Glyptodontidae

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Glyptodontidae
Skeleton reconstruction with the tank of Glyptodon

Skeleton reconstruction with the tank of Glyptodon

Temporal occurrence
Middle Eocene to Lower Holocene
48 million years to 8,000 years
Locations
Systematics
Synapsids (Synapsida)
Mammals (mammalia)
Higher mammals (Eutheria)
Sub-articulated animals
Armored siderails (Cingulata)
Glyptodontidae
Scientific name
Glyptodontidae
Gray , 1869

The Glyptodontidae or Glyptodonten are an extinct family of the sibling animals (Xenarthra) and were common in South America and in some parts of North America . They are closely related to the armadillos still alive today and, like them, had well-developed body armor, which, unlike that of the armadillos, was rigid and had no movable ligaments. In addition, the tail was also fully armored, some shapes had a club-like, elongated end. The representatives of the family reached enormous proportions , especially in the Pleistocene, and weighed up to 2 tons, making them the largest members of the armored articulated animals . In addition to some features of the skull, the glyptodonts also had peculiarities in skeletal construction, including elephant-like hind legs and an extremely ossified spine. As possibly adapted grazers , they also had teeth with high crowns. The glyptodons mostly inhabited open landscapes, influenced by cool to tropical climates, and also occurred in higher mountain regions.

The first members of the family are known from the Middle Eocene around 48 million years ago and were discovered in Patagonia , which is probably the origin of the group. The most recent finds come from the beginning of the Holocene and are around 7500 years old. During the tribal history , there was a strong breakdown of the glyptodons into several lines of development. In the Pliocene around 3.5 million years ago, some representatives also reached North America through the formation of a land bridge, but they did not stay there for very long. Since often only remnants of the armor are fossilized , the systematic structure has not been adequately investigated and is largely based on the design and changes in shape of the individual elements of the back and tail armor. Overall, the glyptodons represent one of the most diverse groups of articular animals alongside the sloths .

The research history of the Glyptodont dates back to the second half of the 18th century and began with the discovery of a back armor in the Pampa region . Further finds followed at the beginning of the 19th century, the generic representative Glyptodon , which gave the family its name , was described in 1839 by Richard Owen . This was based on a partial skeleton from the pampas region south of Buenos Aires , which had been brought to England. Charles Darwin made an important contribution on his trip to South America, who hid numerous remains of glyptodons in the pampas. The first finds in North America were not made until the transition from the 19th to the 20th century. During the same period, Florentino Ameghino worked out a first systematic subdivision of the glyptodonts, which served as the basis for numerous other researches.

features

General and height

Live reconstruction of Panochthus

The Glyptodontidae resembled today's armadillos (Dasypoda), with which they are more closely related, but were significantly larger and had a short-nosed skull and a rigid, dome-like arched back armor and a fully armored, comparatively short tail. Early members of this family were still relatively small. Thus achieved Cochlops from the Lower Miocene kg a total length of about one meter and a weight of about 90 which is about the same age Eucinepeltus brought up to 115 kilos. Above all, the forms of the late Pleistocene had huge proportions in comparison. Panochthus was over 2.6 m long, while the weight was between 1.1 and 1.3 t. Larger proportions had doedicurus with a length of 3.3 m to 4 more and a height of 1.5 m. It weighed between 1.4 and 1.76 t, with a weight of 2.3 t being determined for a very late member of the same genus based on a fragmented humerus . The largest representatives of Glyptodon brought it to a length of over 3.5 m with a weight of around 2 t. This made the glyptodons the largest representatives of the armored collateral animals (Cingulata) and significantly larger than the related armadillos and Pampatheriidae .

Skull and dentition features

Skull of Glyptodon

The skull of the Glyptodon was massive and short and tall. As a result, it had a very characteristic structure, which together with the lower jaw almost resembled a cube . Only historically older forms still had a somewhat elongated rostrum and thus resembled today's armadillos. Smaller forms such as Eosclerocalyptus had skull lengths of about 22 cm, while large ones such as Panochthus were up to 42 cm long, well over twice the length. The markedly shortened rostral area was particularly characteristic , which made the entire skull seem telescopically pushed together. Another striking feature was the extremely massive zygomatic arches , which protruded far and reached a distance of 31 cm in Panochthus . A second outgrowth of bone proved to be conspicuous here, reaching from the front attachment of the zygomatic arch far down to below the row of teeth of the lower jaw and narrowing in front and back. A similar formation can be observed in the sloths , but this is flattened laterally. In both cases, the outgrowth serves as the attachment point for the masseter muscle of the masticatory muscles. Since the muscle is arranged differently in the two groups, one assumes that the bony process is independent in each case. In addition, the Glyptodonts have other skull features that distinguish them from other mammals. Above all, the bone sections in the area of ​​the rostrum, such as the upper jaw and the palatine bone , showed vertically aligned extensions to accommodate the extremely high-crowned ( hypdodontic ) teeth. The frontal sinuses were also greatly enlarged, which may have played a role in chewing the food. The lower jaw was solidly designed, extremely high on the horizontal bone body due to the teeth and provided with a robust symphysis . The end of the joint rose steeply, far beyond the level of the row of teeth. Due to the unusual construction of the skull, the glyptodonts had the highest point of attachment for the joints of the lower jaw (craniomandibular connection) within the mammals in relation to the length of the skull.

As with all articular animals, the structure of the dentition differed from that of the other higher mammals . Incisors and canines were not developed. The posterior teeth were designed in a homodontic manner so that no distinction can be made between premolars and molars . In principle there were eight molar-like teeth in each half of the jaw, so the dentition had a total of 32 teeth. A peculiarity was the lack of tooth enamel , which is also a general characteristic of the secondary articulations. The teeth are largely made of orthodentine and osteodentine , the two hardest variants of the dentin . With the exception of the two anterior teeth, the chewing surfaces of all molars had three transverse flap-like ( trilobate ) formations. Each individual lobe consisted of a shell made of clearly mineralized orthodentine (originally also described as dental cement ) and an interior made of less hardened dentin. In the center of each a small, transverse rib made of very hard osteodentine rose, with a bar running along the central axis of the tooth connecting the three ribs. Thus, the teeth of the glyptodons differed from those of today's armadillos with their generally pen- or nail-like shape. On the other hand, they were similar to the molars of the Pampatheriidae, also armor-bearing relatives of the armadillos, who had two transverse lobes . In general, the teeth were very high crowned and rootless and grew as a result, which, as with other articular animals, did not lead to a change of teeth. In contrast to other groups of higher mammals, only a few cases of the formation of supernumerary teeth are known in glyptodonts. One of the few examples is from an individual of the genus Boreostemma from the Middle Miocene who had an additional anterior tooth in the upper jaw.

Body skeleton

Shell and skeleton of Glyptodon

Special skeletal features are found on the spine. The fusion of the vertebrae of the cervical spine is characteristic of all armored articulated animals, whereby the atlas (foremost cervical vertebra) is usually free to move, but the rear ones represent a fused bone structure. In addition, the second, third and fourth thoracic vertebrae were firmly fused in the Glyptodonts ( trivertebral element ). This was followed by a bony tube consisting of nine thoracic vertebrae. The number of lumbar vertebrae varied within the glyptodont at the species level, representatives of the North American genus Glyptotherium showed between 5 and 9. However, the vertebrae of the lumbar spine were also firmly fused, with which the glyptodons showed the highest degree of vertebral fusions among all mammals. As a result, the xenarthric joints (secondary joints or xenarthrals) typical of the secondary articular animals and giving their name did not appear on the lateral articular processes of the lumbar and posterior thoracic vertebrae, which is to be regarded as a unique feature .

The musculoskeletal system showed particularities, especially on the hind legs. These were designed similar to today's elephants and had adaptations to an extremely heavy physique. In terms of proportions, the glyptodons surpassed today's elephants, making them one of the land vertebrates most adapted to a difficult gait. Furthermore, the basin was completely fused with the tank and was therefore immobile. At the femur , a third trochanter as muscle attachment point formed from typical for Xenarthra, but was very far down and continuously passed into the lower joint end. The tibia and fibula were firmly fused at the ends. The hands and feet had a relatively original structure with no major specializations. The respective first phalanges (finger and toe phalanges) were significantly reduced. Fingers and toes ended in hoof rather than claw-like formations. Overall, the front and rear feet were reminiscent of those of the elephants. Early glyptodonts each had five-pointed hands and feet ( pentadactyl ). The later forms reduced the innermost ray (ray I) at the autopodia and therefore had four-rayed hands and feet ( tetradactyl ), with the exception of Glyptodon and its close relatives, who retained the pentadactyl hind foot.

tank

Body and tail armor from Glyptotherium
Osteoderms of Propalaehoplophorus , the central pattern and the smaller patterns arranged concentrically around it are clearly visible
Propalaehoplophorus osteoderms from various areas of the dorsal and tail armor with important names

The armored secondary articulated animals are the only mammals in which an external, bony armor is formed. In contrast to today's armadillos, the armor of the back of the glyptodons was rigid and immobile. It covered the entire trunk of the animals and consisted of a mosaic of several hundred to 1800 individual bone platelets ( osteoderms ) that formed in the skin and were mostly covered with horn , as in today's armadillos . The shape of the carapace varied and in Doedicurus was roughly arched like a dome, in Glyptodon less distinct and in Neosclerocalyptus it was strikingly flat. In larger forms such as Panochthus and Glyptodon , the carapace measured between 127 and 194 cm in length, taking into account the curvature, the values ​​amount to 146 to 220 cm. The bone platelets also showed a different design depending on the species, but they often had a polygonal outline. Sometimes they had a conspicuous rosette-like ornamentation on the surface with a more or less centrally located pattern, around which more were arranged concentrically in one or more rows. The individual patterns were separated from one another by furrows ( sulci ). In cross-section, the osteoderms had a multilayer structure, analogous to those of the armadillos: Inside and outside there was a solid layer of bone, in between a space with air-filled chambers in which sweat and sebum glands and hair follicles were possibly also embedded. This interior space takes up up to half the volume of the entire bone plate. In the Glyptodon it was built up much more regularly than in comparison to the armadillos.

Tail lobe of Panochthus

The skull was also protected by a head shield that covered the head like a helmet. In addition, osteoderms have also been detected in some representatives on the abdomen and legs, but they were irregularly shaped and not fused together. The rather short tail was also fully armored. The armor here consisted of several bony rings that were loosely connected to each other, giving the tail a high degree of flexibility. Each ring was made up of two or three rows of bone platelets. In some genera, such as Glyptodon , the tail had a relatively short end. Other forms such as Doedicurus , Panochthus or Castellanosia only had rings in the front area. In contrast, the osteoderms of the rear part of the tail were completely fused together and completely enveloped the tail. They formed a tubular, bony club that could be up to over a meter long and had a diameter of up to 30 cm. Sometimes the end was also thickened, but there were often oval depressions with a roughened surface on the surface. In the case of Hoplophorus and Panochthus , conical bone formations protruded from these depressions; in the case of other representatives, some scientists assume that originally thorn-like spines made of keratin were formed there. The tail design is unique among mammals and represents a convergent development to the ankylosaurs . These convergences relate to the stiffening and thickening of the rear tail section, but also the partial agglomeration of the vertebrae and the formation of a bony skin as well as an enormous increase in body weight.

distribution and habitat

The Glyptodontidae family lived exclusively on the American double continent . It originated in the Eocene in South America , possibly in the region of today's Patagonia , and from there gradually spread to the north and south. The finds of the Santa Cruz Formation in the transition from the Lower to the Middle Miocene in the south of the continent are significant here , the numerous finds of which indicate that the glyptodons lived in a mosaic of open and partly closed landscapes in their early development phase. In the Pliocene they reached North America for the first time , which was made possible by the closure of the Isthmus of Panama and the creation of a land bridge, after which the Great American Fauna Exchange began. As a result, the glyptodons had their largest distribution area in the Pleistocene and then came from South America from the 49th parallel south to North America around the 36th north. At that time they inhabited largely open landscapes in both cool to cold, dry, desert-like regions, but also occurred in tropical areas. In addition, they were also found in the Andes region at up to more than 3300 m, in some cases up to more than 4000 m, above sea level, which is one of the highest records of articulated animals. A way of life in swampy areas or floodplain landscapes , sometimes adopted for some forms, is usually rejected with reference to the construction of the limbs and the adaptations to open landscapes.

Paleobiology

Locomotion

Life reconstruction of Glyptodon

With a few exceptions, such as the spherical armadillos, today's armadillos are good graves, which are adapted to a burrowing way of life with specialized forefeet and robust forelimbs. These include the strong and partially elongated middle finger and the very extensive upper articular process of the ulna , the olecranon . It is assumed that the ability to dig was originally developed in all armored articulated animals and that it was only later adapted to the purely ground-dwelling ways of life. A large part of the Glyptodontidae, including the early representatives such as Propalaehoplophorus , but also numerous later forms have a moderately built olecranon that does not indicate specialized graves. The structure of the hand is also rather original and therefore different from that of the armadillos. The extremely large late forms such as Glyptodon and Doedicurus have massive upper ulnar joint processes that are up to 11 cm long with a bone length of 24 cm. Here it is argued that this strongly built forearm is an adaptation to the very high body weight. It also served to cushion the body when an animal returned from a two-footed position to a four-footed position. The fact that the glyptodonts were able to stand up on their hind legs is shown above all by the extremely strong bones of the hind legs, which, according to histological examinations on limb elements of Lomaphorus, are significantly more compact than the front legs. In addition, the center of the body mass is very far back on the torso and thus supports the straightening. Today's armadillos and anteaters are also able to change into a two-legged position, which is mainly used for foraging and defense.

Direct evidence for the locomotion of the glyptodons in the form of trace fossils is so far little known. The most extensive came to light in Pehuén-Có near Bahía Blanca in the Argentine province of Buenos Aires . This site, which was discovered in 1986 and is around 12,000 years old, contains a large number of footsteps from a wide variety of mammals and birds on an area of ​​around 1.5 km², which have been pressed into an originally soft substrate. Underneath there are rounded prints with five or three short toes. In shape and size they match the anatomical reconstruction of the feet of Glyptodon , whereby the five-rayed prints are interpreted as hind feet, the three-rayed ones as forefeet (the forefeet of Glyptodon are actually four-rayed, the outer ray V protrudes to the side and is small, so that he may not have left an imprint). The dimensions of the rear footprint are 18.5 × 18.5 cm, those of the front feet 17 × 10 cm. The finds are assigned to the trace genus Glyptodontichnus . Some researchers explain the rarity of glyptodon trace fossils in contrast to the rich anatomical fossil report by saying that the large animals avoided landscapes with soft subsoil due to the greater risk of injury.

nutrition

Due to the enamel loose teeth no abrasion can be identified, nor is there any remains of food in the form of coprolites to examine more closely to the diet. The extremely high-crowned teeth may lead to a diet mainly derived from grass. For early phylogenetic representatives, however, mixed vegetable food is also assumed to be the main food. This can be deduced from the relative width of the snout, among other things. Today's grass-eating ungulates usually have a comparatively wide mouth compared to the narrow-snouted, more selective leaf-eaters. For numerous early Glyptodonten from the Miocene as Propalaehoplophorus or Cochlops a rather narrow can Rostrum be reconstructed during this late in forms from the Pleistocene as doedicurus or Panochthus was significantly wider. The former two therefore ate more picky than the latter. Similar to Propalaehoplophorus and Cochlops , Glyptodon is also to be assessed , whose snout was generally narrower than that of its Pleistocene contemporaries. In the structure of the teeth, glyptodonts resemble cattle and African rhinos , the former lacking incisors in the upper jaw, the latter also in the lower jaw. Both groups use their flexible lips to take in food, which can also be assumed in the case of the glyptodons. Due to the special design of the skull with the short snout, a different chewing apparatus has developed in the Glyptodonts. The shortening of the skull resulted in the cranium protruding over the back half of the teeth, a condition that does not occur in the related armadillos and leads to a different arrangement of the masticatory muscles.

In comparison to the body size of the animals, the total available chewing surface of the teeth is also rather small. With a body weight of approximately 260 kg , Plohophorus had a chewing surface of almost 900 mm², a plains zebra of about the same size , on the other hand, has 2600 to 2830 mm², the flatland tapir ranges between 1930 and 2240 mm². For Doedicurus , a chewing surface of around 1800 mm² with a body weight of around 1.5 t can be determined. Comparative values ​​for the Java rhinoceros of roughly the same size are around 6250 mm². Due to these features and in connection with the special design of the lower jaw - the rearmost tooth lies directly below the lower jaw joint due to the obliquely forward joint branch and not, as usual, clearly in front of it - it results that glyptodons were obviously less able to chew their food. As the food was reduced to a lesser degree, but the sometimes enormous size of the animals meant that correspondingly large amounts of food had to be consumed, scientists assume an overall low metabolic rate for these animals. However, the hyoid bone shows a robust design that allows a very well defined and flexible tongue to be assumed, which possibly supports the food intake and the digestive process in the mouth.

Social behavior

Little is known about the social and reproductive behavior of the glyptodons, today's armadillos are solitary and only come together during the mating season. The litter size is species-specific and varies between 1 and 3 young animals for most representatives and up to 12 for some members of the long-nosed armadillos . Glyptodon juveniles are partly well documented. Osteoderms found with these are very thick and have a poor surface pattern. This indicates that, analogous to today's armadillos, the armor was only fully formed and hardened after birth. Finds of unborn individuals in the back armor of old animals are extremely rare. One example was discovered in Monte Cercado in southern Bolivia and belongs to Glyptodon , another is from the Sopas formation in Uruguay and represents Neuryurus . Individual teeth had already erupted on the lower jaw of the first-mentioned find, but whether glyptodonts were born fully dentate is unclear.

Function of the tank and defense

It is considered likely that the tank did not originally evolve due to the threat of predators, but possibly acted as a shelter from prickly vegetation. Only secondarily did it protect against larger predators, which at that time were provided in South America by the Phorusrhacidae ("terror birds") and predatory baggers such as the representatives of the Borhyaenidae and the Thylacosmilidae . As a secondary function, it probably served as the closure of a fat store above the chest area, similar to the humps of camels . Here the carapace is not connected to the spine or the ribs and rises freely above it, which gave Doedicurus , among others, a free space with a height of around 25 cm. Since this area is unlikely to have been stressed by muscles, experts assume a fat pad here.

Different tail designs in Doedicurus (left) and Glyptodon (right)

The long and massive tail probably also served as a counterweight for locomotion, which was necessary due to the rigid spine and the pelvis that was firmly attached to it. However, some representatives, where the end was overgrown like a club, could also use it as a weapon. Strong muscles were developed at the base of the tail, as indicated by the massive extensions of the vertebrae and the large diameter of the armor rings. This alone weighed calculations based on the size of the tail in Panochthus around 74 kg, in Doedicurus possibly up to 108 kg. The club-like end is 89 and 105 cm long, respectively, with a weight of 30 and 65 kg. With the existing muscle mass, the club-like end could release up to 3000  J of energy selectively with a blow (this corresponds roughly to the amount that a shot putter needs to push the 7.3 kg ball 16 m far), the point of impact in the back of the tail lobe. Some armor showing fractures, such as a Doedicurus armor with healed scars about 35 cm in diameter , also indicate such use of the tail in intraspecific combat . In some forms, which like Glyptodon did not have a club-like tail end, there were up to three rows of highly modified bone platelets on the edge of the body armor, which enclosed it, but were not firmly attached to it and had a pointed-conical shape. These osteoderms may have had a protective function for particularly important parts of the body such as the neck.

It is unclear whether the glyptodons also use their tails against predators, but today's forehead weapon bearers as a comparison use their horns and antlers only extremely rarely to ward off predators. At least for the largest representatives, such a threat is more likely to be excluded in adulthood. From North America only the skull of a young individual of Glyptotherium from the Pliocene is known, whose head armor was not yet fully developed and which has bite marks that led to the death of the animal. From the Argentine province of Buenos Aires, on the other hand, a tank with remains of the body skeleton of Eosclerocalyptus was found, which also dates to the Pliocene and whose vertebrae in turn have bite marks. These probably come from a small bear , possibly from Chapalmalania , which is fossilized in the region at that time. However, they probably emerged only after the death of the glyptodon and thus go back to scavenging.

brain

Castings of the brain capsules of Glyptodon (left) and Doedicurus (right), after Paul Gervais 1869

Several full skulls allow the structure of the brain to be studied. For large representatives such as Glyptodon , Doedicurus and Panochthus with body weights of 1.2 to 2 tons, pourings from the brain capsule give a volume of 213 to 234 cm³, for smaller ones like Pseudohoplophorus , which weighed only a little more than 200 kg, of 101 cm³. The encephalization quotient is 0.12 to 0.4, with the smaller Pseudohoplophorus having the highest value. The data are in the lower range of today's armadillos (0.44 to 1.06) and also correspond to those of the pampatheria . The glyptodon brain had an extensive olfactory bulb that took up between 4.8 and 9.7% of the total brain . Around two thirds were also taken up by the cerebrum and the rest by the cerebellum . In general, this corresponds to the armadillos, but in the latter the cerebrum is relatively more voluminous and the cerebellum less extensive. In contrast to the armadillos with their wide olfactory bulb, the glyptodonts had long, narrow and pointed, similar to the pampatheria. Corresponding to the armadillos, the relief of the cerebral cortex was relatively simple. Only the sulcus suprasylvianus of the parietal lobe could be made out of furrows , which also applies to the pampatheria. The sulcus praesylvianus, which was additionally formed in the armadillos, was missing.

In general, today's armadillos have relatively smaller brains than anteaters and sloths . The reasons for this are not clear. On the one hand, they could be due to a shorter rearing phase for the offspring, on the other hand, they could also be related to the formation of the shell and the numerous biological and functional restrictions associated with it. The extremely low metabolism of the armadillos would also be a potential possibility, as it means that less energy flows into the complex development of the brain. A low metabolism is also assumed for the glyptodonts. Their stature, which is much larger than that of the armadillos, allows further reflections on the small brain. The sometimes enormous body size and the massive tank hardly required any defense and escape strategies against large predators, which in turn speaks against a larger brain. Something similar has been reported about the armored ankylosaurs , which also have a small encephalization quotient in relation to unarmored dinosaurs. However, the armor itself can be considered as a restrictive functional component. Due to its compactness, this enabled only weakly developed neck muscles that support and stabilize the head. A reduced brain size thus supported the weight reduction of the skull, which then had a stronger effect, especially in the giant forms of the Pleistocene with their large skulls.

Parasites and pathologies

Glyptotherium osteoderms with surface changes caused by ectoparasites

Some dorsal and tail armor, including those of Glyptodon and Panochthus , sometimes have rounded perforations with a diameter of just a few millimeters and a conical cross-section. They resemble parasitic feeding marks caused by fleas , especially of the genus Tunga . Similar findings have also been reported for fossil armadillos and are also known from today's representatives of the group. Since such depressions can be proven in both the armadillos and the glyptodonts and have already been documented in fossils from the Miocene , a long coevolution between the armored animals and the fleas can be assumed. In addition, there are superficial changes to the ornaments of the bone platelets. These are possibly due to inflammation caused by bacteria or fungi .

So far, pathological changes can rarely be detected. These include bone deformations on the feet, especially in the area of ​​the joints. They are usually due to enthesopathies and pseudogout . Individual individuals may show several clinical pictures at the same time, which may indicate the susceptibility of the large glyptodons in particular to such joint diseases.

Systematics

External system

Relationship between glyptodons and armadillos according to morphological data from Billet et al. 2011
 Cingulata  

 Peltephilidae 


 Dasypoda  

 Dasypodidae 


  Chlamyphoridae  

 Tolypeutinae


   

 Chlamyphorinae


  Euphractinae  


 Eutatini


  Glyptodonta  

 Pampatheriidae


   

 Glyptodontidae




   

 Euphractini







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The Glyptodontidae form a family within the order of the armored collateral animals (Cingulata). This order was once represented in many forms in South and North America. It also includes the extinct Pampatheriidae , the Peltephilidae , the Palaeopeltidae , the Pachyarmatheriidae and the armadillos (Dasypoda) that still exist today . The armored articulated animals in turn represent a part of the superordinate hierarchy of the articulated animals (Xenarthra), which also include the tooth arms (Pilosa) with today's sloths (Folivora) and the anteaters (Vermilingua). The secondary articulated animals represent one of the four main lines of the higher mammals , which stands opposite the other three (collectively as Epitheria ) as a sister group . A general common feature of the secondary articular animals is found in the eponymous xenarthric joints (secondary joints, also xenarthral joints ) on the articular processes of the posterior thoracic and lumbar vertebrae. However, the Glyptodontidae are an exception here, as their lumbar vertebrae are fused together as a unique feature . The origin is still unknown, the oldest fossil finds come from South America and date to the Paleocene more than 56 million years ago, which are considered to be armadillos. Molecular genetic studies revealed that the secondary animals split off from the other higher mammals as early as the lower Cretaceous , about 103 million years ago. The armadillos as the closest relatives of the glyptodons living today separated from the common line with the tooth arms at the beginning of the Paleocene about 65 million years ago.

Holmesina from the group of the Pampatheriidae , the closest relatives of the Glyptodons

The internal structure of the armored articulated animals is not fully secured and is currently in flux. Of greater importance here are the Pampatheriidae, which are considered to be the closest relatives of the glyptodons in a classical view. These are characterized by armor on the back, which is similar to the armadillos with movable bands between a rigid shoulder and pelvic shield. In contrast to the armadillos with their variable number of movable ligaments, there were mostly three in the Pampatherien. With a weight of over 200 kg for some forms of the late Pleistocene , such as Pampatherium and Holmesina , they also reached significantly larger dimensions than the armadillos, but the design of the forelegs distinguishes them as not as skilful graves. Due to the distinctive construction of the tank, the Pampatheria were originally thought to be more closely related to the armadillos and were run within them as a subfamily. Studies on skulls and teeth, however, have shown that pampatheria and glyptodonts are closer. This resulted, among other things, from the structure of the ear canal and the construction of the chewing apparatus, such as the high lower jaw, but also due to the more complex teeth. The latter are simply built like nails in the armadillos, but are characterized by two transverse lobes in the pampatheria and three in the glyptodons. Both families together form the parent group of the Glyptodonta . The other groups - Palaeopeltidae, Peltephilidae and Pachyarmatheriidae - are more or less closely related to the other armored siblings, the latter possibly representing the sister group of the Glyptodonta.

Relationship between glyptodons and armadillos according to molecular genetic data from Delsuc et al. 2016
 Dasypoda  

 Dasypodidae 


  Chlamyphoridae  

 Euphractinae


   

 Glyptodontidae (Glyptodontinae?)


   

 Chlamyphorinae


   

 Tolypeutinae






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The relationships between the glyptodons and today's armadillos are subject to an ongoing scientific debate. The initially favored division of armored articulated animals into the large groups of armadillos and glyptodon relatives was largely eliminated in 2006 by more modern phylogenetic studies based on anatomical features. The analysis showed that the glyptodonts (glyptodonts and pampatheria) were much more closely bound to the armadillos and placed them in an evolutionary line of development. As a result, the Glyptodonta formed a common clade with the Euphractinae , which include today's six-banded armadillo . Refined studies in connection with further fossil material in 2011 moved the Glyptodonta even deeper into the armadillos, as they suggested a close relationship to the Eutatini , the latter, named after the character form Eutatus , represent an extinct branch of the euphractin armadillos the armadillos in themselves a paraphyletic group. The skeletal anatomical analyzes are contrasted with molecular genetic studies from 2016, which included the armadillos as well as the extinct glyptodon genus Doedicurus . They generally confirm the very close relationship between the two groups. In contrast to the relationship to the Euphractinae established by skeletal anatomy, the genetic data support a close relationship between the glyptodons and a clade consisting of the Chlamyphorinae, which make up the girdle mole, and the Tolypeutinae , within which the spherical armadillos , the bare- tailed armadillos and the giant armadillo are united. Accordingly, the Glyptodonts would be the sister group of these two subfamilies, the separation of the two lines occurred in the transition from the Eocene to the Oligocene about 35 million years ago. According to both the anatomical and the molecular genetic investigation results, the glyptodonts would only be seen as a side branch of the armadillos and would not represent an independent line of development within the armored articulated animals. There are different views on the exact taxonomic status of the glyptodons. Some researchers see the glyptodonts only at the level of a subfamily (Glyptodontinae) within the armadillos, others retain the family status for the time being (the position of the pampatheria is unclear because no DNA studies are yet available).

Internal system

Internal systematics of the glyptodons according to Croft et al. 2007
  Glyptodontidae  

 Parapropalaehoplophorus


   

 Glyptatelinae


   

 Propalaehoplophorinae


   

 " Hoplophorinae " 


   

 Doedicurinae


   

 Glyptodontinae 







Template: Klade / Maintenance / Style

In one oft-quoted view, the glyptodon family is divided into five subfamilies. The Glyptatelinae appear very primeval as a basal group. They were limited to the Eocene and the Oligocene and still had very original features of the osteoderms and teeth. The former are not as symmetrical, rosette-like as those of the younger Glyptodonts, but have a decentralized main pattern. Overall, however, they are considered to have been little explored. The Propalaehoplophorinae first appeared in the Upper Oligocene, but disappeared again in the Upper Miocene. They were significantly more developed than the Glyptatelinae, but less modern than the later Glyptodonts. Compared to these, they are characterized by slightly longer skulls with a more extended snout, as well as by the presence of up to six movable ligaments on the side edges of the back armor, which is considered to be the original state within the glyptodons. However, the osteoderms already have a symmetrical shape and mostly a central pattern. At around 70 to 115 kg, they lagged significantly behind their later forms in height.

The other three subfamilies include the " Hoplophorinae " (= Sclerocalyptinae), the Doedicurinae and the Glyptodontinae . The "Hoplophorinae" represent the most diverse group with about 60% of all known taxa. The subfamily is characterized by a few, insufficiently defined features, so some studies do not consider them a monophyletic group. It is therefore partly understood as a collective group for representatives who appear more modern than the Propalaehoplophorinae, but are outside the Glyptodontinae and Doedicurinae. The great heterogeneity of the subfamily is also reflected in the listing of at least six other subgroups (such as the Hoplophorini, Panochthini, Plohophorini), some of which, taken in and of themselves, are again heavily questioned. Occasionally as a result of this, individual specialists released various other subfamilies from the "Hoplophorinae". One of the most important developments within the "Hoplophorinae" is the formation of a club-like tail end. The Doedicurinae and Glyptodontinae comprise phylogenetically younger Glyptodonts, which were more modern than the Propalaehoplophorinae. The Doedicurinae are characterized by a pronounced tail lobe and bone platelets that are barely ornamented. The group is largely little researched except for the eponymous representative and is almost only known from the armor. The Glyptodontinae, in turn, largely retained the shape of the osteoderms of the Propalaehoplophorinae, but developed short tails without a club-like end.

Alternative internal systematics of the glyptodons according to Fernicola et al. 2008 and 2018
  Glyptodontia  

 Propalaehoplophoridae


  Glyptodontoidea  
  Panochthidae  

 Panochthini


   

 Neoscleroscalyptini



  Glyptodontidae  

 Lomaphorini


   

 Plohophorinae


  Glyptodontinae  

 Doedicurini


   

 Glyptodontini







Template: Klade / Maintenance / Style

A classification scheme of the glyptodonts deviating from this traditional structure resulted in cladistic analyzes of the year 2008. The entire group of glyptodonts in the classical view was raised to the level of subordination to the glyptodontia. The actual Glyptodontidae are limited to the more modern phylogenetic forms. Within the Glyptodontidae the Doedicurinae were united with the Glyptodontinae, a division is only made here on the level of the tribe . To the side of the Glyptodontinae are the Plohophorinae and individual other tribes. The Panochthidae, which are differentiated into the two tribes of the Panochthini and the Neosclerocalyptini, proved to be independent of the Glyptodontidae. The Glyptodontidae and the Panochthidae in turn form the superfamily of the Glyptodontoidea. The Propalaehoplophorinae, often assumed to be paraphyletic, turned out to be monophyletic, but now form a line outside of the Glyptodontoidea as an independent family. The Glyptatelinae were also given family status here. The problematic “Hoplophorinae”, on the other hand, have been resolved, but most of the genera contained cannot be assigned to a more precise group in this classification scheme, so they are considered incertae sedis . In this view, the Glyptodontia currently consist of four families: the Glyptatelidae, the Propalaehoplophoridae, the Panochthidae and the Glyptodontidae.

Overview of the genera

A total of about 65 genera of the Glyptodontidae are known, making the family very diverse. Most of the taxa , however, are not well known, their description is usually based on remains of the armor. Skulls or parts of the skeleton of the body are only known for very few forms. Traditionally, the distinction between the individual species and genera of the glyptodons is based on the shape of the bone platelets of the carapace and the design of the tail armor and not, as is usual in mammals , with the help of skull and dentition features. For this reason, it is sometimes difficult to assign isolated skeletal material to specific taxa that are otherwise only known from armor remains. The classification system used here largely follows McKenna and Bell 1997, with recent developments being taken into account.

  • Family: Glyptodontidae Gray , 1869
  • Boreostemma Carlini, Zurita, Scillato-Yané, Sánchez & Aguilera , 2008
  • Glyptotherium (= Brachyostracon , Boreostracon , Neothoracophorus ?, Xenoglyptodon ) Osborn , 1903

Tribal history

Adaptive Radiation and Origins

Numerous lines of the Glyptodonts experienced a strong increase in body size, especially in the last section of the Pleistocene . This is partly explained by Bergmann's rule , which means that the animals grew larger under the cooler conditions of the Ice Age. Other explanations cite the competitive pressure created by immigrating herbivores from North America in the course of the Great American Faun Exchange since the Pliocene . Further evolutionary changes concern the reduction in the number of toes in several lines and the overprinting of the shape of the bone platelets and the tail armor. It turns out, however, that similarly designed structures do not necessarily indicate a closer relationship, but are in part an expression of a convergent development .

The earliest known finds, some bone platelets from the carapace, come from Patagonia and belong to the Middle Eocene with an age of 48 to 42 million years . They are assigned to the genus Glyptatelus from the group of Glyptatelinae . The material was presented at the end of the 19th century and is said to have been recovered from correspondingly old rock units, but more precise data on the locations are not available. This means that the oldest evidence of the glyptodons is somewhat younger than that of the related armadillos, which was found in deposits of the Paleocene with an age of over 56 million years ago in southern Brazil . Possibly the region of origin of both groups can also be found in the southern part of the continent. In the Upper Eocene, Clypeotherium from the same family group has been proven.

Oligocene

Clypeotherium continues to occur during the Oligocene . In the late Oligocene, a time with greater diversification of the Glyptodonts, representatives of the Propalaehoplophorinae , one of the best-studied basal groups of these armadillos relatives, have come down for the first time . They are also considered the starting group for the emergence of the more modern glyptodons. Similar to the Glyptatelinae, the earliest records of the Propalaehoplophorinae from Patagonia are available, for example the site El Pajarito in the Argentine province of Chubut is important . But already in the Upper Oligocene they had reached regions further north, as some osteoderms from the Fray Bentos Formation in the Argentine province of Entre Ríos show. In addition to these two important lines, Pseudoglyptodon is a genus that has mixed features of the sloths and the glyptodons. The set of teeth with five teeth per upper jaw and four per lower jaw are reminiscent of the sloths, while the design of the occlusal surfaces with three transverse lobes is reminiscent of the glyptodon. In contrast to the glyptodonts, the teeth lacked the raised ridges made of hard dentin within the lobes . Significant finds come from Salla-Luribay in Bolivia and the Tinguiririca fauna from central Chile . Maybe, but is Pseudoglyptodon sloths closer.

Miocene

Remnants of armor from Propalaeohoplophorus

The glyptodons appeared in the Miocene with a wealth of forms. In the Lower Miocene and in the transition to the Middle Miocene from 18 to 16 million years ago, the Santa Cruz Formation is important in Patagonia, where the glyptodons with the Propalaehoplophorinae appear quite numerous. With Propalaehoplophorus , Cochlops , Asterostemma and Eucinepeltus, at least four genera are proven , the latter also being represented by at least four species. All representatives were significantly larger than today's armadillos and weighed between 67 kg for Propalaehoplophorus and 115 kg for Eucinepeltus . For the first time for glyptodons, complete skeletons have also been preserved from the Santa Cruz formation. As a result, the Propalaehoplophorinae are considered to be well studied. The members of the Propalaehoplophorinae lived mainly on a mixed vegetable diet and only moved on the ground. In addition to the finds of the Santa Cruz Formation, other important ones are known from other areas of South America. These include bone platelets and extensive remains of the body skeleton of Propalaehoplophorus from the somewhat more recent Río Mayo Formation in the Chubut Province of Argentina. A partial skeleton of Parapropalaehoplophorus has been reported from the Chucal Formation in northern Chile. The find is about the same as that of the Santa Cruz Formation, due to different characteristics of the teeth and the osteoderms, the latter hardly show any ornamentation, its exact systematic allocation is unclear.

In the Middle Miocene, a few early representatives still appear, such as the Paraeucinepeltus from southern Argentina , which is equipped with movable bands on the lateral armor edges . In addition, some more modern forms have also been proven, such as Eonaucum . The earliest representatives of the Glyptodontinae appear for the first time in the Middle Miocene around 12 million years ago and belong to the genus Boreostemma . A complete 1.4 m long specimen of Boreostemma from the Monkey Beds of the Villavieja Formation in La Venta in Colombia is available from this period. Since other early finds also come from northern South America, for example from the richly shaped Fitzcarrald local fauna of the western Amazon region in Peru, the group can be assumed to have originated in this region. From the north, the modern glyptodonts spread to the south, so that in the late Miocene they are also detectable in the pampa region with Glyptodontidium and possibly already Glyptodon . A very extensive glyptodon fauna of the Upper Miocene is from the Conglomerado osífero of the Ituzaingó Formation , which is exposed in north-eastern Argentina on the lower reaches of the Río Paraná near the city of Paraná and includes around a dozen genera.

Plio and Pleistocene

Skeletal reconstruction of Hoplophorus

Especially in the Pliocene and Pleistocene there was a strong fragmentation of the glyptodons, which was possibly accompanied by the spread of open landscapes as a result of the cooling climate. Were in the Pliocene with Boreostemma from the major Codore lineup in Venezuela to meet even early forms of modern Glyptodonten is in the Early Pleistocene before about 1.8 million years ago for the first time clearly Glyptodon handed down in southern South America. One of the earliest finds is a 28 cm long skull find from near Tarija in southern Bolivia. Finds of similar age come from the adjacent pampas region , so that the probable origin of the genus lay in this entire region. In the following time Glyptodon is the dominant representative of the Glyptodonts. The genus came mainly from southern Brazil to the southern areas of the continent, but was also found further north in the Andes . Their distribution area extended between the 20th and 38th parallel south. The originally assumed biodiversity of the genus is, however, possibly a relic of research, as modern analyzes can only distinguish two forms in the lowlands, which largely differ in time, plus a third from the Andes. Representatives of other lines also inhabited large areas of the continent. So Panochthus had a greater geographical range and ecological tolerance and settled both tropical areas and temperate areas from today's northeastern Brazil to far into southern Argentina. Also includes Panochthus next Glyptodon of the few members of the Armored Xenarthra, which at that time up in high mountain regions of partly penetrated 4000 m. Other forms, on the other hand, were more regionally limited. These include Hoplophorus , which predominantly occurred in tropical landscapes, and Doedicurus and Plaxhaplous in the pampas and in the neighboring Mesopotamia . Neosclerocalyptus also had a similar distribution area, with an estimated 250 kg body weight, which was the smallest of all the Pleistocene representatives of the glyptodons. A strongly ossified nasal region was unique to him.

Skeletal reconstruction of Glyptotherium

With the formation of the Isthmus of Panama around 3.5 million years ago during the Pliocene and the associated formation of a land bridge between South and North America, the Great American Fauna Exchange took place . The earliest glyptodon remnants north of South America, which contain only a few osteoderms, were discovered in the San Miguel Allende Basin of the Mexican state of Guanajuato in the central part of the country and date from around 3.6 to 3.9 million years ago. At the end of the Pliocene, the North American representative Glyptotherium was found for the first time , the bone platelets of which are more ornamented than those of its relative Glyptodon . At first rather small members of this genus appeared, weighing only around 230 kg, only in the course of the Pleistocene did forms weighing up to 790 kg develop. In North America Glyptotherium , the so far only recognized representative of the Glyptodonts there, was mainly distributed along the Gulf Coast , numerous finds are known from Texas , Arizona and Florida . One of the most important and richest sites is occupied with the 111 Ranch in Arizona. In Oklahoma , Glyptotherium reached its northernmost occurrence at 36 ° 37 ', as a back armor from Carmen in Alfalfa County indicates. In the course of the Middle Pleistocene , the glyptodons gradually disappeared from their northernmost refuges, but remained very widespread over today's Mexico. Obviously at this time there was a partial migration back to South America, since in the end of the Pleistocene Glyptotherium is also known from Venezuela and Brazil and inhabited coastal flatlands there.

die out

At the end of the Pleistocene in the transition to the Holocene , the glyptodons died out as part of the Quaternary extinction wave . For Glyptodon , the most recent directly dated finds are around 25,500 years old from Inciarte in Venezuela. More recent dates from 9600 to 10,500 years from Pay Paso in Uruguay were obtained from found charcoal , although a joint deposition of the remains of glyptodon and charcoal is not certain. Doedicurus survived significantly longer . It was found during the colonization of South America by early humans, which began about 14,500 years ago. But whether humans are causally responsible for the disappearance of the large mammal fauna is part of a much discussed debate. In the Pampa region is held doedicurus even to the early Holocene. Finds of the genus, which include a lower jaw, several cervical vertebrae, bones of the foot skeleton and other elements of the body skeleton as well as the shell, together with stone artifacts from early hunter-gatherer groups from La Moderna in northeastern Argentina , have been dated to an age of 7500 years BP . It is unclear here whether the humans hunted the animal themselves or cut up a carcass. A fragment of a humerus, which was found south of Buenos Aires, is a little older at around 8480 years of BP.

Research history

First discoveries in the 18th and 19th centuries

Dàmaso Antonio Larrañaga (1771-1848) created the first description of a glyptodon in June 1814

The earliest became known find of a Glyptodonten to the year 1774 back when the English Jesuit Thomas Falkner by a 2.7-meter body armor consisting reported of hexagonal bone plates that on the banks of the Rio Carcarañá near Santa Fe in Argentina had been discovered and which he compared to that of today's armadillos . In 1814 Dàmaso Antonio Larrañaga (1771-1848) created the first scientific description of a glyptodon in his Diario de Historia Natural based on a back armor, a thigh bone and a tail armor. He added the name Dasypus ( Megatherium Cuv) to this, which was introduced in 1823/24 by Georges Cuvier (1769-1832) in the second edition of his work Recherches sur les ossemens fossiles , one of the basic works for the development of paleontology . The indication of the subgenus Megatherium , actually a giant ground sloth, which Cuvier himself had scientifically introduced in 1796, led to the view of armored giant sloths being taken in the following time. In 1827, for example, Christian Samuel Weiss described the remains of a glyptodon from today's Uruguay and Brazil as belonging to Megatherium . In a publication from 1835, William Clift also linked a megatherium skeleton from the area around Villanueva on the Río Salado in the pampas region south of Buenos Aires with remains of glyptodont tanks that were also found there and also depicted them.

Six years later, Joseph Eduard d'Alton re- examined the material used by Weiss and also included skeletal remains that were found. After extensive anatomical comparisons, he came to the conclusion that they were giant armadillos. Heinrich Georg Bronn (1800–1862) used the same collection of finds for the establishment of the genus Chlamydotherium . However, the name Chlamydotherium is problematic, on the one hand because it was used almost simultaneously by Peter Wilhelm Lund for a representative of the Pampatheriidae , close relatives of the Glyptodons, and on the other hand because Bronn introduced it as the genus coelebs (unbound genus), without the genus one assign specific type. Chlamydotherium is now synonymous with Glyptodon . During the same period, an expedition led by Teodoro Vilardebó discovered several bones and armor in Uruguay, which they published in a local newspaper in 1838 after a scientific analysis. The term Dasypus antiquus used therein is, however, invalid. It was only in 1844, after the remains had been sent to the Muséum national d'histoire naturelle in Paris, that the Swiss zoologist François Jules Pictet was able to classify them as belonging to Glyptodon .

Scientific naming

Life reconstruction of Glyptodon after Richard Owens first description from 1839
Richard Owen (1804-1892) named the genus Glyptodon in 1839

The genus name Glyptodon was introduced by the English paleontologist Richard Owen (1804-1892) in 1839, another detailed presentation of the genus here, also combined with the species addition Glyptodon clavipes, followed two years later. The first description was based on a partial skeleton that had been found in the 1830s in the area of Cañuelas on the Río Matanza-Riachuelo south of Buenos Aires . Woodbine Parish , a senior British diplomat in Buenos Aires, sent the fossil remains to England, where they examined Owen in detail and discovered that it was a relative of the armadillos. He also revised all alleged remains of armor that were associated with Megatherium . The name Glyptodon is made up of the Greek words γλύφειν ( glyphine “cut in”; past participle γλύπτω , glypto ) and ὀδούς ( odoús “tooth”) and refers to the special design of the teeth that Owen saw in comparison to the armadillos.

Around the same time, Charles Darwin (1809–1882) undertook his research expedition with the HMS Beagle, which was important for evolutionary research, and landed several times on the coast of Argentina between 1832 and 1834. There he collected over 5000 fossil finds, mostly from the Pleistocene , which he forwarded to the Royal College of Surgeons in London (of which only 175 objects survived the bombing of London in 1941). It was there that Owen began studying fossils in 1836. Darwin himself was convinced of the armored giant sloths based on Cuvier's information on the body armor of Megatherium in the new edition of his work Recherches sur les ossemens fossiles and assigned numerous finds to them in his travel notes, which Owen corrected in the following. In parallel to Darwin's undertakings and Owen analyzes, the Danish researcher Peter Wilhelm Lund (1801–1880) worked in Brazil, where he brought together a collection of over 12,000 fossils from around 800 sites, mostly caves, in the Rio das Velhas region and shipped it to Copenhagen . These activities led Lund to describe Hoplophorus in 1838 .

Herrmann Burmeister's reconstruction of Glyptodon as a "two-shell" from 1866
John Edward Gray (1800–1875) introduced the family name of the Glyptodontidae in 1869

In 1869, John Edward Gray (1800–1875) established the name Glyptodontidae, which is used today for the family. In addition to Glyptodon, he also included Hoplophorus , Panochthus and Schistopleurum , the latter is now a synonym for Hoplophorus . The main characteristics he named:

Dorsal shield entire, not revolute, immovable affixed to the upper part of the very large pelvis.

"Back shield complete, not rollable, fixed immovably on the upper part of the very large basin."

A few years earlier, the German researcher Hermann Burmeister (1807-1892), who worked for years in South America, introduced the term biloricata ("two-shell") for the glyptodonts, assuming that the one on the stomach also had an, albeit thinner, shell trained. Gray worked this consideration into his description, adding that the head could be retracted into the shell, analogous to the turtles .

In 1875, remains of glyptodonts in the form of a complete armor in the valley of Mexico were discovered for the first time outside of South America , which were followed by a few more until the beginning of the 20th century, such as a complete armor and some skeletal parts in 1912. As early as 1888, Edward Drinker Cope (1840–1897) described individual osteoderms from Texas, thus providing the earliest evidence of glyptodons north of Mexico. It was not until 1903 that Henry Fairfield Osborn (1857–1935) published an almost complete skeleton of Glyptotherium , thus describing the only recognized genus in North America today.

The turn from the 19th to the 20th century

In the transition from the 19th to the 20th century, the works of the brothers Carlos and Florentino Ameghino stand out. In 1889, Florentino Ameghino first developed a systematic classification of the glyptodons, which included 13 of the 19 genera known at the time. He arranged them into three families (Glyptodontidae, Doedicuridae and Hoplophoridae). The basis of its systematic division was the different design of the osteoderms and the tail armor. Ameghino was the first to recognize the two basic tail types of the Glyptodons: on the one hand, the tail, which was completely surrounded and ended in a short tip, typical of Glyptodon, and, on the other hand, the tail with a club-like end, as it was found in Doedicurus . This method of subdividing the glyptodons was later used by other scientists. Other outstanding researchers on the glyptodons include Lucas Kraglievich in the 1930s and Robert Hoffstetter in the 1950s.

literature

  • Richard M. Fariña, Sergio F. Vizcaíno and Gerardo de Iuliis: Megafauna. Giant beasts of Pleistocene South America. Indiana University Press, 2013, ISBN 978-0-253-00230-3
  • Paul S. Martin and Richard G. Klein (Eds.): Quaternary Extinctions. A Prehistoric Revolution. The University of Arizona Press, Tucson AZ 1984, ISBN 0-8165-1100-4

Individual evidence

  1. ↑ a b c d e f g Sergio F. Vizcaíno, Juan C. Fernicola and M. Susana Bargo: Paleobiology of Santacrucian glyptodonts and armadillos (Xenarthra, Cingulata). In: Sergio F. Vizcaíno, Richard F. Kay and M. Susana Bargo (eds.): Early Miocene paleobiology in Patagonia: High-latitude paleocommunities of the Santa Cruz Formation. Cambridge University Press, New York, 2012, pp. 194-215
  2. ↑ a b c Leopold Héctor Soibelzon, Martín Zamorano and Gustavo Juan Scillato-Yané: Un Glyptodontidae de gran tamaño en el Holoceno Temprano de la Region Pampeana, Argentina. Revista Brasileira de Paleontologia 15 (1), 2012, pp. 105-112
  3. ↑ a b Kieren J. Mitchell, Agustin Scanferla, Esteban Soibelzon, Ricardo Bonini, Javier Ochoa and Alan Cooper: Ancient DNA from the extinct South American giant glyptodont Doedicurus sp. (Xenarthra: Glyptodontidae) reveals that glyptodonts evolved from Eocene armadillos. Molecular Ecology 25 (14), 2016, pp. 3499-3508, doi: 10.1111 / mec.13695
  4. ↑ a b Sergio F. Vizcaíno, R. Ernesto Blanco, J. Benjamí Bender and Nick Milne: Proportions and function of the limbs of glyptodonts. Lethaia 44, 2011, pp. 93-101
  5. ↑ Fariña, Vizcaíno and de Iuliis 2013. pp. 223–234
  6. ↑ a b c d e f Alfredo A. Carlini, Alfredo E. Zurita, Gustavo J. Scillato-Yané, Rodolfo Sánchez and Orangel A. Aguilera: New Glyptodont from the Codore Formation (Pliocene), Falcón State, Venezuela, its relationship with the Asterostemma problem, and the paleobiogeography of the Glyptodontinae. Paläontologische Zeitschrift 82 (2), 2008, pp. 139–152
  7. ↑ a b c d e f Richard A. F. Farina Sergio Vizcaíno: Carved teeth and strange jaws. How glyptodonts masticated. Acta Palaeontologica Polonica 46 (2), 2001, pp. 219-234
  8. ^ A b Sergio F. Vizcaíno, Richard A. Fariña, M. Susana Bargo, and Gerardo de Iuliis: Functional and phylogenetic assessment of the masticatory adaptations in Cingulata (Mammalia, Xenarthra). Ameghiniana 41 (4), 2004, pp. 651-664
  9. ^ Alfredo Eduardo Zurita and Silvia A. Aramayo: New remains of Eosclerocalypto tapinocephalus (Cabrera) (Mammalia, Xenarthra, Glyptodontidae): Description and implication for its taxonomic status. Revista Italiana di Paleontologia e Stratigrafia 113 (1), 2007, pp. 57-66
  10. ^ Alfredo Eduardo Zurita, Martín Zamorano, Gustavo J. Scillato-Yané, Laureano R. González-Ruiz, Santiago Rodríguez-Bualó, Boris Rivas Durán and Ricardo Céspedes Paz: An exceptional Pleistocene specimen of Panochthus Burmeister (Xenarthra, Glyptodontoidea) from Bolivia: Its contribution to the understanding of the Early-Middle Pleistocene Panochthini. Comptes Rendus Palevol 10, 2011, pp. 655-664
  11. ↑ a b c H. Gregory McDonald: Xenarthran skeletal anatomy: primitive or derived? Senckenbergiana biologica 83, 2003, pp. 5-17
  12. ↑ a b c d e f g h i David D. Gillette and Clayton E. Ray: Glyptodonts of North America. Smithonian Contributions to Paleobiology 40, 1981, pp. 1-251
  13. ↑ Sergio F. Vizcaíno: The teeth of the “toothless”: novelties and key innovations in the evolution of xenarthrans (Mammalia, Xenarthra). Paleobiology 35 (3), 2009; Pp. 343-366
  14. ^ Daniela C. Kalthoff: Microstructure of Dental Hard Tissues in Fossil and Recent Xenarthrans (Mammalia: Folivora and Cingulata). Journal of Morphology 272, 2011, pp. 641-661
  15. ↑ Laureano R. González-Ruiz, Martin R. Ciancio, Gabriel M. Martin and Alfredo E. Zurita: First Record of Supernumerary Teeth in Glyptodontidae (Mammalia, Xenarthra, Cingulata). Journal of Vertebrate Paleontology 35 (1), 2015, p. E885033
  16. ↑ Kevin F. Downing and Richard S. White: The cingulates (Xenarthra) of the Leisey Shell Pit local fauna (Irvingtonian), Hillborough County, Florida. Bulletin of the Florida Museum of Natural History 37 (Part II), 1995, pp. 375-396
  17. ↑ a b c Kleberson de O. Porpino, Juan C. Fernicola and Lílian P. Bergqvist: Revisiting the Intertropical Brazilian Species Hoplophorus euphractus (Cingulata, Glyptodontoidea) and the Phylogenetic Affinities of Hoplophorus. Journal of Vertebrate Paleontology 30 (3), 2010, pp. 911-927
  18. ↑ a b c Alfredo Eduardo Zurita, Martín Zamorano, Gustavo Juan Scillato-Yané, Sergio Fidel, Martín Iriondo and David D. Gillette: A new species of Panochthus Burmeister (Xenarthra, Cingulata, Glyptodontidae) from the Pleistocene of the Eastern Cordillera, Bolivia . Historical Biology 29 (8), 2017, pp. 1076-1088, doi: 10.1080 / 08912963.2016.1278443
  19. ↑ a b c d Francisco Cuadrelli, Alfredo E. Zurita, Pablo Toriño, Angel R. Mio-Boilini, Daniel Perea, Carlos A. Luna, David D. Gillette and Omar Medina: A new species of glyptodontine (Mammalia, Xenarthra, Glyptodontidae ) from the Quaternary of the Eastern Cordillera, Bolivia: phylogeny and palaeobiogeography. Journal of Systematic Palaeontology, 2020, doi: 10.1080 / 14772019.2020.1784300
  20. ^ Robert V. Hill: Comparative Anatomy and Histology of Xenarthran Osteoderms. Journal of Morphology 267, 2005, pp. 1441-1460
  21. ↑ a b c Alfredo Eduardo Zurita, Leopoldo Hector Soibelzon, Esteban Soibelzon, Germán Mariano Gasparini, Marcos Martín Cenizo and Héctor Arzani: Accessory protection structures in Glyptodon Owen (Xenarthra, Cingulata, Glyptodontidae). Annales de Paléontologie 96, 2010, pp. 1–11
  22. ^ A b R. Ernesto Blanco, Washington W. Jones and Andrés Rinderknecht: The Sweet Spot of a Biological Hammer: The Center of Percussion of Glyptodont (Mammalia: Xenarthra) Tail Clubs. Proceedings of the Royal Society B 276 (1675), 2009, pp. 3971-3978
  23. ↑ Victoria M. Arbor and Lindsay E. Zanno: The evolution of tail weaponization in amniotes. Proceedings of the Royal Society B 285, 2018, S. 20172299, doi: 10.1098 / rspb.2017.2299
  24. ↑ Victoria M. Arbor and Lindsay E. Zanno: Tail Weaponry in Ankylosaurs and Glyptodonts: An Example of a Rare but Strongly Convergent Phenotype. The Anatomical Record 303 (4), 2020, 988-998, doi: 10.1002 / ar.24093
  25. ^ A b Alfredo E. Zurita, Ángel R. Miño-Boilini, Analía Francia and José E. Arenas-Mosquera: The Pleistocene Glyptodontidae Gray 1869 (Xenarthra: Cingulata) of Colombia and some considerations about the South American Glyptodontidae. Revista Brasileira de Paleontologia 15 (3), 2012, pp. 273-280
  26. ^ François Pujos and Rodolfo Salas: A systematic reassessment and paleogeographic review of fossil Xenarthra from Peru. Bulletin de l'Institut Français d'Etudes Andines 33 (2), 2004, pp. 331-377
  27. ↑ a b Jim I. Mead, Sandra L. Swift, Richard S. White, H. Greg McDonald and Arturo Baez: Late Pleistocene (Rancholabrean) Glyptodont and Pampathere (Xenarthra, Cingulata) from Sonora, Mexico. Revista Mexicana de Ciencias Geológica 24 (3), 2007, pp. 439-449
  28. ↑ Nick Milne, Sergio F. Vizcaíno and Juan Carlos Fernicola: A 3D geometric morphometric analysis of digging ability in the extant and fossil cingulated humerus. Journal of Zoology 278, 2009, pp. 48-56
  29. ↑ Fiona R. Straehl, Torsten M. Scheyer, Analía M. Forasiepi, Ross D. MacPhee and Marcelo R. Sánchez Villagra: Evolutionary Patterns of Bone Histology and Bone Compactness in Xenarthran Mammal Long Bones. PLoS ONE 8 (7), 2013, p. E69275, doi: 10.1371 / journal.pone.0069275
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  120. ↑ Barnum Brown: Brachyostracon, a New Genus of Glyptodonts from Mexico. American Museum of Natural History Bulletin 31 (17), 1912, pp. 167-177
  121. ^ Henry Fairfield Osborn: Glyptotherium texanum, a New Glyptodont, from the Lower Pleistocene of Texas. Bulletin of the American Museum of Natural History 19 (17), 1903, pp. 491-494
  122. ↑ Florentino Ameghino: Contribución al conocimiento de los mamíferos fósiles de la República Argentina. Actas de la Academia Nacional de Ciencias de la República Argentina 6, 1889, pp. 1–1027 (pp. 758–852) ( [14] )
  123. ↑ Fariña, Vizcaíno and de Iuliis 2013. pp. 183–186

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This article was added to the list of excellent articles on April 24, 2020 in this version .