /> However, the evolution of a rigid carapace as opposed to a flexible one in extant armadillos as well as a weakly developed [[Humerus|deltoid crest]] on the humerus (upper arm bone) provided evidence against fossorial hypotheses. The elbow had a great range of movement, as with digging cingulates, but this is more likely to be due to size adaptations.<ref
=== Endocranial anatomy ===
[[File:Glyptodontidae brains - Gervais 1869.png|alt=Endocasts of Glyptodon and Doedicurus |left|thumb|[[Endocast
Several complete skulls of ''Glyptodon'' enable the endocranial anatomy to be analyzed, as well as compared to other well-preserved taxa like ''Doedicurus'' and ''Panochthus''. The brain cavities of the larger glyptodontines ''Glyptodon, Doedicurus,'' and ''Panochthus'' had a braincase volume of {{Convert|213 to 234|cm3|U.S.oz}}. The [[encephalization quotient]] of these taxa are 0.12 to 0.4, lower than most modern armadillos (0.44-1.06) and corresponds to those of pampatheres. The brain of the glyptodontines had an extensive [[olfactory bulb]] that took up between 4.8 and 9.7% of the entire brain, while around two thirds of it were occupied by the [[cerebrum]] and the rest by the [[cerebellum]]. Overall, this is akin to that of other armadillos, but in the latter the cerebrum is smaller relative to the cerebellum and the braincase's total volume. Deviating from the armadillos with their wide olfactory bulb, glyptodontines and pampatheres have elongated and triangular olfactory systems. Several other neuroanatomical characteristics differ between glyptodontines and armadillos, such as the
In general, living cingulates have smaller brains than anteaters and sloths for reasons unknown. Several theories have been made as to why, such as a shorter rearing phase of offspring, dedication of resources to the development of the carapace, and other biological and functional handicaps. Members of Cingulata also tend to have extremely low metabolisms, causing less energy flow to the development of the brain's neurons. The pattern of large bodies bearing adequate protection and a reduction of intelligence is found in several other groups such as [[Ankylosauria|ankylosaurs]] and [[Stegosauria|stegosaurs]], two types of armored [[dinosaur]]. However, the carapace itself is considered as a restrictive functional component as it prohibited much neck movement and forced a reduced brain size. This reduction thus resulted in weight loss in the skull, which had a great
=== Feeding and diet ===
[[File:Glyptodon sp teeth.jpg|thumb|204x204px|Teeth, with the distinct hypdodonty of glyptodontines.|alt=Teeth.|left]]
Two main groups of glyptodontines can be distinguished by their feeding habits: narrow-muzzled Miocene propalaehoplophorids and wide-muzzled post-Miocene glyptodontines.<ref>{{Cite web |title=Bargo M. S. Vizcaíno S. F. — Paleobiology of Pleistocene ground sloths (Xenarthra, Tardigrada) : biomechanics, morphogeometry and ecomorphology applied to the masticatory apparatus. Ameghiniana |url=https://www.researchgate.net/publication/262718932 |access-date=2015-10-30 |website=ResearchGate}}</ref> The propalaehoplophorids were selective feeders, while the post-Miocene glyptodontines were [[bulk
Like most other xenarthrans, glyptodontines had lower energy requirements than contemporary mammal groups.<ref>{{Cite journal |last1=Vizcaíno |first1=Sergio F. |last2=Cassini |first2=Guillermo H. |last3=Fernicola |first3=Juan C. |last4=Bargo |first4=M. Susana |year=2011 |title=Evaluating Habitats and Feeding Habits Through Ecomorphological Features in Glyptodonts (Mammalia, Xenarthra) |url=https://www.researchgate.net/publication/257931077 |journal=Ameghiniana |pages=305–319 |doi=10.5710/AMGH.v48i3(364) |hdl=11336/69574 |access-date=2015-10-29 |s2cid=85793531}}</ref> The stomachs of glyptodontids are mysterious
=== Intraspecific combat ===
Glyptodonts are believed to have taken part in [[Intraspecific competition|intraspecific fighting]]. It was presumed that since the tail of ''Glyptodon'' was very flexible and had rings of bony plates, it was used as a weapon in fights. Although its tail could be used for defense against predators, evidence suggests that the tail of ''Glyptodon'' was primarily for attacks on its own kind. A ''G. reticulatus'' fossil displays damage done on the surface of its carapace. A study based on this specimen calculated that ''Glyptodon'' tails would have been able to generate enough force to break the carapace of another ''Glyptodon''. This suggests that they likely fought each other to settle territorial or mating disputes through the use of their tails, much like male-to-male fighting among deer using their antlers
▲Glyptodonts are believed to have taken part in [[Intraspecific competition|intraspecific fighting]]. It was presumed that since the tail of ''Glyptodon'' was very flexible and had rings of bony plates, it was used as a weapon in fights. Although its tail could be used for defense against predators, evidence suggests that the tail of ''Glyptodon'' was primarily for attacks on its own kind. A ''G. reticulatus'' fossil displays damage done on the surface of its carapace. A study based on this specimen calculated that ''Glyptodon'' tails would have been able to generate enough force to break the carapace of another ''Glyptodon''. This suggests that they likely fought each other to settle territorial or mating disputes through the use of their tails, much like male-to-male fighting among deer using their antlers. .<ref name="Alexander1999">{{cite journal |last1=Alexander |first1=R. M. |last2=Fariña |first2=R. A. |last3=Vizcaíno |first3=S. F. |date=May 1999 |title=Tail blow energy and carapace fractures in a large glyptodont (Mammalia, Xenarthra) |journal=Zoological Journal of the Linnean Society |volume=126 |issue=1 |pages=41–49 |doi=10.1111/j.1096-3642.1999.tb00606.x |doi-access=free}}</ref>
=== Ontogeny ===
In 2009, a partial skeleton of a prenatal individual of ''Glyptodon'' was described that had been found inside of the pelvic region of a carapace of an adult.<ref name="Zurita-2009">{{Cite journal |last1=Zurita |first1=Alfredo Eduardo |last2=Miño-Boilini |first2=Angel R. |last3=Soibelzon |first3=Esteban |last4=Scillato-Yané |first4=Gustavo J. |last5=Gasparini |first5=Germán M. |last6=Paredes-Ríos |first6=Freddy |date=2009-09-01 |title=First record and description of an exceptional unborn specimen of Cingulata Glyptodontidae: Glyptodon Owen (Xenarthra) |url=https://www.sciencedirect.com/science/article/pii/S1631068309000694 |journal=Comptes Rendus Palevol |language=en |volume=8 |issue=6 |pages=573–578 |doi=10.1016/j.crpv.2009.04.003 |bibcode=2009CRPal...8..573Z |hdl=11336/37710 |issn=1631-0683}}</ref> The skeleton had been collected from the Pleistocene-aged deposits in the [[Tarija|Tarija Valley]] of Bolivia and included a partial skull, partial mandible, and fragments from the scapulae and femora. The skeleton is the only known prenatal specimen of a
In the osteoderms of juvenile ''Glyptodon reticulatus,'' the central figures are larger than the peripheral osteoderms. These central figures are planar, sometimes even concave, and elevated compared to the peripherals. The peripherals in younger individuals are also less distinct and bear weakly marked or absent furrows (grooves that separate osteoderms). On the other hand, peripherals and central figures of adults are similarly sized, distinct, and of similar heights.<ref name="Cuadrelli-2018" /><ref name="Zurita-2018" />
=== Posture ===
Several interpretations of
=== Sexual dimorphism and group behavior ===
No evidence of [[sexual dimorphism]] in ''Glyptodon'' has been described, but it has been observed in the close relative ''Glyptotherium'' based on fossils found in Pliocene deposits in Arizona''.'' In the genus, the caudal aperture of males and females differ in that the marginal osteoderms of males are much more conical and convex than those of females. Even in the carapaces of newborn ''Glyptotherium'', the marginal osteoderms are either conical or flat, which enables their sex to be determined.<ref name="Gillette-2016" /> No direct evidence of
== Distribution and paleoecology ==
''Glyptodon'' is one of the most common Pleistocene glyptodontines with a large range from the lowland [[Pampas]] to the towering [[Andes|Andean Mountains]] of Peru and Bolivia, some fossils found at elevations reaching over {{Convert|4100|m|ft}} above sea level.<ref>{{Cite journal |last1=Pujos |first1=François |last2=Salas |first2=Rodolfo |date=2004-08-01 |title=A systematic reassessment and paleogeographic review of fossil Xenarthra from Peru |url=https://journals.openedition.org/bifea/5746 |journal=Bulletin de l'Institut français d'études andines |volume=33 |language=en |issue=2 |pages=331–377 |doi=10.4000/bifea.5746 |s2cid=130927479 |issn=0303-7495}}</ref><ref name="Zurita-2017">{{Cite journal |last1=Zurita |first1=Alfredo Eduardo |last2=Zamorano |first2=Martín |last3=Scillato-Yané |first3=Gustavo Juan |last4=Fidel |first4=Sergio |last5=Iriondo |first5=Martín |last6=Gillette |first6=David D. |date=2017-11-17 |title=A new species of Panochthus Burmeister (Xenarthra, Cingulata, Glyptodontidae) from the Pleistocene of the Eastern Cordillera, Bolivia |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2016.1278443 |journal=Historical Biology |language=en |volume=29 |issue=8 |pages=1076–1088 |doi=10.1080/08912963.2016.1278443 |s2cid=91031708 |issn=0891-2963}}</ref><ref name="Cuadrelli-2020" /> Only ''G. munizi'' is found in the early-middle Pleistocene, whereas other species are younger.<ref name="Soibelzon-2006" /><ref name="Cuadrelli-2020" /> ''G. reticulatus'' is specifically noted to be known from 60ka to as recent as 7ka possibly, though confirmed records only extend to 11 ka.<ref name="Ubilla-2018" />The genus had a generalist diet, which allowed it to fill niches in areas that were inaccessible by grazing genera, with ''G. reticulatus'' representing up to 90% of the glyptodontinefossils in the Tarija Valley of Bolivia.<ref name="Zurita" /> However in regions such as the Pampas, Mesopotamia, and Uruguay, an array of glyptodontines are known.<ref name="Ubilla-2018" /><ref name="Cuadrelli-2018" /> Further evidence of ''Glyptodon''{{'}}s adaptability is found in the Pampas, which were semihumid and temperate from 30,000 to 11,000 ka, alternating between the [[Wet season|rainy]] and [[dry season]]s, over a large area consisting mostly of grasslands dotted with forests and mixed shrubbery.<ref>Blasi, A., Castiñeira Latorre, C., Del Puerto, L., Prieto, A. R., Fucks, E., De Francesco, C., ... & Young, A. (2010). [http://www.scielo.org.ar/pdf/lajsba/v17n2/v17n2a02.pdf Paleoambientes de la cuenca media del río Luján (Buenos Aires, Argentina) durante el último período glacial (EIO 4-2).] ''Latin American journal of sedimentology and basin analysis'', ''17''(2), 85-111.</ref><ref>{{Cite journal |last1=Prado |first1=José L. |last2=Martinez-Maza |first2=Cayetana |last3=Alberdi |first3=María T. |date=2015-05-01 |title=Megafauna extinction in South America: A new chronology for the Argentine Pampas |url=https://www.sciencedirect.com/science/article/pii/S0031018215000899 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=425 |pages=41–49 |doi=10.1016/j.palaeo.2015.02.026 |bibcode=2015PPP...425...41P |issn=0031-0182}}</ref><ref name="Cuadrelli-2018" /> This is in stark contrast to the [[Bermejo Formation]] of [[Formosa Province]], Argentina where the climate and fauna suggest a more arid environment with fewer grasslands.<ref>Kruck, W., Helms, F., Geyh, M. A., Suriano, J. M., Marengo, H. G., & Pereyra, F. (2011). Late pleistocene-holocene history of chaco-pampa sediments in Argentina and Paraguay. ''E&G Quaternary Science Journal'', ''60''(1), 14.</ref><ref>Zurita, A. E., M. Taglioretti, M. De los Reyes, C. Oliva, and F. Scaglia. 2014. First Neogene skulls of Doedicurinae (Xenarthra, Glyptodontidae): morphology and phylogenetic implications. Historical Biology 28:423–432.</ref> ''G. jatunkhirkhi'' specifically is known only from Andean climate of Eastern Cordillera in Bolivia, causing it to evolve to be smaller in size than lowland species due to less support for larger masses.<ref name="Cuadrelli-2020" /><ref name="Zurita-2017" /> ''G. jatunkhirkhi'' is not the only example of this in Xenarthra, with species of ''Panochthus'' and ''Pleurolestodon'' evolving to be smaller in size in mountainous regions.<ref name="Zurita-2017" /><ref name="Cuadrelli-2020" />▼
[[File:SerraGeralI.jpg|left|thumb|240x240px|The [[Pampas|Argentine Pampas]], where ''Glyptodon'' was a grazer|alt=The Argentine Pampas grasslands.]]▼
In areas such as Uruguay, fossils of ''Glyptodon'' have been unearthed alongside the contemporary glyptodontines ''Doedicurus, [[Neuryurus]],'' ''Panochthus;'' armadillos ''[[Chaetophractus]], [[Propaeopus]],'' and ''[[Eutatus]];'' and the insectivorous pampathere ''[[Pampatherium]].'' As for their distant relatives the ground sloths, the humongous ''Megatherium'' is known, in addition to two species of the [[Scelidotheriidae|scelidothere]] ''[[Catonyx]]'', and the [[Mylodontidae|mylodontid]] genera ''[[Mylodon]]'' and ''[[Glossotherium]]''. Some other groups are known, including the unusual [[litopterna]]ns ''[[Macrauchenia]]'' and ''[[Neolicaphrium]],'' [[Notoungulata|notoungulate]] ''[[Toxodon]]'', massive [[proboscidea]]n ''[[Notiomastodon]]'', and the [[Equidae|equids]] ''[[Equus neogeus]]'' and ''[[Hippidion]]''. Various [[Even-toed ungulate|artiodactyls]] have been recorded, including the peccaries ''[[Catagonus]]'' and [[White-lipped peccary|''Tayassu peccari'']], extinct deers ''[[Morenelaphus]]'' and ''[[Antifer]]'', and two genera of llamas including ''[[Hemiauchenia]]'' and ''[[Lama (genus)|Lama]].'' A variety of carnivorans have been recorded, such as the "saber-toothed" ''[[Smilodon]],'' the bear ''[[Arctotherium]]'', and the wolf-like canids ''[[Protocyon]],'' and ''[[Dusicyon]]''.<ref>{{Cite journal |last1=Prevosti |first1=F.J. |last2=Ubilla |first2=M. |last3=Perea |first3=D. |date=2009 |title=Large extinct canids from the Pleistocene of Uruguay: systematic, biogeographic and paleoecological remarks |url=http://www.tandfonline.com/doi/abs/10.1080/08912960903249337 |journal=Historical Biology |language=en |volume=21 |issue=1–2 |pages=79–89 |doi=10.1080/08912960903249337 |hdl=11336/103575 |s2cid=85140125 |issn=0891-2963}}</ref><ref>{{Cite journal |last1=Manzuetti |first1=Aldo |last2=Perea |first2=Daniel |last3=Rinderknecht |first3=Andrés |last4=Ubilla |first4=Martín |date=2018-09-01 |title=New Canid Remains from Dolores Formation, late Pleistocene-early Holocene, Uruguay |url=https://doi.org/10.1007/s10914-017-9387-8 |journal=Journal of Mammalian Evolution |language=en |volume=25 |issue=3 |pages=391–396 |doi=10.1007/s10914-017-9387-8 |s2cid=254701482 |issn=1573-7055}}</ref> Rodents too have been found, such as ''[[Holochilus]], [[Hydrochoerus]]'' (capybara), ''[[Cavia]],'' and ''[[Microcavia]]''.<ref name="Ubilla-2018">{{Cite journal |last1=Ubilla |first1=Martín |last2=Rinderknecht |first2=Andrés |last3=Corona |first3=Andrea |last4=Perea |first4=Daniel |date=2018-06-01 |title=Mammals in Last 30 to 7 ka Interval (Late Pleistocene-Early Holocene) in Southern Uruguay (Santa Lucía River Basin): Last Occurrences, Climate, and Biogeography |url=https://doi.org/10.1007/s10914-017-9380-2 |journal=Journal of Mammalian Evolution |language=en |volume=25 |issue=2 |pages=291–300 |doi=10.1007/s10914-017-9380-2 |s2cid=254697032 |issn=1573-7055}}</ref> Notably, some of the youngest "terror-bird" fossils of the genus ''[[Psilopterus]]'' have been unearthed in the area.<ref>{{Cite journal |last1=Jones |first1=Washington |last2=Rinderknecht |first2=Andrés |last3=Alvarenga |first3=Herculano |last4=Montenegro |first4=Felipe |last5=Ubilla |first5=Martín |date=2018-06-01 |title=The last terror birds (Aves, Phorusrhacidae): new evidence from the late Pleistocene of Uruguay |url=https://doi.org/10.1007/s12542-017-0388-y |journal=PalZ |language=en |volume=92 |issue=2 |pages=365–372 |doi=10.1007/s12542-017-0388-y |s2cid=134344096 |issn=1867-6812}}</ref>▼
[[File:Glyptodonts.jpg|thumb|right|The contemporary [[glyptodonts]] ''Glyptodon'', ''[[Doedicurus]]'', and ''[[Panochthus]]'']]
Material previously assigned to ''Glyptodon'' in northeast Brazil has been reassigned to ''Glyptotherium'', restricting the Brazilian distribution of ''Glyptodon'' to the southern provinces. However, two osteoderms with characteristics similar to those of ''Glyptodon'' have been found in [[Sergipe]] state in the northeast, suggesting that both genera occurred in this region during the Pleistocene.<ref name="Dantas2013">{{cite journal |last1=Dantas |first1=M. A. T. |last2=França |first2=L. M. |last3=Cozzuol |first3=M. A. |last4=Rincón |first4=A. D. |year=2013 |title=About the occurrence of ''Glyptodon'' sp. in the Brazilian intertropical region |journal=Quaternary International |volume=305 |pages=206–208 |bibcode=2013QuInt.305..206D |doi=10.1016/j.quaint.2011.06.024}}</ref> ''Glyptodon''<nowiki/>'s northernmost locality comes from Pleistocene deposits o in central Colombia,<ref name="Zurita-2012" /> though many specimens formerly attributed to the genus come from the bordering country of [[Venezuela]].<ref name="Carlini-2022" /> ▼
▲''Glyptodon'' is one of the most common Pleistocene glyptodontines with a large range from the lowland [[Pampas]] to the towering [[Andes|Andean Mountains]] of Peru and Bolivia, some fossils found at elevations reaching over {{Convert|4100|m|ft}} above sea level.<ref>{{Cite journal |last1=Pujos |first1=François |last2=Salas |first2=Rodolfo |date=2004-08-01 |title=A systematic reassessment and paleogeographic review of fossil Xenarthra from Peru |url=https://journals.openedition.org/bifea/5746 |journal=Bulletin de l'Institut français d'études andines |volume=33 |language=en |issue=2 |pages=331–377 |doi=10.4000/bifea.5746 |s2cid=130927479 |issn=0303-7495}}</ref><ref name="Zurita-2017">{{Cite journal |last1=Zurita |first1=Alfredo Eduardo |last2=Zamorano |first2=Martín |last3=Scillato-Yané |first3=Gustavo Juan |last4=Fidel |first4=Sergio |last5=Iriondo |first5=Martín |last6=Gillette |first6=David D. |date=2017-11-17 |title=A new species of Panochthus Burmeister (Xenarthra, Cingulata, Glyptodontidae) from the Pleistocene of the Eastern Cordillera, Bolivia |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2016.1278443 |journal=Historical Biology |language=en |volume=29 |issue=8 |pages=1076–1088 |doi=10.1080/08912963.2016.1278443 |bibcode=2017HBio...29.1076Z |s2cid=91031708 |issn=0891-2963}}</ref><ref name="Cuadrelli-2020" /> Only ''G. munizi'' is found in the early-middle Pleistocene, whereas other species are younger.<ref name="Soibelzon-2006" /><ref name="Cuadrelli-2020" /> ''G. reticulatus'' is specifically noted to be known from 60ka to as recent as 7ka possibly, though confirmed records only extend to 11 ka.<ref name="Ubilla-2018" /> The genus had a generalist diet, which allowed it to fill niches in areas that were inaccessible by grazing genera, with ''G. reticulatus'' representing up to 90% of the
▲[[File:SerraGeralI.jpg|left|thumb|240x240px|The [[
▲During the Ensenadan and Marplatan, ''Glyptodon'' coexisted with a variety of mammals unique to the period such as the [[Notoungulata|notoungulate]] ''[[Mesotherium]]'', canid ''[[Theriodictis]],'' and a species of the giant bear ''[[Arctotherium]]''.<ref name=":4" /> In areas such as Uruguay, fossils of ''Glyptodon'' have been unearthed alongside the contemporary glyptodontines ''Doedicurus, [[Neuryurus]],'' ''Panochthus;'' armadillos ''[[Chaetophractus]], [[Propaeopus]],'' and ''[[Eutatus]];'' and the
▲Material previously assigned to ''Glyptodon'' in northeast Brazil has been reassigned to ''Glyptotherium'', restricting the Brazilian distribution of ''Glyptodon'' to the southern provinces. However, two osteoderms with characteristics similar to those of ''Glyptodon'' have been found in [[Sergipe]] state in the northeast, suggesting that both genera occurred in this region during the Pleistocene.<ref name="Dantas2013">{{cite journal |last1=Dantas |first1=M. A. T. |last2=França |first2=L. M. |last3=Cozzuol |first3=M. A. |last4=Rincón |first4=A. D. |year=2013 |title=About the occurrence of ''Glyptodon'' sp. in the Brazilian intertropical region |journal=Quaternary International |volume=305 |pages=206–208 |bibcode=2013QuInt.305..206D |doi=10.1016/j.quaint.2011.06.024}}</ref> ''Glyptodon''<nowiki/>'s northernmost locality comes from Pleistocene deposits
=== Predation and relationship with humans ===
''Glyptodon'' coexisted with a variety of large predators including the cat ''Smilodon,'' [[jaguar]]s'','' and canid ''Protocyon''.<ref name="Bocherens-2016">{{Cite journal |last1=Bocherens |first1=Hervé |last2=Cotte |first2=Martin |last3=Bonini |first3=Ricardo |last4=Scian |first4=Daniel |last5=Straccia |first5=Pablo |last6=Soibelzon |first6=Leopoldo |last7=Prevosti |first7=Francisco J. |date=2016-05-01 |title=Paleobiology of sabretooth cat Smilodon populator in the Pampean Region (Buenos Aires Province, Argentina) around the Last Glacial Maximum: Insights from carbon and nitrogen stable isotopes in bone collagen |url=https://www.sciencedirect.com/science/article/pii/S0031018216000912 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=449 |pages=463–474 |doi=10.1016/j.palaeo.2016.02.017 |bibcode=2016PPP...449..463B |hdl=11336/43965 |issn=0031-0182}}</ref><ref>Montalvo, C. I., Zárate, M. A., Bargo, M. S., & Mehl, A. (2013). [https://www.researchgate.net/profile/M-Susana-Bargo/publication/259484888_Registro_Faunistico_y_Paleoambientes_del_Cuaternario_Tardio_Provincia_de_la_Pampa_Argentina/links/5433d9210cf2bf1f1f264a74/Registro-Faunistico-y-Paleoambientes-del-Cuaternario-Tardio-Provincia-de-la-Pampa-Argentina.pdf Registro faunístico y paleoambientes del Cuaternario tardío, provincia de La Pampa, Argentina.] ''Ameghiniana'', ''50''(6), 554-570.</ref> This belief is furthered by the discovery of fractured dorsal armor, which implies that ''Glyptodon'' had been in physical conflict with other animals.<ref name="Zurita2010" /> However, [[Isotope analysis|isotope analyses]] of the [[collagen]] from ''Glyptodon'' and other mammals of the Pampas region by Bocherens ''et al''. (2015) discovered little evidence to support the idea of predators feeding on ''Glyptodon''.<ref name="Bocherens-2016" /> Instead, it was found that ''Glyptodon'' as well as herbivorous mammals living in denser forests made up a smaller portion of carnivore diets, whereas open grazers such as ''Lestodon'' and ''Macrauchenia'' were consumed more often''.''<ref name="Bocherens-2016" /> Furthermore, the appearance of secondary armor in the dermis of ''Glyptodon'' coincides with the arrival of North American predators in South America during the [[Great American Interchange]].<ref name=":0" /> For this reason, it was hypothesized that the osteoderms developed as a defensive/offensive mechanism to combat the new arrivals of the area.<ref name="Zurita2010" />
''Smilodon'' may have occasionally preyed upon glyptodontines, based on a skull of ''Glyptotherium texanum'' which bears the distinctive elliptical puncture marks that best match those of the [[machairodont]] cat, indicating that the predator successfully bit into the skull through the armored cephalic shield.<ref>{{cite book |last=Antón |first=Mauricio |title=Sabertooth |date=2013 |publisher=University of Indiana Press |isbn=978-0-253-01042-1 |location=Bloomington, Indiana |pages=203–204}}</ref> The ''Glyptotherium'' in question was a juvenile, with a still-developing head shield, making it far more vulnerable to the cat's attack.<ref name="Gillette2">{{cite journal |last=Gillette |first=D. D. |date=Spring 2010 |title=Glyptodonts in Arizona |url=http://azgeology.azgs.arizona.edu/archived_issues/azgs.az.gov/arizona_geology/spring10/article_feature.html |journal=Arizona Geology |publisher=[[Arizona Geological Survey]] |access-date=2018-08-17}}</ref> Although originally theorized by George Brandes to be possible in 1900,<ref name=":25">Brandes, G. (1900) : Ueber eine Ursache des Aussterbens Diluvialer Säugethiere. Corrblatt d. Deutsch . ''Ges. f. Anthropol. Jahrg. 31. Munichen'' 1901.</ref> ''Smilodon'' canines could not pierce the thick carapace osteoderms of glyptodontines.<ref name=":26">Bohlin, B. (1940). 8. Food habit of the machairodonts, with special regard to ''Smilodon''.</ref> Brandes imagined that the evolution of thick
The coexistence of early [[hunter-gatherer]] humans and glyptodontines in South America was first hypothesized in 1881 based on fossil discoveries from the Pampas,<ref>Vogt, C. (1881). Squelette humain associe aux glyptodontidae. Bulletin de la Societe d’Antropologie de Paris, 3(4), 693–699</ref> and many fossil discoveries from the Late Pleistocene to Early Holocene have been unearthed since that exhibit human predation on glyptodontines. No fossils of ''Glyptodon'' preserving direct interactions have been unearthed, but it did inhabit this region alongside humans. At the site of Pay Paso 1, an archaeological site in northwestern Uruguay preserving human-made spear points and other signs of culture were found associated with fossils of ''Glyptodon'' and the horse ''Equus.'' These were used for radiocarbon dating using collagen, supposedly dating to around 9,000 to 9,500 BP but these dates cannot be verified.<ref>{{Cite journal |last=Suárez |first=Rafael |date=2015 |title=The Paleoamerican Occupation of the Plains of Uruguay: Technology, Adaptations, and Mobility |url=http://www.tandfonline.com/doi/full/10.1179/2055556314Z.00000000010 |journal=PaleoAmerica |language=en |volume=1 |issue=1 |pages=88–104 |doi=10.1179/2055556314Z.00000000010 |s2cid=128985617 |issn=2055-5563}}</ref><ref name="Zurita-2018" /> During this period, a wide array of Xenarthrans inhabited the Pampas were hunted by humans, with evidence demonstrating that the small ({{convert|300|–|450|kg|abbr=on|disp=comma}}) glyptodontine ''Neosclerocalyptus,''<ref name="Quiñones">{{Cite journal |last1=Quiñones |first1=Sofía I. |last2=De los Reyes |first2=Martin |last3=Zurita |first3=Alfredo E. |last4=Cuadrelli |first4=Francisco |last5=Miño-Boilini |first5=Ángel R. |last6=Poiré |first6=Daniel G. |date=2020-11-01 |title=Neosclerocalyptus Paula Couto (Xenarthra, Glyptodontidae) in the late Pliocene-earliest Pleistocene of the Pampean region (Argentina): Its contribution to the understanding of evolutionary history of Pleistocene glyptodonts |url=https://www.sciencedirect.com/science/article/pii/S0895981120302443 |journal=Journal of South American Earth Sciences |language=en |volume=103 |pages=102701 |bibcode=2020JSAES.10302701Q |doi=10.1016/j.jsames.2020.102701 |issn=0895-9811 |s2cid=225024450}}</ref> the armadillo ''Eutatus'', and the gigantic (2 ton) glyptodontine ''Doedicurus'', the largest
== Extinction ==
{{Main|Late Pleistocene extinctions}}
Some evidence suggests that humans drove glyptodontines to extinction.<ref name = "islands&continents">{{Cite book▼
''Glyptodon'', along with all other glyptodonts became extinct around the end of the Late Pleistocene, as part of [[Late Pleistocene extinctions|a wave of extinctions of most large mammals]] across the Americas.
▲Some evidence suggests that humans drove glyptodontines to extinction.<ref name
| last1 = Martin | first1 = P. S. | author-link = Paul Schultz Martin
| last2 = Steadman | first2 = D. W. | author2-link = David Steadman
Line 264 ⟶ 274:
| isbn = 978-0-306-46092-0
| oclc = 41368299 | access-date = 2015-11-07
}}</ref> Evidence from the Campo Laborde and La Moderna archaeological sites in the Argentine Pampas suggest that ''Glyptodon'<nowiki/>''s relatives ''Doedicurus'' and ''Panochthus'' survived until the Early Holocene, coexisting with humans for a minimum of 4,000 years.<ref name="Politis2008">{{cite journal|last1=Politis|first1=G. G.|last2=Messineo|first2=P. G.|title=The Campo Laborde site: New evidence for the Holocene survival of Pleistocene megafauna in the Argentine Pampas|journal=Quaternary International|volume=191|issue=1|date= November 2008 |pages=98–114 |doi=10.1016/j.quaint.2007.12.003|bibcode=2008QuInt.191...98P }}</ref> This overlap provides support for models showing that the South American Pleistocene extinctions resulted from a combination of climatic change and anthropogenic causes.<ref name="Politis2008" /> These sites have been interpreted as ones used for butchering megafauna (''Megatherium'' and ''Doedicurus''); however, some of the chronology has been problematic and controversial, due to poor preservation of the collagen used for dating.<ref name="Politis2008" /> The extinction rates in South America during the
The extinction of ''Glyptodon'' notably coincides with the end of the [[Antarctic Cold Reversal]] period in which, for 1,700 years, temperatures dropped before spiking after ending at 12.7 ka.<ref>{{Cite journal |last1=Orombelli |first1=Giuseppe |last2=Maggi |first2=Valter |last3=Delmonte |first3=Barbara |date=2010-06-01 |title=Quaternary stratigraphy and ice cores |url=https://www.sciencedirect.com/science/article/pii/S1040618209003474 |journal=Quaternary International |series=Plio-Pleistocene Correlation and Global Change |language=en |volume=219 |issue=1 |pages=55–65 |doi=10.1016/j.quaint.2009.09.029 |bibcode=2010QuInt.219...55O |issn=1040-6182}}</ref><ref>{{Cite journal |last1=Weber |first1=M. E. |last2=Clark |first2=P. U. |last3=Kuhn |first3=G. |last4=Timmermann |first4=A. |last5=Sprenk |first5=D. |last6=Gladstone |first6=R. |last7=Zhang |first7=X. |last8=Lohmann |first8=G. |last9=Menviel |first9=L. |last10=Chikamoto |first10=M. O. |last11=Friedrich |first11=T. |last12=Ohlwein |first12=C. |date=2014 |title=Millennial-scale variability in Antarctic ice-sheet discharge during the last deglaciation |url=https://www.nature.com/articles/nature13397 |journal=Nature |language=en |volume=510 |issue=7503 |pages=134–138 |doi=10.1038/nature13397 |pmid=24870232 |bibcode=2014Natur.510..134W |s2cid=205238911 |issn=1476-4687}}</ref> Many climatic fluctuations occurred during the late Pleistocene between humid and dry cycles, with ''Glyptodon'' preferring drier climates. Following the Antarctic Cold Reversal, temperatures rose and the climate became more consistently humid, which then led C3 grasses to become increasingly replaced by C4 grasses and southern beech trees. These changes led vulnerable, grazing-specialized forms like glyptodontines, toxodonts, and some ground sloths to become extinct.<ref>{{Cite journal |last1=Villavicencio |first1=Natalia A. |last2=Werdelin |first2=Lars |date=2018-09-01 |title=The Casa del Diablo cave (Puno, Peru) and the late Pleistocene demise of megafauna in the Andean Altiplano |url=https://www.sciencedirect.com/science/article/pii/S0277379118301318 |journal=Quaternary Science Reviews |language=en |volume=195 |pages=21–31 |doi=10.1016/j.quascirev.2018.07.013 |bibcode=2018QSRv..195...21V |s2cid=134626837 |issn=0277-3791}}</ref><ref>{{Cite journal |last1=Hubbe |first1=Alex |last2=Hubbe |first2=Mark |last3=Karmann |first3=Ivo |last4=Cruz |first4=Francisco W. |last5=Neves |first5=Walter A. |date=2011 |title=Insights into Holocene megafauna survival and extinction in southeastern Brazil from new AMS 14C dates |url=https://www.cambridge.org/core/journals/quaternary-research/article/abs/insights-into-holocene-megafauna-survival-and-extinction-in-southeastern-brazil-from-new-ams-14c-dates/38AB0ACF3EE6A9405937A0E22A613F8F |journal=Quaternary Research |language=en |volume=79 |issue=2 |pages=152–157 |doi=10.1016/j.yqres.2012.11.009 |s2cid=129827927 |issn=0033-5894}}</ref> Around 11.5 ka, temperatures peaked before again dropping, resulting in the extinction of several different genera of mammals including some megafauna. ''Glyptodon'' along with genera such as ''Glossotherium'' and ''Morenelaphus'' were wiped out, though several other groups lived for several thousand years after.<ref name="Mitchell-2016" /><ref name="Ubilla-2018" />
== See also ==
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[[Category:Prehistoric placental genera]]
[[Category:Pleistocene xenarthrans]]
[[Category:Pleistocene mammals of South America]]
[[Category:Pleistocene Argentina]]
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