| Glyptodon | |
|---|---|
 | |
| Skeleton ofG. clavipes at theNaturhistorisches Museum,Vienna | |
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 | |
| OtherSpecies | |
 | |
| Distribution ofGlyptodon (green) compared toGlyptotherium's (orange). | |
| Synonyms | |
Genus synonymy
Synonyms ofG. clavipes
Synonyms ofG. reticulatus
Dubious species
| |
Glyptodon (lit. 'grooved or carved tooth'; from Ancient Greek γλυπτός (gluptós) 'sculptured' and ὀδοντ-,ὀδούς (odont-, odoús) 'tooth')[1] is a genus ofglyptodont, an extinct group of large, herbivorousarmadillos, that lived from thePliocene, around 3.2 million years ago,[2] to the early Holocene, around 11,000 years ago, inSouth America. It is one of, if not the, best known genus of glyptodont.Glyptodon has a long and storied past, being the first named extinctcingulate and the type genus of the subfamily Glyptodontinae. Fossils ofGlyptodon have been recorded as early as 1814 from Pleistocene aged deposits from Uruguay, though many were incorrectly referred to theground slothMegatherium by early paleontologists.
Thetype species,G. clavipes, was described in 1839 by notable BritishpaleontologistSir Richard Owen. Later in the 19th century, dozens of complete skeletons were unearthed from localities and described by paleontologists such asFlorentino Ameghino andHermann Burmeister. During this era, many species ofGlyptodon were dubbed, some of them based on fragmentary or isolated remains. Fossils from North America were also assigned toGlyptodon, but all of them have since been placed in the closely related genusGlyptotherium. It was not until the later end of the 1900s and 21st century that full review of the genus came about, restrictingGlyptodon to just five species under one genus.
Glyptodonts were typically large, quadrupedal (four-legged),herbivorous armadillos with armoredcarapaces (top shell) that were made of hundreds of interconnectedosteoderms (structures in dermis composed of bone). Other pieces of armor covered the tails andskull roofs, the skull being tall withhypsodont (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 relativeGlyptotherium[citation needed] orDoedicurus, the largest known glyptodont.Glyptodon is morphologically andphylogenetically most similar toGlyptotherium, however they differ in several ways.Glyptodon is larger on average[citation needed], with an elongated carapace, a relatively shorter tail, and a robustzygoma, or cheek bone.
Glyptodonts existed for millions of years, thoughGlyptodon itself was one of its last surviving members.Glyptodon was one of many South Americanmegafauna, with many native groups such asnotoungulates 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 withGlyptodon, including the "saber-tooth cat"Smilodon, the largecanidProtocyon, and the giant bearArctotherium.
Glyptodon, along with all other glyptodonts, became extinct at the end of the Late Pleistocene, around 12,000 years ago as part of theLate Pleistocene extinctions, along with most large mammals in the Americas. Evidence of hunting of glyptodonts by recently arrivedPaleoindians suggests that humans may have been a causal factor in the extinctions.
The history and taxonomy ofGlyptodon is storied and convoluted, as it involved confusion with other genera anddubious species, as well as a lack of detailed data. The first recorded discovery ofGlyptodon was as early as 1814 when Uruguayan priest, scientist, soldier, and later politicianDámaso Antonio Larrañaga (1771–1848) wrote about the discovery of several unusual fossils in hisDiario de Historia Natural, which included his descriptions of many new species of ants, birds, mammals, and even one of the first figures of the extinctMegatherium, a genus of giantground sloth that was named in 1796 by French scientistGeorges Cuvier (1769–1832).[3][4] This was the first recorded discovery of aglyptodontine or fossilcingulate.[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 thePleistocene aged (ca. 2.5-0.011 mya) strata on the banks of theSolís Grande Creek, Uruguay.[4][5] Larrañaga identified the fossils as those ofDasypus (Megatherium), believing thatMegatherium was a subgenus ofDasypus based on the incorrect referral of glyptodontine osteoderms toMegatherium 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 scientistAuguste Saint Hilaire about the discovery, and the letter was reproduced by Cuvier in 1823 in the second volume of his landmark bookRecherches 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 relatedHoplophorus.[8] The armoredMegatherium hypothesis was further supported later in 1827 when portions of aGlyptodon carapace, as well as a partial femur and some caudal armor, were found by aPrussian traveler toMontevideo, Uruguay named Mr. Sellow, who sent the carapace toBerlin where it was described by Professor von Weiss, who referred it toMegatherium.[9] The femur and caudal armor were recovered from theQuegnay in northern Uruguay, while the carapace had been found in theArapey River.[9][5] Weiss and other paleontologists noted that the osteoderms closely resembled those of armadillos likeDasypus, but Cuvier's hypothesis was popularized based on the incorrect referral of glyptodontine osteodermsMegatherium.[6][7]
Another work on the armoredMegatherium 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, andshoulder girdle. D'Alton recognized the great similarities of the fossils toDasypus and speculated that it was a giant armadillo, contrary to the notion that they were fromMegatherium. 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 mentionMegatherium 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 fromMegatherium 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 paleontologistPeter Wilhelm Lund published a series of memoirs on the fossils ofLagoa 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 andOrycterotherium 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 (originallyThoracophorus but the name was preoccupied by abeetle) in 1889 by Argentine paleontologistFlorentino Ameghino[12] based on isolated osteoderms now identified as those of a juvenileGlyptodon from Patagonia,[13] andLepitherium in 1839 byGeoffroy 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 ofTeleosaurus, a crocodile-like reptile known fromJurassic deposits in France.[15][14]
In 1838, British diplomatSir Woodbine Parish (1796–1882) was sent an isolatedmolariform and a letter about the discovery of several large fossils from theMatanza River inBuenos Aires, Argentina that dated to the Pleistocene.[16][17] Parish later collected several more fossils from localities inLas Averias andVillanueva; 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 theNatural History Museum, London, but the original fossils were destroyed afterGerman aerial bombing raids hit the college duringWorld War II from 1940 to 1941.[13][17]Glyptodon was named byRichard 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 bookBuenos 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 nameGlyptodon ("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 thelectotype byRobert 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 ofG. clavipes is that the material is undiagnostic and indistinguishable from otherGlyptodon species and evenGlyptotherium, making it dubious.
Cuadrelliet al (2018) designated the species aspecies 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, ofG. clavipes from theRiver 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 paleontologistThomas 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 ofGlyptodon by Owen:G. ornatus, G. reticulatus, G. tuberculatus, andG. clavicaudatus in 1847. Of these additional species, onlyG. reticulatus is still considered a valid species ofGlyptodon asG. ornatus was reassigned to the genusNeosclerocalyptus,[22]G. tuberculatus toPanochthus,[23] andG. clavicaudatus toDoedicurus.[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 forGlyptodon species after the 1840s, but many of them are now seen as dubious, species inquirenda, or synonymous with previously named species.[26][17] Par L. Nodot described a new genus and species of glyptodontine in 1857,Schistopleurum typus, on the basis of a caudal tube found in thePampas of Argentina, but it has since been synonymized withG. reticulatus.[27] 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 ofArroyo del Medio,San Nicolás, Argentina.[28][29] 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.[28][17] German zoologistHermann Burmeister described severalGlyptodon fossils in the 1860s, many of them he named as new species ofGlyptodon itself or the synonymSchistopleurum, all of which are now synonyms ofGlyptodon and its species.[30][17] In 1908, Florentino Ameghino named another species ofGlyptodon, G. chapalmalensis, based on a carapace fragment that he had collected from the Atlantic Coast of Buenos Aires Province that dated to theChapadmalalan. In 1932, A. Castellanos made a new genus forG. chapalmalensis,Paraglyptodon, which later included another species,P. uquiensis, that was based on more complete specimens that had been collected fromUquía, Argentina between 1909 and 1912.[31][32] The former species is dubious, but likely notGlyptodon based on its age.[33]P. uquiensis has been synonymized withGlyptodon and is possibly a valid species, though further analysis is necessary to settle its status.[25][17]
In the 1950s, Argentine paleontologistAlfredo Castellanos (1893–1975) erected new generic names for several species ofGlyptodon, the first beingGlyptocoileus and second of these beingGlyptopedius in 1953 that was made for the speciesG. 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 speciesG. reticulatus to the genus, but this unsupported.[17] Yet another genus was erected in 1976 namedHeteroglyptodon genuarioi by F. L. Roselli based on an incomplete skeleton that had been collected from the Pleistocene agedLibertad Formation inNueva Palmira, Uruguay,[34][35] but it has since been found to be an indeterminate specimen ofGlyptodon.[35] SeveralGlyptodon fossils from Pleistocene deposits in Colombia were described in 2012, extending the known range of the genus north greatly.[36]
AnotherGlyptodon species was described in 2020 calledG. jatunkhirkhi by several authors led by Argentine zoologistFrancisco Cuadrelli on the basis of an individual preserving a nearly complete carapace, several caudal rings, and a pelvis that had been collected fromYamparaez, 24 kilometres (15 mi) southeast of the Bolivian city ofSucre. The strata they were found in was made up of fluvial, sandy sediments that dated to theLate 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 inEastern Cordillera, Bolivia including a nearly complete skull and several osteoderms.[26] In a phylogenetic analysis conducted by Cuadrelliet al., 2020,G. jatunkhirki was recovered as the most basalGlyptodon species, despite being the same age as the more derived speciesG. clavipes. Reassessment ofGlyptodon 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 speciesG. reticulatus, G. munizi, andG. jatunkhirkhi definitively valid;G. clavipes andG. uquiensis asspecies inquirendas.[17] However a 2016 review ofG. uquiensis determined thatG. uquiensis was actually a juvenile specimen ofGlyptodon, though the species could not be determined.[33]
Fossils from North America were also described and referred toGlyptodon 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 nearTequixquiac,Mexico, the fossils coming from theRancholabrean stage of the Pleistocene.[37][25] These fossils were the first found of glyptodontines in North America and were named as a new species ofGlyptodon,G. mexicanum, but the fossils have since been lost and the species was synonymized withGlyptotherium 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 genusGlyptotherium, which was named in 1903 by American paleontologistHenry Fairfield Osborn.[39]
Glyptodon is the type genus of Glyptodontinae, an extinctsubfamily of large, heavily armored armadillos that first evolved in theLate Eocene (ca. 33.5 mya) and went extinct in theEarly Holocene during theLate Pleistocene extinctions (ca. 7,000 years ago).[26][40] Owen recognized thatGlyptodon was anedentate, 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 byJohn Edward Gray, who included the generaGlyptodon, Panochthus, andHoplophorus 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 ventralplastron (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 thatGlyptodon was descended from earlier genera.[12][28]
Glyptodontinae was classified in its own family or even superfamily until in 2016, whenancient DNA was extracted from the carapace of a 12,000 year oldDoedicurus specimen, and a nearly completemitochondrial genome was reconstructed (76xcoverage). Comparisons with those of modern armadillos revealed that glyptodonts diverged fromtolypeutine andchlamyphorine armadillos approximately 34 million years ago in thelate Eocene.[42][40] This prompted moving them from their own family, Glyptodontidae, to the subfamily Glyptodontinae within theextantChlamyphoridae.[40] 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.[42] Chylamyphoridae is a group in the order Cingulata, which includes all extant armadillos in addition to other fossil groups likePachyarmatheriidae andPampatheridae. Cingulata is itself within the basal mammal group Xenarthra, which includes an array of American mammal groups likeVermilingua (anteaters) andFolivora (sloths and ground sloths) in the orderPilosa. The following phylogenetic analysis was conducted by Frédéric Delsuc and colleagues in 2016 and represents the phylogeny of Cingulata using ancient DNA fromDoedicurus to determine the position of it and other Glyptodonts:[42][40]
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 theMiddle Miocene (ca. 13 mya) withBoreostemma,[45] but split into two genera,Glyptodon in the south andGlyptotherium in the north,[36] thoughGlyptotherium also lived in some areas of South America like Venezuela and eastern Brazil.[46][47][36]Glyptotherium andGlyptodon lived during the same intervals and are nearly identical toGlyptodon in many aspects, so much so that the first fossils ofGlyptotherium to be described were misidentified as those ofGlyptodon.[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 likeDoedicurus, Hoplophorus, andPanochthus.[50][26]
Below is the phylogenetic analysis conducted by Cuadrelliet al., 2020 of Glyptodontinae, with Glyptodontidae as a family instead of subfamily, that focuses on advanced glyptodonts:[26]
| Chlamyphoridae |
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||
Like the extant armadillos and all other glyptodontines,Glyptodon had a large, bony carapace that covered much of its torso, as well as smallercephalic 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), withGlyptodon carapaces preserving a total of 1,800 osteoderms each. The anatomy of differentGlyptodon species varies greatly, mostly in the speciesG. jatunkhirkhi which is more similar toGlyptotherium in certain aspects.[26]
In theaxial skeleton, glyptodontines had strongly fusedvertebrae andpelves 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 andGlyptodon'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 likeNeosclerocalyptus andDoedicurus.[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 ofGlyptotherium 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] andG. munizi weighed 1,150 kilograms (2,540 lb).[56] A partial skeleton ofG. 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 likeG. munizi andG. reticulatus measured 2.2 metres (7.2 ft) and 2.19 metres (7.2 ft) long respectively.[25]
Glyptodont dentition contains entirelyhypsodont 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; themaxilla andpalatine are enlarged vertically to make space for the molariforms, while the braincase is brachycephalic, short and flat.[59] InGlyptodon 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 orforamina. Sutures separating osteoderms are well marked, as inPanochthus.[28][60] Other Pleistocene glyptodontines are known by complete/sub-complete skulls, allowing for comparisons toGlyptodon.[13][43]Glyptotherium'szygoma are narrow, slender, almost parallel, and close to thesagittal plane in frontal view; inGlyptodon, this structure is broader, robust, divergent rather than parallel and more laterally placed.[51][25]
The nasal passage was reduced with heavymuscle attachments for some unknown purpose. Some have speculated that the muscle attachments were for aproboscis, ortrunk, much like that of atapir orelephant. The lower jaws were very deep and helped support massive chewing muscles to help chew coarse fibrous plants. Some paleontologists have proposed thatGlyptodon and some glyptodontines also had aproboscis or large snout similar to those inproboscideans andtapirs,[61] but few have accepted this hypothesis.[43][62] Another suggestion, made by A.E. Zurita and colleagues, is that the large nasal sinuses could be correlated with the cold arid climate ofPleistocene South America.[62][63] 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, theinfraorbital foramina are narrow and not visible in anterior view inGlyptotherium, but inGlyptodon they are broad and clearly visible in anterior view. In lateral view, the dorso-ventral height between theskull roof and thepalatal plane inGlyptodon decreases anteriorly, contrary toGlyptotherium; the nasal tip is in a lower plane with respect to thezygomatic arch inGlyptodon, but inGlyptotherium is higher than the zygomatic arch plane. The 1st molariform (molaiform is abbreviated as mf#) ofGlyptodon is distinctlytrilobate (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.[25] These traits separate the two genera.[25] Within the genusGlyptodon this trait varies as well, withG. reticulatus having triloby to a greater degree thanG. munizi.[17]
Themandibles ofGlyptotherium andGlyptodon are very similar, butGlyptotherium's mandible is smaller by about 10% in total size. The angle between theocclusal plane (part of the jaw where upper and lower teeth contact) and the anterior margin of the ascendingramus is approximately 60 inGlyptotherium, while it is 65° inGlyptodon. The ventral margin of the horizontal ramus is more concave inGlyptodon than inGlyptotherium. Thesymphysis area is extended greatly inGlyptotherium antero-posteriorily compared toGlyptodon. The mf1 is ellipsoidal inGlyptotherium and the mf2 is "submolariform", while inGlyptodon both teeth are trilobate.[25]
)_BHL39943960.jpg)
Glyptodon has 7cervical vertebrae, of which the first 3 cervicals were fused together while the rest of the cervicals were free except for the 7th.[5] The 7th cervical and the first 2 dorsal vertebrae were fused together into a trivertebral, a broad, flat bone with very smallspinous processes (projections from a vertebra) and large articular surfaces that held ribs.[5] 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.[5]Spinous processes in these vertebrae are also heavily reduced, with some being only a thin blade of bone ankylosed with other vertebrae.[5]Sacral vertebrae inGlyptodon are also fused and 13 in number, which preserve very unusual oval-shaped, thin, and slightly concave ends on the centra.[5] The pelves are also unusual, as they preserve giantilia and are fused to the rest of the skeleton.[5]
Glyptodon's osteoderms were attached bysynotoses (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 justconical. The sulci between these raised structures were deep and wide with parallel lines.[64] The carapace ofGlyptodon was strongly elongated compared to those ofBoreostemma andGlyptotherium, with the carapace being relatively 65% longer than the former and 14% than the latter. InGlyptodon, the top-bottom height of the carapace represents 60% of its total length, whereas inGlyptotherium 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 mostGlyptodon species, while inGlyptotherium andGlyptodon jatunkhirkhi it was at the center of the midline. The carapace of most species ofGlyptodon is arched subtly, whileGlyptotherium andGlyptodon 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 inGlyptotherium they are less ankylosed and more flexible.[25] The osteoderms of the caudal aperture (large conical osteoderms that protect the base of the tail) are more conical inGlyptodon and more rounded inGlyptotherium, though in the latter the anatomy of the caudal aperture osteoderms varies by sex while inGlyptodon it varies by age.[25][65] The caudal aperture is more vertically oriented in the latter genus, while inGlyptotheirum it is angled posteriorily.[25] Although frequently used to differentiate the two taxa,Glyptodon andGlyptotherium have similar osteoderm morphologies that differ only in several areas. Both genera have tall, thick osteoderms compared to those of many other glyptodontines such asHoplophorus andNeosclerocalyptus. Glyptodon sometimes preserves a "rosette" pattern, where the osteoderm's central figure is surrounded by a row of peripheral figures, while other specimens lack them completely.G. reticulatus varies from a complete rosette pattern to a reticular surface, which has convex central and peripheral figures.[65][25]Glyptotherium however always preserves rosettes.[65][25] The central and radialsulci are deeper and broader inGlyptodon (ca. 4–6 mm) than inGlyptotherium (ca. 1–2.4 mm). The osteoderms inGlyptodon andGlyptotherium have 5-11 peripheral figures, rugose exposed surfaces, and heights up to 47 millimetres (1.9 in).[25]
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.[30] This hypothesis has since been disproven, but in the early 2000s, the presence of osteoderms onGlyptodon's face, hind legs, and underside was confirmed in several species. The fossils with these characteristics were from the Pleistocene, evolving in younger species likeG. reticulatus . These small to medium-sized ossicles were actually embedded in the dermis and did not connect in a pattern.[64]
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 conicaltubercles by the third caudal ring. The more posterior the rings were, the larger they were, with the exception of the 2nd ring which was the largest and 1st complete ring in the series, creating a cone-shaped tail. The distal scutes are larger, and their free margins are rounded producing a fan-like shape. Most of the osteoderms of the distal row (some individuals preserving up to 12) bear prominent conical outlines, in stark contrast to more advanced glyptodontines likeDoedicurus andPanochthus, which had completely fused tails that formed an inflexible mace or club.[66] The caudal tube at the distalmost end of the tail is cylinder-shaped with smaller conical osteoderms and is stubbier proportionally inGlyptodon. InGlyptotherium, this caudal tube represents ca. 20% of the total length of the caudal armor, whereas inGlyptodon, this structure represents 13% of the total length.[25] InGlyptodon, the caudal armor length represents circa 30-40% of the carapace's total length in contrast toGlyptotherium, where this value is greater at around 50%.[25] For example, in specimen MCA 2015 ofGlyptodon reticulatus, the terminal tube measured only 73.23 millimetres (2.883 in) long in comparison toGlyptotherium texanum specimen UMMP 34 826's 210 millimetres (8.3 in) long tube.[25]
.jpg)
Many armadillo species have digging capabilities, with large claws adapted for scraping dirt in order to make burrows or forage for food underground.[67][68] Much of armadillo diets consist of insects and other invertebrates that live underground,[69] in contrast to the herbivorous diets ofGlyptodon and related genera.[70] Being a large armadillo,Glyptodon's fossorial capabilities have been researched on several occasions. Owen (1841) opposed this idea,[16] though pushback came from Nodot (1856) and Sénéchal (1865) who believed digging was possible for the genus.[27][71] However, the evolution of a rigid carapace as opposed to a flexible one in extant armadillos as well as a weakly developeddeltoid 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.[55][72]
Several complete skulls ofGlyptodon enable the endocranial anatomy to be analyzed, as well as compared to other well-preserved taxa likeDoedicurus andPanochthus. The brain cavities of the larger glyptodontinesGlyptodon, Doedicurus, andPanochthus had a braincase volume of 213 to 234 cubic centimeters (7.2 to 7.9 U.S. fl oz). Theencephalization 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 extensiveolfactory bulb that took up between 4.8 and 9.7% of the entire brain, while around two thirds of it were occupied by thecerebrum and the rest by thecerebellum. 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 pronouncedsulcus praesylvianus.[73][74]
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 asankylosaurs andstegosaurs, two types of armoreddinosaur. 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 likeGlyptodon.[73][74]
Two main groups of glyptodontines can be distinguished by their feeding habits: narrow-muzzled Miocene propalaehoplophorids and wide-muzzled post-Miocene glyptodontines.[75] The propalaehoplophorids were selective feeders, while the post-Miocene glyptodontines werebulk 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.[76]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.[76] They had a well-developed snout musculature, along with a mobile neck region that helped them secure food.[77] The hyoid shows a robust design that suggestsGlyptodon had a large and robust tongue, which may have aided in food intake and processing.[78][79]
Like most other xenarthrans, glyptodontines had lower energy requirements than contemporary mammal groups.[80] 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.[81] This in conjugation with the proposed idea that aquatic grazing may have caused the isotopes strongly associated with herbivory observed inGlyptodon fossils.[81] However, aquatic grazing inGlyptodon is little supported[82] though more backing for this hypothesis has been found in the relatedGlyptotherium.[83][84][38] A carbon isotopic analysis ofGlyptodon bones by Françaet al (2015) found that it consumed a variety of bothC3 plants andC4 grasses at lower latitudes while it ate exclusively C3 grasses at higher ones, implying an ecological shift based on the climate. A 2012 analysis of isotopes supports this, but the isotopic results are not backed by morphological evidence.[76] The isotopic conclusion would placeGlyptodon as a mixed browser in most environments, similar to some other glyptodontines.[85][86] The 2012 paper also noted thatGlyptodon may have had a more flexible diet than previously imagined,[87] with a mix of slightly wooded and slightly open habitats as implied by the consumption of C3 and C4 material.[86][56] The C4 plants include groups likePoaceae,Cyperaceae,Asteraceae, andAmaranthaceae based on palynological evidence, meaning thatGlyptodon likely ate C4flowering plants in addition to C3 grasses.[88][85] A mesowear analysis supported their conclusion, however, finding that mixed-feeding causing blunt wear that suggests a more abrasion-dominated diet. This is similar to that ofNeosclerocalyptus, but in contrast toHoplophorus which had sharper wear ends.Neosclerocalyptus favored more open environments despite this, as found by isotopic studies.[43][56] The mesowear angles ofGlyptodon were noted to possess abimodal distribution, implying a difference between populations, sexes, or species in diet.[70]
Glyptodonts are believed to have taken part inintraspecific fighting. It was presumed that since the tail ofGlyptodon 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 ofGlyptodon was primarily for attacks on its own kind. AG. reticulatus fossil displays damage done on the surface of its carapace. A study based on this specimen calculated thatGlyptodon tails would have been able to generate enough force to break the carapace of anotherGlyptodon. 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.[89]
In 2009, a partial skeleton of a prenatal individual ofGlyptodon was described that had been found inside of the pelvic region of a carapace of an adult.[90] The skeleton had been collected from the Pleistocene-aged deposits in theTarija 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 immatureGlyptodon known, though dozens of isolated osteoderms from juveniles are known.[90] The preserved skull measures only 51 mm long, but still bears many characteristics ofGlyptodon such as a subtriangularnaris, a lateral margin on the naris that forms an acute angle of 30-degrees, oval infraorbital foramina, and several other traits.[90] However, the mandible differs in that the ascending ramus is at a 90-degree angle in contrast to the 60-70 degree angles preserved in adults. Interestingly, this mandibular morphology is alike to that in some specimens ofGlyptotherium cylindricum.[60][90]
In the osteoderms of juvenileGlyptodon 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.[17][25]
Several interpretations of glyptodontine posture have been made,[91] starting with those by Richard Owen in 1841 using comparative anatomy.[16] Owen theorized that the phalanges were weight-bearing due to their short and broad physiology, in addition to evidence provided in the postcranial skeleton.[16] It was also proposed that an upright posture was possible forGlyptodon, first by Sénéchal (1865) who stated that the tail could be an equilibrium for the front half of the body as well as a method of supporting the legs.[71] Linear measurements were later taken which provided insight into this hypothesis, finding thatbipedalism would be possible.[92][93] Thepatellar articulation with the femur suggests rotation of the lower leg during knee extension and potentially even knee-locking were feasible.[94]
No evidence ofsexual dimorphism inGlyptodon has been described, but it has been observed in the close relativeGlyptotherium 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 newbornGlyptotherium, the marginal osteoderms are either conical or flat, which enables their sex to be determined.[65] No direct evidence of glyptodontine group behavior has been described, though some localities preserving juveniles, subadults, and adults ofGlyptotherium together are known.[65][51] Living armadillos are loners and only come together during mating season, with the number of offspring varying between one and even twelve babies depending on the species.[95]
Glyptodon is one of the most common Pleistocene glyptodontines with a large range from the lowlandPampas to the toweringAndean Mountains of Peru and Bolivia, some fossils found at elevations reaching over 4,100 metres (13,500 ft) above sea level.[96][97][26] OnlyG. munizi is found in the early-middle Pleistocene, whereas other species are younger.[28][26]G. reticulatus is specifically noted to be known from 60ka to as recent as 7ka possibly, though confirmed records only extend to 11 ka.[98] The genus had a generalist diet, which allowed it to fill niches in areas that were inaccessible by grazing genera, withG. reticulatus representing up to 90% of the glyptodontine fossils in the Tarija Valley of Bolivia.[13] However, in regions such as the Pampas, Mesopotamia, and Uruguay, an array of glyptodontines are known.[98][17] Further evidence ofGlyptodon's adaptability is found in the Pampas, which were semihumid and temperate from 30,000 to 11,000 ka, alternating between therainy anddry seasons, over a large area consisting mostly of grasslands dotted with forests and mixed shrubbery.[99][100][17] Temperatures in this region were lower than the present, with an estimatedmean annual temperature 4.2 °C (39.6 °F) in the Pampas compared to 16.4 °C (61.5 °F) in Buenos Aires today. The Pampas specifically was a mix of semi-arid Patagonian and tropical Brazilian climates during the middle Pleistocene before the expansion of the drier climates.[101] This is in stark contrast to theBermejo Formation ofFormosa Province, Argentina where the climate and fauna suggest a more arid environment with fewer grasslands.[102][103]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.[26][97]G. jatunkhirkhi is not the only example of this in Xenarthra, with species ofPanochthus andPleurolestodon evolving to be smaller in size in mountainous regions.[97][26]
During the Ensenadan and Marplatan,Glyptodon coexisted with a variety of mammals unique to the period such as thenotoungulateMesotherium, canidTheriodictis, and a species of the giant bearArctotherium.[101] In areas such as Uruguay, fossils ofGlyptodon have been unearthed alongside the contemporary glyptodontinesDoedicurus,Neuryurus,Panochthus; armadillosChaetophractus,Propaeopus, andEutatus; and the herbivorous pampatherePampatherium. As for their distant relatives the ground sloths, the giantMegatherium is known, in addition to two species of thescelidothereCatonyx, and themylodontid generaMylodon andGlossotherium. Some other groups are known, including the unusuallitopternsMacrauchenia andNeolicaphrium, notoungulateToxodon, massiveproboscideanNotiomastodon, and theequidsEquus neogeus andHippidion. Variousartiodactyls have been recorded, including the peccariesCatagonus andTayassu peccari, extinct deerMorenelaphus andAntifer, and two genera of llamas includingHemiauchenia andLama. A variety of carnivorans have been recorded, such as the "saber-toothed"Smilodon, the bearArctotherium bonariense, and the wolf-like canidsProtocyon, andDusicyon.[104][105] Rodents too have been found, such asHolochilus,Hydrochoerus (capybara),Cavia, andMicrocavia.[98] Notably, some of the youngest "terror-bird" fossils of the genusPsilopterus have been unearthed in the area.[106]
Material previously assigned toGlyptodon in northeast Brazil has been reassigned toGlyptotherium, restricting the Brazilian distribution ofGlyptodon to the southern provinces. However, two osteoderms with characteristics similar to those ofGlyptodon have been found inSergipe state in the northeast, suggesting that both genera occurred in this region during the Pleistocene.[47]Glyptodon's northernmost locality comes from Pleistocene deposits in central Colombia,[36] though many specimens formerly attributed to the genus come from the bordering country ofVenezuela.[51]
Glyptodon coexisted with a variety of large predators including the catSmilodon,jaguars, and canidProtocyon.[107][108] This belief is furthered by the discovery of fractured dorsal armor, which implies thatGlyptodon had been in physical conflict with other animals.[64] However,isotope analyses of thecollagen fromGlyptodon and other mammals of the Pampas region by Bocherenset al. (2015) discovered little evidence to support the idea of predators feeding onGlyptodon.[107] Instead, it was found thatGlyptodon as well as herbivorous mammals living in denser forests made up a smaller portion of carnivore diets, whereas open grazers such asLestodon andMacrauchenia were consumed more often.[107] Furthermore, the appearance of secondary armor in the dermis ofGlyptodon coincides with the arrival of North American predators in South America during theGreat American Interchange.[2] For this reason, it was hypothesized that the osteoderms developed as a defensive/offensive mechanism to combat the new arrivals of the area.[64]
Smilodon may have occasionally preyed upon glyptodontines, based on a skull ofGlyptotherium texanum which bears the distinctive elliptical puncture marks that best match those of themachairodont cat, indicating that the predator successfully bit into the skull through the armored cephalic shield.[109] TheGlyptotherium in question was a juvenile, with a still-developing head shield, making it far more vulnerable to the cat's attack.[110] Although originally theorized by George Brandes to be possible in 1900,[111]Smilodon canines could not pierce the thick carapace osteoderms of glyptodontines.[112] Brandes imagined that the evolution of thick glyptodontine armor and long machairodont canines was an example ofcoevolution,[111] but Birger Bohlin argued in 1940 that the teeth were far too fragile to do damage against glyptodontine armor.[112][64]
The coexistence of earlyhunter-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 ofGlyptodon 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 ofGlyptodon and the horseEquus. 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) glyptodontineNeosclerocalyptus,[115] the armadilloEutatus, and the gigantic (2 ton) glyptodontineDoedicurus, 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]
Glyptodon, along with all other glyptodonts became extinct around the end of the Late Pleistocene, as part ofa 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 thatGlyptodon's relativesDoedicurus andPanochthus 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 andDoedicurus); 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 ofGlyptodon, 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 ofGlyptodon notably coincides with the end of theAntarctic 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, withGlyptodon 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 asGlossotherium andMorenelaphus were wiped out, though several other groups lived for several thousand years after.[44][98]
