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Glyptodon

Glyptodon is a genus of glyptodont, an extinct group of large, herbivorous armadillos that lived from the Pliocene, around 3.2 million years ago, to the early Holocene, around 11,000 years ago, in South America. It is one of, if not the, best-known genera of glyptodont. Glyptodon has a long and storied past, being the first named extinct cingulate and the type genus of the subfamily Glyptodontinae. Fossils of Glyptodon have been recorded as early as 1814 from Pleistocene aged deposits from Uruguay, though many were incorrectly referred to the ground sloth Megatherium by early paleontologists.

History
Confusion with Megatherium (1769–1832), describer of Megatherium|alt=Portrait of Georges Cuvier, describer of 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). This was the first recorded discovery of a glyptodontine or fossil cingulate. 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. Larrañaga also noted that similar fossils had been found in "analogous strata near Lake Mirrim, on the frontier of the Portuguese colonies" (southern Brazil). The femur and caudal armor were recovered from the Queguay in northern Uruguay, while the carapace had been found in the Arapey River. The fossils included osteoderms comparable to those described earlier by Larrañaga, as well as teeth, skull fragments, limb bones, and other elements. Pachypus by Eduard D'Alton in 1839, Saint-Hilaire 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. Parish later collected several more fossils from localities in Las Averias and Villanueva; the latter including a partial skeleton containing a mandible fragment and a set of partial limbs. This skeleton was deposited in Parish's collection at the Royal College of Surgeons upon his return to the United Kingdom. Within this book, Owen erroneously believed the fossils from Las Averias and Villanueva were all from the same specimen, which he named Glyptodon ("grooved tooth") based on the anatomy of the molariform. However, the lectotype of G. clavipes was undiagnostic and indistinguishable from other Glyptodon species and even Glyptotherium, making it dubious. 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, G. tuberculatus to Panochthus, and G. clavicaudatus to Doedicurus. 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 Natural History Museum (NHMUK) as well, being used to diagnose the species. 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. but it has since been synonymized with G. reticulatus. 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, Argentina. 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. P. uquiensis has been synonymized with Glyptodon and is possibly a valid species, though further analysis is necessary to settle its status. but it has since been found to be an indeterminate specimen of Glyptodon. 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, 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 above sea level. 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. 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. == Taxonomy ==
Taxonomy
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). 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. 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. One tribe, Glyptodontini (typically labeled Glyptodontinae) is a group of younger, larger glyptodontines that evolved in the Middle Miocene (ca. 13 mya) with Boreostemma, but split into two genera, Glyptodon in the south and Glyptotherium in the north, 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: == Description ==
Description
''|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 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. Glyptodon sizes vary between species and individuals. G. clavipes, the type species, was estimated to weigh , G. reticulatus weighed a mere to , and G. munizi weighed . A partial skeleton of G. clavipes measured with a carapace length of , 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. 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. Other Pleistocene glyptodontines are known by complete/sub-complete skulls, allowing for comparisons to Glyptodon. 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. A distinctive bar of bone projects downwards on the cheek, extending over the lower jaw, perhaps providing an anchor for powerful snout muscles. In turn, the infraorbital foramina are narrow and not visible in anterior view in Glyptotherium, but in Glyptodon they are broad and clearly visible in anterior view. In lateral view, the dorso-ventral height between the skull roof and the palatal plane in Glyptodon decreases anteriorly, contrary to Glyptotherium; the nasal tip is in a lower plane with respect to the zygomatic arch in Glyptodon, but in Glyptotherium is higher than the zygomatic arch plane. The 1st molariform (molaiform is abbreviated as mf#) of Glyptodon is distinctly trilobate (three-lobed) both lingually and labially, nearly as trilobate as the mf2; on the contrary, Glyptotherium shows a very low trilobation of mf1, which is elliptical in cross-section, the mf2 is weakly trilobate, and the mf3 is trilobate. In both genera, the mf4 to mf8 are fully trilobate and serially identical. 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 jatunkhirkhis 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. The caudal aperture is more vertically oriented in the latter genus, while in Glyptotheirum it is angled posteriorily. The caudal tube at the distalmost end of the tail is cylinder-shaped with smaller conical osteoderms and is stubbier proportionally in Glyptodon. In Glyptotherium, this caudal tube represents ca. 20% of the total length of the caudal armor, whereas in Glyptodon, this structure represents 13% of the total length. In Glyptodon, the caudal armor length represents circa 30-40% of the carapace's total length in contrast to Glyptotherium, where this value is greater at around 50%. For example, in specimen MCA 2015 of Glyptodon reticulatus, the terminal tube measured only long in comparison to Glyptotherium texanum specimen UMMP 34 826's long tube. == Paleobiology ==
Paleobiology
Digging abilities |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. Much of armadillo diets consist of insects and other invertebrates that live underground, in contrast to the herbivorous diets of Glyptodon and related genera. Being a large armadillo, Glyptodon's fossorial capabilities have been researched on several occasions. Owen (1841) opposed this idea, Endocranial anatomy s of Glyptodon (left) and Doedicurus (right).|256x256px 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 . 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 presence of a pronounced sulcus praesylvianus. 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 ankylosaurs and 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 effect on the skulls of large-headed glyptodontines like Glyptodon. The propalaehoplophorids were selective feeders, while the post-Miocene glyptodontines were bulk feeders (obtain nutrients by consuming an entire plant). However, because of their body form and fusion of the cervical vertebrae glyptodontines would have needed to forage near the ground. Their craniomandibular joint limited their jaw to side-to-side movement. Glyptodon's jaws had large ridges of osteodentine which could effectively be used to grind food particles before shearing and pushing them via the constant motion of the mandible. The hyoid shows a robust design that suggests Glyptodon had a large and robust tongue, which may have aided in food intake and processing. Like most other xenarthrans, glyptodontines had lower energy requirements than contemporary mammal groups. The stomachs of glyptodontids are mysterious due to being entirely herbivorous, in contrast to modern, omnivorous armadillos which have simple stomachs instead of the chambered ones of sloths. This in conjugation with the proposed idea that aquatic grazing may have caused the isotopes strongly associated with herbivory observed in Glyptodon fossils. though more backing for this hypothesis has been found in the related Glyptotherium. The 2012 paper also noted that Glyptodon may have had a more flexible diet than previously imagined, with a mix of slightly wooded and slightly open habitats as implied by the consumption of C3 and C4 material. 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. The skeleton had been collected from the Pleistocene-aged deposits in the 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 glyptodontine and is one of the most complete specimens of an immature Glyptodon known, though dozens of isolated osteoderms from juveniles are known. starting with those by Richard Owen in 1841 using comparative anatomy. Linear measurements were later taken which provided insight into this hypothesis, finding that bipedalism would be possible. The patellar articulation with the femur suggests rotation of the lower leg during knee extension and potentially even knee-locking were feasible. Trackways The first ever glyptodont footprints were recovered from the late Pleistocene Pehuén-Có fossil site, Argentina, and referred to Glyptodon. Described as the ichnotaxon Glyptodontichnus pehuencoensis in 2015, the rarity of glyptodont trackways despite their frequency in the fossil record suggests that glyptodonts may have avoided walking on muddy substrates as they'd be especially vulnerable to getting trapped. 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. == Distribution and paleoecology ==
Distribution and paleoecology
Glyptodon, Doedicurus, and Panochthus Glyptodon is one of the most common Pleistocene glyptodontines with a large range from the lowland Pampas to the towering Andean Mountains of Peru and Bolivia, some fossils found at elevations reaching over above sea level. 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. 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. Rodents too have been found, such as Holochilus, Hydrochoerus (capybara), Cavia, and Microcavia. Notably, some of the youngest "terror-bird" fossils of the genus Psilopterus have been unearthed in the area. 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. Glyptodon's northernmost locality comes from Pleistocene deposits in central Colombia, This belief is furthered by the discovery of fractured dorsal armor, which implies that Glyptodon had been in physical conflict with other animals. The Glyptotherium in question was a juvenile, with a still-developing head shield, making it far more vulnerable to the cat's attack. Although originally theorized by George Brandes to be possible in 1900, Smilodon canines could not pierce the thick carapace osteoderms of glyptodontines. Brandes imagined that the evolution of thick glyptodontine armor and long machairodont canines was an example of coevolution, 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. the armadillo Eutatus, and the gigantic (2 ton) glyptodontine Doedicurus, the largest glyptodontine known, were hunted. 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. Glyptodon was also a victim of parasitism, as evidenced by findings of Karethraichnus kulindros on an articulated carapace of G. clavipes, which are believed to represent traces made by tungid fleas that were related to Tunga perforans. == Extinction ==
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. 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. This overlap provides support for models showing that the South American Pleistocene extinctions resulted from a combination of climatic change and anthropogenic causes. 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. 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. 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. == See also ==
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