Glyptodon
Glyptodon | |
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Skeleton of G. clavipes at the Naturhistorisches Museum, Vienna
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Cingulata |
Family: | Chlamyphoridae |
Subfamily: | †Glyptodontinae |
Genus: | †Glyptodon Owen, 1839 |
Type species | |
†Glyptodon clavipes Owen, 1839
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Other Species | |
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Distribution of Glyptodon (green) compared to Glyptotherium's (orange). | |
Synonyms | |
Genus synonymy
Synonyms of G. clavipes
Synonyms of G. reticulatus
Dubious species
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Glyptodon (lit. 'grooved or carved tooth'; from
The type species, G. clavipes, was described in 1839 by notable British paleontologist 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, quadrapedral (four-legged), herbivorous armadillos with armored carapaces (top shell) that were made of hundreds of interconnected osteoderms (structures in dermis composed of bone). Other pieces of armor covered the tails and skull roofs, the skull being tall with hypsodont (high-crowned) teeth. As for the postcranial anatomy, pelves fused to the carapace, an amalgamate vertebral column, short limbs, and small digits are found in glyptodontines. Glyptodon reached up to 2 meters (6.56 feet) long and 400 kilograms (880 pounds) in weight, making it one of the largest glyptodontines but not as large as its close relative Glyptotherium or Doedicurus, the largest known glyptodont. Glyptodon is morphologically and phylogenetically most similar to Glyptotherium, however they differ in several ways. Glyptodon is larger on average, with an elongated carapace, a relatively shorter tail, and a robust zygoma, or cheek bone.
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 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 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 Paleoindians suggests that humans may have been a causal factor in the extinctions.
History
Confusion with Megatherium
The history and taxonomy of Glyptodon is storied and convoluted, as it involved confusion with other genera and 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 (1771-1848) wrote about the discovery of several unusual fossils in his Diario de Historia Natural, which included his descriptions of many new species of ants, birds, mammals, and even one of the first figures of the extinct Megatherium, a genus of giant ground sloth that was named in 1796 by French scientist Georges Cuvier (1769-1832).[3][4] This was the first recorded discovery of a glyptodontine or fossil cingulate.[4] The unusual fossils consisted of a femur, carapace fragments, and a caudal tube (an armored tail covering found in glyptodontines) that he collected from the Pleistocene aged (ca. 2.5-0.011 mya) strata on the banks of the Solís Grande Creek, Uruguay.[4][5] Larrañaga identified the fossils as those of Dasypus (Megatherium), believing that Megatherium was a subgenus of Dasypus based on the incorrect referral of glyptodontine osteoderms to Megatherium years earlier by Spanish scientist Juan Bautista Bru de Ramón, which misled other scientists to believe that glyptodontine fossils were actually those of armored megatheres.[6][4]
Larrañaga wrote to French scientist 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.[7] Larrañaga also noted that similar fossils had been found in "analogous strata near Lake Merrim, on the frontier of the Portuguese colonies (southern Brazil)."[7][6] These fossils were also likely those of glyptodontines, possibly the closely related Hoplophorus.[8] The armored Megatherium hypothesis was further supported later in 1827 when portions of a Glyptodon carapace, as well as a partial femur and some caudal armor, were found by a Prussian traveler to Montevideo, Uruguay named Mr. Sellow, who sent the carapace to Berlin where it was described by Professor von Weiss, who referred it to Megatherium.[9] The femur and caudal armor were recovered from the Quegnay in northern Uruguay, while the carapace had been found in the Arapey River.[9][5] Weiss and other paleontologists noted that the osteoderms closely resembled those of armadillos like Dasypus, but Cuvier's hypothesis was popularized based on the incorrect referral of glyptodontine osteoderms Megatherium.[6][7]
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, 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.[5]
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 in Brazil, dating to the Pleistocene.[10][8] The fossils included osteoderms comparable to those described earlier by Larrañaga, as well as teeth, skull fragments, limb bones, and other elements.[5][8] 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,[11] Pachypus by Eduard D'Alton in 1839 based on Sellow's 1833 material,[5] Neothoracophorus (originally Thoracophorus but the name was preoccupied by a beetle) in 1889 by Argentine paleontologist Florentino Ameghino[12] based on isolated osteoderms now identified as those of a juvenile Glyptodon from Patagonia,[13] and Lepitherium in 1839 by Geoffroy Saint-Hilaire based on Sellow's osteoderms.[14][15] 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.[15][14]
Richard Owen and referred species
In 1838, British diplomat Sir Woodbine Parish (1796–1882) was sent an isolated molariform and a letter about the discovery of several large fossils from the Matanza River in Buenos Aires, Argentina that dated to the Pleistocene.[16][17] Parish later collected several more fossils from localities in Las Averias and Villanueva; the latter preserved the most complete skeleton which included a mandible fragment, partial limbs, and unguals of a single individual. They were deposited in Parish's collection at the Royal College of Surgeons in the United Kingdom that year. Some of these fossils were cast at the Natural History Museum, London, but the original fossils were destroyed after German aerial bombing raids hit the college during World War II from 1940 to 1941.[13][17] Glyptodon was named by Richard Owen (1804-1892), one of the most influential British naturalists of the Victorian Era, writing a chapter about the animal and publishing a reconstruction of its skeleton in the book Buenos Ayres, and the provinces of the Rio de La Plata: their present state, trade, and debt in 1839.[18][19] Within this book, Owen erroneously believed they were all from the same specimen, the name Glyptodon ("grooved tooth") based on the anatomy of the molariform.[19][17] A later study found the molariform to actually be from another glyptodontine, Panochthus, and the Villanueva individual was designated the lectotype by Robert Hoffstetter in 1955.[20] The Las Averias individual consists of a carapace that was only mentioned in Owen's description, but was used in later reconstructions of the animal and has since been lost. An issue with the lectotype of G. clavipes is that the material is undiagnostic and indistinguishable from other Glyptodon species and even Glyptotherium, making it dubious.
Cuadrelli et al (2018) designated the species a species inquirenda due to this issue and commented that more analyses are necessary.[17] In 1860, Signor Maximo Terrero collected a partial skeleton, including a skull and carapace, of G. clavipes from the 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.[5] This skeleton was also destroyed during WWII, but Huxley published several illustrations that presented great amounts of new information on the taxon.[5][21]
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,[22] G. tuberculatus to Panochthus,[23] and G. clavicaudatus to Doedicurus.[24] 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.[25]
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. In the 1950s, Argentine paleontologist Alfredo Castellanos (1893-1975) erected new generic names for several species of Glyptodon, the first being Glyptocoileus and second of these being Glyptopedius in 1953 that was made for the species G. elongatus that had been named by Robert Burmeister in 1866 on the basis of a single carapace,[30] though its validity is disputed.[26][17] Castellanos also referred the species G. reticulatus to the genus, but this unsupported.[17] Yet another genus was erected in 1976 named Heteroglyptodon genuarioi by F. L. Roselli based on an incomplete skeleton that had been collected from the Pleistocene aged Libertad Formation in Nueva Palmira, Uruguay,[34][35] but it has since been found to be an indeterminate specimen of Glyptodon.[35] Several Glyptodon fossils from Pleistocene deposits in Colombia were described in 2012, extending the known range of the genus north greatly.[36]
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 Yamparaez, 24 kilometres (15 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 4,100–2,500 metres (13,500–8,200 ft) above sea level.[26] Several additional paratypes were referred to the species from other Late Pleistocene sites in Eastern Cordillera, Bolivia including a nearly complete skull and several osteoderms.[26] 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. Reassessment of Glyptodon species began in the late 20th and early 21st centuries, with various hypotheses developing on the number of valid species. Numbers varied, with some authors considering up to 4 species valid, while phylogenetic analyses in 2018 and 2020 only found the species G. reticulatus, G. munizi, and G. jatunkhirkhi definitively valid; G. clavipes and G. uquiensis as species inquirendas.[17] However a 2016 review of G. uquiensis determined that G. uquiensis was actually a juvenile specimen of Glyptodon, though the species could not be determined.[33]
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 stage of the Pleistocene.[37][25] 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.[25][38] 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.[39]
Glyptodon is the type genus of Glyptodontinae, an extinct subfamily of large, heavily armored armadillos that first evolved in the Late Eocene (ca. 33.5 mya) and went extinct in the Early Holocene during the Late Pleistocene extinctions (ca. 7,000 years ago).[26][40] Owen recognized that Glyptodon was an edentate, but did not recognize it as being a part of a new subfamily as there were no other recognized glyptodontines in 1839.[19] The family Glyptodontidae was not named until 1869 by John Edward Gray, who included the genera Glyptodon, Panochthus, and Hoplophorus within the group and believed that it was diagnosed by an immovable carapace that was fused to the pelvis.[41] However, Hermann Burmeister proposed the name Biloricata for the family, believing that glyptodontines possessed a ventral plastron (bottom shell) and could pull their heads inside their carapaces like turtles. This name lost all use and his theory has not been supported.[30][23] The internal phylogenetics of Glyptodontidae was analyzed in greater detail by Florentino Ameghino during his descriptions of earlier members of the clade, which proposed that Glyptodon was descended from earlier genera.[12][28]
Glyptodontinae was classified in its own family or even superfamily until in 2016, when Reassessment and Glyptotherium
Taxonomy
Cingulata |
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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.[43][8] Glyptodontinae's tribes were long-considered subfamilies before the 2016 analysis.[44] One tribe, Glyptodontini (typically labeled Glyptodontinae) is a group of younger, larger glyptodontines that evolved in the Middle Miocene (ca. 13 mya) with Boreostemma,[45] but split into two genera, Glyptodon in the south and Glyptotherium in the north,[36] though Glyptotherium also lived in some areas of South America like Venezuela and eastern Brazil.[46][47][36] Glyptotherium and Glyptodon lived during the same intervals and are nearly identical to Glyptodon in many aspects, so much so that the first fossils of Glyptotherium to be described were misidentified as those of Glyptodon.[48][49][25] Glyptodontini is distinguishable from other groups for example in that it has large, conical tubercular osteoderms absent or only present on the caudal (tailward) notch on the posterior end of the carapace and different ornamentation of the armor on the carapace than the tail.[25] Glyptodontini is often recovered as more basal to most other glyptodontines like Doedicurus, Hoplophorus, and Panochthus.[50][26]
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:[26]
Chlamyphoridae |
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Description
Like the extant armadillos and all other glyptodontines, Glyptodon had a large, bony carapace that covered much of its torso, as well as smaller 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.[26]
In the axial skeleton, glyptodontines had strongly fused vertebrae and pelves completely connected to the carapace, traits convergently evolved in turtles.[5][38] The large tails of glyptodontines likely served as a counterbalance to the rest of the body and Glyptodon'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.[38] Glyptodon had graviportal (weight-bearing), short limbs that are very similar to those in other glyptodontines, being indistinguishable from those of some other taxa.[25] 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.[38]
During the Pleistocene, the diversity of glyptodontines diminished but body size increased, with the largest known glyptodont, Doedicurus, evolving in the Pleistocene.[51][52] Glyptodon sizes vary between species and individuals. G. clavipes, the type species, was estimated to weigh 2,000 kilograms (4,400 lb),[53][54] G. reticulatus weighed a mere 401 kilograms (884 lb) to 862 kilograms (1,900 lb),[55] and G. munizi weighed 1,150 kilograms (2,540 lb).[56] A partial skeleton of G. clavipes measured 3.5 metres (11 ft) with a carapace length of 1.7 metres (5.6 ft),[4][21] while the carapaces of other species like G. munizi and G. reticulatus measured 2.2 metres (7.2 ft) and 2.19 metres (7.2 ft) long respectively.[25]
Skull, mandible, and dentition
Glyptodont dentition contains entirely hypsodont molariforms, which have one of the most extreme examples of hypsodonty known from terrestrial mammals.[57] The dentition is typical of other armadillos, but is fluted on each side by deep grooves. The anterior teeth were compressed, while the posterior teeth were cylindrical.[58] Glyptodont skulls have several unique features; the maxilla and palatine are enlarged vertically to make space for the molariforms, while the braincase is brachycephalic, short and flat.[59] In Glyptodon and many other glyptodontines, the roof of the skull was covered by a shield composed of polygonal, irregular osteoderms that were variable in size and ankylosed together to form a robust cephalic shield that had a smoothly convex exterior surface without ornamentation.[28] Each osteoderm has a rugose and slightly convex dorsal surface, with ornamentation pattern defined by a central figure, slightly elevated and surrounded by an area without peripheral figures or foramina. Sutures separating osteoderms are well marked, as in Panochthus.[28][60] Other Pleistocene glyptodontines are known by complete/sub-complete skulls, allowing for comparisons to Glyptodon.[13][43] Glyptotherium's zygoma are narrow, slender, almost parallel, and close to the sagittal plane in frontal view; in Glyptodon, this structure is broader, robust, divergent rather than parallel and more laterally placed.[51][25]
The nasal passage was reduced with heavy
Vertebrae and pelvis
Glyptodon has 7
Carapace and osteoderms
The coexistence of early hunter-gatherer humans and glyptodontines in South America was first hypothesized in 1881 based on fossil discoveries from the Pampas,[113] 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.[114][25] During this period, a wide array of Xenarthrans inhabited the Pampas were hunted by humans, with evidence demonstrating that the small (300–450 kg, 660–990 lb) glyptodontine Neosclerocalyptus,[115] the armadillo Eutatus, and the gigantic (2 ton) glyptodontine Doedicurus, the largest glyptodontine known, were hunted.[116] The only other records of human predation from outside the Pampas area a partial carapace, which was eviscerated by humans, and several skulls preserving signs that they were dispatched by human tools. All were found in Venezuela.[117][51] The discoveries there showed the first signs of human hunting on the skulls of glyptodontines.[118] Hunters may have used the shells of dead animals as shelters in inclement weather.[119][120]
Extinction
Glyptodon, along with all other glyptodonts became extinct around the end of the Late Pleistocene, as part of a wave of extinctions of most large mammals across the Americas.
Some evidence suggests that humans drove glyptodontines to extinction.[121] Evidence from the Campo Laborde and La Moderna archaeological sites in the Argentine Pampas suggest that Glyptodon's relatives Doedicurus and Panochthus survived until the Early Holocene, coexisting with humans for a minimum of 4,000 years.[122] This overlap provides support for models showing that the South American Pleistocene extinctions resulted from a combination of climatic change and anthropogenic causes.[122] 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.[122] The extinction rates in South America during the late Pleistocene were the highest out of any continent, with all endemic animals weighing over 100 kilograms (220 lb) going extinct by the middle Holocene.[4] This supports the idea of human hunting as a drive for the extinction of Glyptodon, as the arrival of humans around 16,000 years BP to such a formerly isolated continent may have caused extinction rates to become higher.[118][4][123]
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.[124][125] 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.[126][127] 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.[44][98]
See also
- Paleontology portal
- Prehistoric mammals portal
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