Armadillos

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Armadillos
Nine-banded armadillo (Dasypus novemcinctus)

Nine-banded armadillo ( Dasypus novemcinctus )

Systematics
without rank: Synapsids (Synapsida)
Class : Mammals (mammalia)
Subclass : Higher mammals (Eutheria)
Superordinate : Sub-articulated animals (Xenarthra)
Order : Armored siderails (Cingulata)
without rank: Armadillos
Scientific name
Dasypoda
Quenstedt , 1885 ( Hoernes , 1886)

The armadillos (Dasypoda) form the only surviving group of mammals of the armored collateral animals (Cingulata). Today they are divided into two families with a total of around 20 currently occurring species , and numerous other extinct representatives are known. The group is distributed over the South American continent and the southeastern parts of North America ; a large part of today's species is concentrated in central and northern South America.

As the only known group of mammals today, the armadillos have a bony outer shell. The pangolins armored with horn scales used to be mistakenly considered to be closely related to them. Armadillos appear largely solitary and to varying degrees live underground in earthworks, whereby they are adapted to this burrowing way of life through several skeletal modifications. A wide variety of insects are consumed as main food, but some species also consume plant-based food or devour smaller vertebrates . Armadillo habits have not been well studied in all cases, especially when it comes to reproduction.

Some of the meat is used by humans as a food resource, some skeletal parts are also used locally as handicraft raw materials. Some species are considered land nuisances because of their ground-burrowing activities. The armadillos first became known in Europe at the beginning of the 16th century, the first description of the Dasypoda group, recognized today, dates from 1885/1886.

The phylogenetic development, documented by fossil finds , can be traced back to the Paleocene , more than 56 million years ago. However, the currently existing species appeared relatively late. The closest relatives of the armadillos are the anteaters and sloths . All three groups have special skeletal formations on the spine, which together allow them to form the superordinate order of the secondary articulated animals (Xenarthra), one of the four great main lines of the higher mammals .

anatomy

Habitus

Belt gullet ( Chlamyphorus truncatus )

Armadillos are predominantly small to rarely medium-sized mammals with a head-trunk length of 11 to 100 cm. In addition, there is a tail which, with the exception of the ball armadillos ( Tolypeutes ) and the belt mole, often makes up about half the body length, but is sometimes significantly longer. Similar to body size, the weight varies greatly and ranges from 100 g for the gullet ( Chlamyphorus truncatus ) to 45 kg for the giant armadillo ( Priodontes giganteus ). Extinct armadillos could also grow considerably larger and, with Macroeuphractus , which weighed between 90 and 120 kg, reached the weight of today's smaller domestic pigs . A large part of the known species remained in the size variability of the recent representatives.

The head has a narrow, pointed shape, the ears protrude upwards like a mouse, the eyes are very small. In some species the snout is elongated like a tube. On the upper side, the head is covered with bone plates that form a head armor. Characteristic is the usually strongly arched back armor , which covers almost the entire body surface of the animals. The front and rear body it is often too rigid back shields fused (shoulder and lap plate, also Buckler called), is located between them a varying number of open to the stomach side, transverse rings, belts or bands, after which the group its German trivial name received . Connections with overlapping skin folds ensure a high flexibility of the armor in the area of ​​the ligaments. In most species, the pointed tail is also surrounded by rings of bones. On the other hand, there is no armor on the stomach, but sometimes there are individual bone platelets. Body hair is not developed in all species, if it is present, it sprouts in bristle hair from the back shields or on the sides of the body and is woolly close to the belly.

The short legs always have five toes at the back, but four to five, rarely three, toes at the front, which have strongly curved claws , with those of the hind feet in all species being significantly smaller than those of the front ones. The claws on the middle toes of the forefeet of the giant armadillo hold up to 20 cm in length, the record for the largest claws in the entire animal kingdom today.

Skull and dentition features

Six-banded armadillo skull ( Euphractus sexcinctus )

The different types of armadillos have varying skull shapes, which mostly reflect the diet. Species that feed mainly on insects usually have a lighter skull and lower jaw with less developed muscle attachment points. However, those who live largely as omnivores and in part also eat plants are characterized by more massive skulls with clearly developed chewing muscles. In addition to the general structure of the mammalian skull, the armadillos have two special anatomical features. In the anterior area of ​​the nasal cavity, a bone formation known as the septomaxilla (os nariale) occurs, which is otherwise only formed in mammals and reptiles that are historically older . The function of this bone in the armadillos is largely unknown; it is probably used to close the nostrils while digging so that dust is not inhaled. Many, but not all, armadillo species have a tympanic ring on the ear canal instead of a tympanic bladder .

Brown-bristle armadillo ( Chaetophractus villosus ) skull

The lower jaw of many species is mostly long, narrow and built low on the bone body. The not firmly ossified symphysis that connects the two halves of the lower jaw in the anterior region is striking . In tooth construction, there are also differences in diet with small teeth in insectivores and large teeth in omnivores. In addition, the teeth differ from those of other mammals: They are designed homodontically , incisors or canines or premolars are not formed. The entire set of teeth consists of simple, pencil- like tooth formations reminiscent of molars . These do not have any tooth enamel - only the long-nosed armadillos ( Dasypus ) and some extinct species have a very thin layer of enamel that is chewed off very quickly - and consist almost entirely of dentine , mainly orthodentin , the hardest component of the dentin. Some species have a very thin outer layer of dental cement instead of tooth enamel , which is the case with the giant armadillo or the dwarf armadillo ( Zaedyus pichiy ). The number of teeth varies from species to species and often also within the species and ranges from 7 to 25 per jaw arch. The teeth usually grow for a lifetime, only in the long-nosed armadillos a change of teeth from deciduous to permanent dentition has been proven. This can sometimes happen very late, sometimes not until adulthood, and obviously varies from person to person.

Armor

Schematic representation of the armor of an armadillo with the names of the individual elements
Armor of an armadillo, here the extinct form Eutatus

The formation of an outer, bony body armor that covers the head (head shield) and the back (carapace) and with the exception of the bare- tailed armadillos ( cabassous ) also the tail (tail armor) and parts of the legs is unique among mammals and gives the armadillos and give their extinct relatives a special position within this class of animals. It consists of bone platelets called osteoderms , which are formed in the skin . They are arranged in rows on both the back armor and the tail. The number of shell-forming osteoderms varies depending on the species. In the spherical armadillos it is over 660, in the dwarf armadillo around 950 and in the bristle armadillos ( Chaetophractus ) over 1040. The osteoderms are covered by horn shields that are arranged in two different ways: on the one hand, a horn shield covers several, sometimes up to four neighboring osteoderms at the same time, on the other hand only one or a maximum of two. The former variant is considered to be rather primitive for the armadillos and is almost exclusively developed only in the long-nosed armadillos, the latter applies to most other species.

The osteoderms of the rigid parts of the back armor often have a square, four- to polygonal shape. On the other hand, they are mostly elongated and rectangular in shape on the movable bands and have an articulated surface at the front end that connects them to the corresponding element of the adjacent row. A single osteoderm of a movable ligament thus lies beneath its counterpart of the previous row with the joint surface and covers the respective joint surface of the subsequent bone platelet in the rear section. The surface of the osteoderm is usually ornamented. This surface drawing usually includes a larger, central pattern, around which smaller patterns are arranged concentrically or laterally. The shape and surface structure of the bone platelets vary between the species, so both features have taxonomic value. The horn shields are mostly round in shape and overlap the bone platelets. In the long-nosed armadillos, whose horny shields cover several osteoderms, an irregular pattern is sometimes created on the shoulder and pelvic shields, while in the other armadillos the linear structure of the armor is repeated.

The bone platelets are multilayered in cross-section, with a firm layer of bone above and below and softer material inside, in which there are cavities for sweat and sebum glands as well as bone marrow and, in hairy species, also for hair follicles. In general, the individual cavities have their own character, depending on their function, and can therefore be assigned relatively safely even with fossil forms. Such hair follicles are mostly elongated and are grouped at the edge of the osteoderm, glands often have a spherical shape and sit in the middle, while those for bone marrow usually have no opening to the surface. The bony substance is made up of hard, strongly mineralized material and collagen-rich , fibrous structures ( Sharpey fibers ), which give the individual bone platelets a high degree of elasticity. Inside, the armor is supported by mostly 11 pairs of very wide ribs.

The creation of the shell begins in the fetal phase in the womb. Newborns, however, have a soft, leathery shell that gradually hardens through ossification of the dermis . The typical, depending on the species, differently shaped, polygonal to round bone platelets are formed, on which the horn platelets sit and sort themselves into solid armor parts and belts. The hardening of the tank takes place differently depending on the species during youth growth, which is interpreted as an adaptation to the respective environmental conditions.

Whether the tank was formed as protection against predators has not been fully scientifically established. However, it protects well against the thorny vegetation of dry landscapes, where numerous species of armadillos live. At the same time, it reduces the infestation with external parasites .

Body skeleton

Giant armadillo skeleton ( Priodontes maximus )

Special skeletal features that differ from other mammals are found primarily in the area of ​​the spine . The so-called xenarthric joints (secondary joints), which gave the parent Xenarthra its name, are mainly formed on the posterior thoracic and lumbar vertebrae. These additional joints are found on the lateral processes of the vertebrae and connect the preceding with the following (more precisely the accessory process with the transversus and mammilar processes ), but their function is largely unclear. The articular surfaces of the caudal vertebrae are also strongly developed and intermesh deeply, which enables the armadillos to use the tail as a support when straightening up on the hind legs. In the area of ​​the cervical spine, there are adhesions of the posterior cervical with the anterior thoracic vertebrae in some species, which is known as postcervical bone and is most likely connected with the burrowing way of life of the animals. In addition, armadillos only have an average of 11 thoracic vertebrae and usually between 3 and 4 lumbar vertebrae, which is significantly lower than that of their closest relatives, the anteaters and sloths and, especially in the case of the lower number of lumbar vertebrae, gives the back more stability when straightening up. Such a reduction in the number of all vertebrae in comparison to closely related groups also seems to be typical for armor-bearing vertebrates , since the same has been proven in turtles and some extinct parareptiles with armor formation.

The rest of the musculoskeletal system also shows some peculiarities. The strong front legs with the big claws are an excellent adaptation to a digging way of life. In the bone structure, this is particularly evident in the ulna , the proximal (= near the trunk) articular process (the olecranon ) is very pronounced; its length makes up between 37 (giant armadillo) and 54% (girdle mole) of the total bone length. The percentage of the olecranon on the ulna is greater, the more the individual species are adapted to a burrowing way of life. The proximal as well as the distal (= distal end of the humerus) also show reinforcements in burrowing species, which serve the attachment of the corresponding musculature.

The hind limbs, on the other hand, are not excessively strong, which is due to the fact that they are not used for digging, but primarily for locomotion when foraging. The greater roll hillock (greater trochanter) of the thigh bone is located approximately at the level of the head of the femur, which promotes the extension of the leg in the hip joint. This happens, among other things, in the burrows when clawing into the ground when an animal is attacked by a predator. In addition to the large and small hillocks (trochanter lesser), the femur typically has a third hillock (trochanter tertius); the latter is not regularly found in mammals, but is a characteristic feature of secondary articulated animals. In armadillo species with a higher body mass, the tertius trochanter is more distal (towards the knee joint) than in those with less.

Internal organs and sensory performance

Ingestion of food takes place with the help of the long, extensively extendable tongue, which also has a surface roughened by mushroom, circular or thread-shaped papillae in most of the armadillo species examined. The digestion is supported by saliva containing sialic acid , which is formed in three different main groups of salivary glands (ear, sub-lingual and lower salivary glands ). Much of the food decomposition takes place in the stomach , the pylorus of which is heavily muscled . The stomach is mostly sack-like and simple, but in some armadillo species, such as the southern seven-banded armadillo , it appears almost bilobed when it is drawn in. It is also quite large and can measure around 150 cm³ in the nine-banded armadillo . The spleen reaches between 0.21 and 0.38% of the body weight and has a blood cell-forming function over the entire lifespan of an animal.

The uterus of the long-nosed and spherical armadillos is simple and trapezoidal, which makes it resemble that of the primates . All other armadillos have a two-horned uterus. Females have a single pair of mammary glands , with the exception of the long-nosed armadillos, which have two pairs. The penis is compared to the height of one of the longest among all mammals and achieved in some species in the erect state up to 50% of the body length. Because of the body armor, copulation would not be possible without such a large penis.

Above all, the sense of smell is very pronounced and is often used when searching for food. This is also reflected in the brain , which has a large olfactory brain . The sense of sight , on the other hand, is underdeveloped, which can also be diagnosed by a less pronounced area optica on the brain.

distribution and habitat

Armadillos live exclusively on the American double continent. The occurrence of most species is restricted to South America , where they are distributed over most of the land area. The origin of the armadillos can also be found here, going back to the end of the Paleocene around 58 million years ago. There are only two species in Central America , including the best-known armadillo, the nine-banded armadillo ( Dasypus novemcinctus ), which inhabits large parts of South and Central America up to and including the southeastern USA . The ancestors of the North and Central American species probably only immigrated from South America about 3 million years ago after the land bridge between North and South America was formed on the Isthmus of Panama and the associated Great American Fauna Exchange in the Pliocene .

Numerous armadillo species prefer dry and open habitats such as semi-deserts , savannas and steppes with thorny bush vegetation or dry forests and also need a loose surface to build their burrows. As a result, many representatives can be found in the Gran Chaco , but also in the pampas regions of central South America from southern Brazil through Bolivia and Paraguay to northern Argentina ; in Paraguay alone, 12 of the 21 species still alive today have been identified. However, some species also live in the tropical rainforest , in the Yungas forests of the high mountain flanks or in wetlands such as the Pantanal . The high elevations of the Andes have only opened up individual armadillo species, such as the small bristle armadillo ("Andean bristle armadillo "; Chaetophractus vellerosus ).

Way of life

Territorial behavior

Pygmy armadillo ( Zaedyus pichiy )

The way of life of most armadillo species is little explored. Reliable data usually refer to the more widespread and more common species, such as the nine -banded or six-banded armadillo ( Euphractus sexcinctus ). Armadillos are predominantly nocturnal loners; however, some species also appear during the day. A total of three life models can be distinguished:

  • predominant soil dwellers who rarely dig underground burrows, such as the spherical armadillos ( Tolypeutes );
  • predominantly burrowing animals that go in search of food superficially, such as the long-nosed armadillos ( Dasypus ) and bristle armadillos ( Chaetophractus );
  • generally burrowing animals that also feed underground, such as the bare- tailed armadillos ( Cabassous ) and both representatives of the belt mole ( Chlamyphorus and Calyptophractus ).

Armadillos spend a large part of the waking phase searching for food, mostly nimble and permanently, whereby the well-developed sense of smell detects the prey up to 20 cm deep in the ground, which is then usually dug up. Some representatives of the armadillos are able to hold their breath for up to six minutes in order to keep their airways free. In spite of the clumsy and apparently rigid physique, the armadillos can move astonishingly nimble. Some species are even good swimmers; So that they do not get lost in their armor, they first pump air into the stomach and intestines . To sleep, they dig themselves into the ground or look for an existing building, but the re-use of such buildings differs significantly from species to species. The caves themselves are also different depending on the species and can be small, but also branched and several meters long underground. In some cases, enlarged nesting areas are created at the end of the corridors.

nutrition

Erected nine-banded armadillo ( Dasypus novemcinctus ) takes in the scent

The armadillos have a wide range of food, but insects are more or less eaten by all representatives depending on the amount. However, different, species-dependent nutritional strategies can be distinguished here:

  • predominant carnivores and omnivores , such as the six-banded armadillo ( Euphractus ), the dwarf armadillo ( Zaedyus ) and the bristle armadillos ( Chaetophractus ); In addition to insects, these also devour small vertebrates such as lizards and mice , and more rarely carrion and vegetable food ;
  • opportunistic insectivores, such as the ball armadillos ( Tolypeutes ), the long-nosed armadillos ( Dasypus ) and the belted gullet ( Chlamyphorus ); these feed on a wide variety of insects such as ants , termites , but also beetles and their larvae, as well as grasshoppers and spiders ;
  • specialized insectivores such as the giant armadillo ( Priodontes ) and the bare- tailed armadillos ( Cabassous ); their diet largely includes ants and termites.

Similarly, the extinct armadillos basis of examined teeth fed largely omnivorous or insektivor, but there is also evidence of a significantly fleischhaltigere food, such as when large Macroeuphractus whose second tooth significantly to a canine remembered and possibly up to hare large animals captured. Representatives from the Eutatini group , such as Eutatus , in turn were more specialized in plant-based food, which could be determined from the structure of the lower jaw and the traces of abrasion on the teeth.

Food is usually consumed with the long, sticky tongue . The long claws of the forelegs are used to tear up the ant and termite mounds, which can be very hard at times, or to dig holes in which to look for food. When foraging for food, armadillos often stand up on their hind legs to follow a scent trail.

Energy balance and consequences of the tank

In general, the armadillos have a very low metabolic rate ; it is around 40 to 60% of that which would be expected in mammals of the same size. Along with that of the monotremes , it is one of the lowest of the mammals. The more the animals are adapted to ants and termites as primary food resources, the lower the metabolism. The slow metabolism is caused by the low energy density of the food; the additional soil material taken in when rummaging around in the insect nests also reduces this. The effect is sometimes more pronounced in larger species than in smaller ones, as in the giant armadillo, among other things, the metabolic rate sometimes only reaches 29%. In contrast to most mammals, the higher oxygen demand that may arise depending on the activity is not covered by deeper but faster breaths. The necessity of this behavior results from the formation of the armor, which constricts the chest and thus limits its freedom of movement.

The slow metabolic rate is also associated with a low and fluctuating body temperature and high thermal conductivity. The average body temperature of tropical forms is around 33 to 34.5 ° C, which is slightly lower than that of animals from cooler regions. The high thermal conductivity is favored by the formation of the shell and the lack of a dense fur, it promotes the release of body heat to the environment and thus prevents excessive heating under warm climatic conditions. In order to prevent body heat being released too quickly, the animals withdraw into their underground burrows with significantly more balanced environmental conditions when there are strong local temperature fluctuations. Species that occur in colder regions, such as the dwarf armadillo, react to inhospitable phases with a torpor or, as with the nine-banded armadillo, encounter them with reduced activity. Sometimes these more cold-resistant representatives also accumulate a pad of fat. There is hardly any evidence of such a food storage strategy for many tropical species. An additional pad of fat, for example, would no longer enable the spherical armadillos to completely curl up in their shell.

The general constitution and the energy balance mean that armadillos are more common in warm climatic landscapes and prevent a stronger spread in cool climatic latitudes. Another limiting factor is the availability of ants and termites to meet their nutritional needs. The colony-building insects do not thrive in cold winter conditions. All of this limits, for example, the further northern expansion of the nine-banded armadillo considerably. Furthermore, the diet and the resulting metabolism have a direct influence on the life strategies of different animal groups. In the armadillos, the low-energy diet and low metabolism lead to an underground way of life. The trained tank in turn offers a certain protection against both predators and collapsing tunnels and the like. As a result, they also retained a more or less "primeval" locomotion with a sole gait and short limbs, which in principle allow fast, but less effective locomotion through short steps. The same is also known by other insectivorous mammals with external protection training, be it the pangolins or the diverse, equipped with spiky fur dress animals like the hedgehog or different representatives of tenrecs . On the other hand, the rabbits, which were comparable in size to the armadillos, developed a highly mobile musculoskeletal system due to their high-energy diet and high metabolic rate, which is characterized by long limbs. With their high speeds and long strides, they are effective escape animals, so that body armor was not necessary here.

Reproduction

Reproduction has only been well researched for the fewest and, above all, the most widespread species. In general, the rutting season is the only phase in which several individuals of the armadillos come together. The gestation period is quite different and can vary between two and four months in the more well-known species. In the case of the nine banded armadillo, it has been proven that the fertilized egg cells are stored in the body for several months before they are nidated and further developed. This prevents the females from giving birth to offspring with no prospects in unfavorable times. In addition, polyembryonia is known in the nine-banded armadillo and also in the southern seven-banded armadillo , so that they produce genetically identical offspring, the number of which is two to twelve per litter. This form of reproduction is also assumed for all other representatives of the long-nosed armadillos . However, it is still unclear whether the polyembryony is related to the development of the simply built uterus in these armadillo representatives. In four other genera the birth of non-identical twins is attested, the remaining species only give birth to one offspring per litter. The boys initially have soft, leathery skin that gradually hardens to form a shell, and they are usually only suckled for a few weeks. The life expectancy of the armadillos in the wild is up to 18 years, in human care some species can even reach more than 30 years, although not all species have been successful in breeding.

Predators and enemy behavior

Standing and curled up southern spherical armadillo ( Tolypeutes matacus )

In case of danger, armadillos usually go to the nearest burrow or dig themselves in, where they then ram their claws into the ground and spread the bone plates and press them into the tunnel, anchoring themselves firmly in the ground. In the open air, the claws are also pressed into the ground, the animals also press themselves firmly to the ground so that only the armor can be attacked and the soft belly is protected. However, the armor then does not offer complete protection, as it is only 2 to 3 mm thick with some representatives and can easily be cracked by some larger predators . Only the spherical armadillos can curl up into a completely protected sphere; the carapace is so seamlessly interlocked and the muscle connection so tight that hardly any predator can bite this shell. The claws can also be used well for defense. Potential predators include larger predatory cats such as Jaguar and Puma , but also smaller predators such as coyote and ocelot or birds, the Aguja and the fence Adler . Wild dogs and cats also hunt armadillos. Young animals can also be killed by numerous other predators.

Ecological importance

Armadillos exert a great influence on their respective local habitats . The digging representatives make a contribution to bioturbation and thus to aeration and loosening of soil substrates. The amount of earth moved, depending on the size of the cause, is between 0.002 and 0.6 m³ per building. For an examined region in Colombia, this resulted in a volume of approximately 99 m³ with 157 registered burrows of the giant armadillo. The burrows not only serve as a shelter for the armadillos, but are also used by numerous other animals. Since the armadillos thus create habitat for various living things, scientists classify them as ecosystem engineers . This has been studied particularly well in the case of the giant armadillo, whose burrows sometimes serve as a habitat for more than two dozen different species. Armadillos also have a share in the nutrient cycle through their burrowing and migrating way of life . This does not only apply to the nitrogen content , for example , which can increase significantly locally due to defecation with a corresponding population density . In addition, armadillos take in significant amounts of soil due to their diet, which is then excreted elsewhere. Some species, sometimes herbivorous, also act as seed dispersers . This includes more than three dozen species of plants, the fruits of which have been shown to belong to the diet of the armadillos. Other issues include controlling the spread of disease by killing a wide variety of insect species . However, armadillos are also carriers of several pathogens that can be dangerous to humans. These include leprosy and Chagas disease . Last but not least, armadillos form part of the food web, as they not only live predators to a certain extent, but are also preyed on by other predators.

Systematics

External system

Relationship of 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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Holmesina , a great representative of the Pampatherien from the Late Pistocene of America

The armadillos (Dasypoda) are a group from the order of the armored articulated animals (Cingulata) and the superordinate order of the articulated animals (Xenarthra). The xenarthric joints on the spine are a common feature of the otherwise very diverse articular animals. Also belonging to the Xenarthra are the sloths (Folivora) and the anteaters (Vermilingua), both of which represent a more closely related group and, as an order of the tooth arms (Pilosa), stand opposite the armadillos. Molecular genetic analyzes showed that the separation of the common lineage of the sloths and anteaters from that of the armadillos had already occurred at the beginning of the Paleocene around 65 million years ago. The Xenarthra, whose origins go back to the Upper Cretaceous , form one of the four major superordinate groups of the Upper Mammals , the other three are partially combined in a superordinate taxon , the Epitheria , which is opposite to the Xenarthra as a sister group. All three recent orders are limited to America today .

The composition and systematic structure of armored articulated animals is complex and under discussion. In general, besides the armadillos, some other larger, only fossilized lines can be distinguished. Of importance here are the Pampatheriidae and Glyptodontidae , which, according to phylogenetic studies, are considered to be very closely related and are therefore grouped together in the higher taxonomic unit of the Glyptodonta . The Pampatherien appeared for the first time in the Middle Miocene and also have a rigid shoulder and pelvic armor, between which there are individual, mostly three, movable ligaments. However, especially in the later phase of their tribal history , the representatives were much larger than today's armadillos. The North American genus Holmesina , which only became extinct at the end of the Pleistocene around 10,000 years ago, reached a length of two meters and a body weight of up to 220 kg; Pampatherium , which was widespread in South America at the same time, had similar dimensions . Originally, the Pampatherien was considered to be closely related to the armadillos, and in some cases they were also run as a subfamily within them. The closer relationship between the pampatheria and glyptodonts results, among other things, from the structure of the auditory canal and the construction of the chewing apparatus, such as the high lower jaw and the more complex teeth. The glyptodonts in turn form one of the largest groups within the Cingulata. They are characterized by a rigid armor and by the absence of xenarthric joints; due to their high-crowned teeth, they followed a rather grass-eating way of life, similar to the pampatheria. Glyptodonts appeared for the first time in the Middle Eocene and were initially quite small, with the Propalaehoplophorus from the Lower Miocene weighing only around 74 kg. In the Pleistocene, however, they reached enormous proportions with a weight of over 2 tons as in Doedicurus .

During the 20th century, most scientists viewed the glyptodon and armadillo kin as two separate lines of development. Skeletal anatomical studies from 2006, carried out by Timothy J. Gaudin and John R. Wible , however, referred the Glyptodonta to be related to the armadillos. The immediate environment of the six-banded armadillo ( Euphractus ) and the bristle armadillos ( Chaetophractus ) proved to be particularly important here. However, this gave first indications of a paraphyletic origin of the armadillos. This was substantiated by further anatomical studies in 2011 that pushed the Glyptodonta even deeper into armadillo control. More recent genetic analyzes from 2016 on the fossil genome of the Glyptodontidae, namely Doedicurus , came to a comparable result. According to this, at least the glyptodonts are grouped within the Chlamyphoridae, but unlike the anatomical studies, a closer relationship to the relatives around the spherical armadillos ( Tolypeutes ) and the belt mole is more likely. On the one hand, this confirmed the paraphyletic origin of the armadillos as a whole, including extinct groups, but they also showed that the glyptodons do not represent an independent line of development within the armored articulated animals. To what extent a similar position also applies to the pampatheria is currently unclear from a genetic point of view, as no DNA material has yet been sequenced.

There are also a few other extinct groups within the Cingulata. These include the Peltephilidae , the Palaeopeltidae and the Pachyarmatheriidae . Their relationship to the armadillos is not entirely clear. The former could be very basal in the family tree of the armored articulated animals, the latter possibly form the sister group of the glyptodons and pampatherias.

Internal system

Internal systematics of the armadillos according to Gibb et al. 2015
  Dasypoda  
  Dasypodidae  

 Dasypus


  Chlamyphoridae  
  Euphractinae  

 Euphractus


   

 Chaetophractus


   

 Zaedyus




   
  Chlamyphorinae  

 Chlamyphorus


   

 Calyptophractus



  Tolypeutinae  

 Priodontes


   

 Tolypeutes


   

 Cabassous







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According to molecular genetic studies from 2015, today's armadillos are divided into two families, which are composed of nine genera and 20 species . One family includes the Dasypodidae, the long-nosed armadillos ( Dasypus ), the other, called Chlamyphoridae , includes all other known genera. According to the genetic data, the division of today's armadillos into these two lines of development began very early and took place during the Middle Eocene around 45 million years ago. However, the current armadillos represent only part of the former biodiversity. There are a total of four main lines (subfamilies) within the two families, to which a noteworthy part of the fossil-based representatives is assigned. The four main lines can be clearly demarcated both genetically and anatomically. The oldest line is represented by the Dasypodinae with the long-nosed armadillos ( Dasypus ). They also form the tribe of the Dasypodini . Furthermore, the extinct lines of the Stegotheriini and the Astegotheriini are to be added, the latter comprising the most primitive group of armadillos, which is already fossilized in the late Paleocene .

The other three lines appeared in quick succession. The Euphractinae were still formed in the Upper Eocene 37 million years ago , but fossils have been found for at least 42 million years. Includes, in addition to the eponymous six-banded armadillo ( Euphractus ) and the Pichi ( Zaedyus ) and the chaetophractus ( Chaetophractus on). They are united together in the tribe of the Euphractini , which also includes several extinct representatives. Opposite them are the Eutatini , an extinct line with numerous genera. The Eutatini belonged to the most diverse representatives of the armadillos, but they lived exclusively in southern South America, where they have been proven on the basis of partially well-preserved fossil finds, which also allow the determination that an extremely thick coat was developed as an adaptation to cool climates.

In the Oligocene around 33 million years ago, the Chlamyphorinae and the Tolypeutinae split off from each other, the last two great lines. The two representatives of the girdle mole ( Chlamyphorus and Calyptophractus ) are assigned to the Chlamyphorinae . No significant fossil material is available from either species, one extinct genus is documented with Chlamydophractus from the Upper Miocene . The Tolypeutinae bring together the ball armadillos ( Tolypeutes ), the giant armadillos ( Priodontes ) and the bare- tailed armadillos ( Cabassous ). The last two form the tribe of the Priodontini , while the former is part of the tribe of the Tolypeutini . Extinct lines are not distinguished here, but with Kuntinaru a very basic representative of the Tolypeutinae from the Oligocene is documented.

Overview of the recent and fossil genera of the armadillos

Around 60 genera are distinguished within the armadillos, some of these genera, such as Dasypus , Chasicotatus , Eutatus or Stegotherium , are very diverse in shape and sometimes include over half a dozen species. The structure is based on the overview by Malcolm C. McKenna and Susan K. Bell from 1997, but also takes into account recent developments in the higher classification of armadillos as well as newly introduced genera:

  • Dasypoda Quenstedt , 1885 ( Hoernes , 1886)
  • Family Dasypodidae Gray , 1821
  • Eocoleophorus Oliveira, Ribeiro & Bergqvist , 1997
  • Anadasypus Carlini, Vizcáıno & Scillato-Yané , 1997
  • Propraopus Ameghino , 1881
  • Dasypodon Castellani , 1925
  • Pliodasypus Castro, Carlini, Sánchez & Sánchez-Villagra , 2014
  • Dasypus Linnaeus , 1758 (long-nosed armadillos; including seven recent species)
  • Family Chlamyphoridae Bonaparte , 1850
  • Kuntinaru Billet, Hautier, de Muizon & Valentin , 2011
  • Pedrolypeutes Carlini, Vizcaíno. & Scillato-Yané , 1997
  • Tolypeutes Illiger , 1811 (spherical armadillos; including two recent species)
  • Priodontes Cuvier , 1825 (giant armadillo; including a recent species)
  • Cabassous McMurtrie , 1831 (Naked- tailed armadillos ; including four recent species)
  • Chlamydophractus Barasoain, Tomassini, Zurita, Montalvo & Superina , 2019
  • Chlamyphorus Harlan , 1825 (girdle gullet; including a recent species)
  • Calyptophractus Fitzinger , 1871 (Burmeister-Gürtelmull; including a recent species)

Taxonomy

One of the oldest drawings of an armadillo by Pierre Belon from 1553

Armadillos were first known in Europe at the beginning of the 16th century with the conquest of America by the Spanish conquistadors . Francisco Hernández de Córdoba , who discovered the Yucatán Peninsula in 1517 , used the local Aztec name Azotochtli as the name for the nine-banded armadillo , which translates as "turtle hare" and refers to the appearance of the animal. In 1553, Pierre Belon published one of the oldest known drawings of an armadillo and named the animals a Tatou . He put the animals next to the hedgehogs . More than 200 years later, in 1763, Georges-Louis Leclerc de Buffon gave a comprehensive overview of the species known at the time in his multi-volume work Histoire naturelle, générale et particuliére . Only five years earlier, however, Linnaeus had introduced the scientific name Dasypus for the nine-banded armadillo in the 10th edition of the Systema Naturae . He referred to Hernández for the origin of the word. It is assumed that Linnaeus considered the Aztec name Azotochtli unsuitable for a Latinization and thus for use in binary nomenclature and therefore used Dasypus (the word dasypus is of Greek origin ( δασύπους ) and means directly translated "rough feet", but was partially also related to the hare). At the same time, he finished the seven-banded armadillo ( Dasypus septemcinctus ), the six-banded armadillo ( Euphractus sexcinctus ), the Northern banded armadillo ( Tolypeutes tricinctus ) and the Southern naked-tailed armadillo ( Cabassous unicinctus ) also with the generic name Dasypus . He differentiated the individual species based on the number of movable belts.

John Edward Gray

A surname derived from Dasypus was introduced by John Edward Gray in 1821, with which he is now regarded as the first person to describe the family. At that time he chose Dasipidae as the name of the family and recognized Dasypus as the only genus . His brief description was:

Face long, conical: teeth, grinders cylindrical, separate from one another, seven or eight in each jaw, inside without enamel: tongue smooth, slightly extensible: body covered with shelly, scaly armor: ear and tail long: cœecum none.

“Face long, conical. Teeth, molars cylindrical, separated from each other, seven or eight in each jaw, without enamel inside. Tongue smooth, easily stretchable. Body covered with peeled, scaly armor. Long ears and tail. Appendicum missing. "

The correct form of the name, Dasypodidae, was first used in 1838 by Charles Lucien Jules Laurent Bonaparte , and he was also considered to be the family's first descriptor. For a long time the term was used for the entire group of armadillos. The division of the armadillos into two families (Dasypodidae and Chlamyphoridae), which was based on the results of molecular genetic studies in 2015, required a new name for the superordinate group. The name “Dasypoda” was chosen. The first use of the name is not clear. Friedrich August Quenstedt used it in 1885 in his handbook of petrefacts . However, Rudolf Hoernes used it during the same period. Reference is often made to the work Manuel de paleontologie , which appeared in Paris in 1886. The name Dasypoda can, however, already be found in the German-language original Elements of Paleontology published two years earlier . Both authors included the glyptodons in the Dasypoda, due to which the name was mostly synonymous with Cingulata . At the end of the 1970s, George F. Engelmann used the name Dasypoda to separate the armadillos from the Glyptodonta, in the latter he summarized the Glyptodontidae and the Pampatheriidae . This approach only lasted for a short time, among other things because Engelmann was only able to list a few separating characteristics between the two large groups. That the Dasypoda and the Glyptodonta form a line of development in Engelmann's sense was later shown by the skeletal anatomical investigations by Gaudin and Wible from 2006. This was also supported by the aforementioned genetic investigations on the fossil DNA of the glyptodons in 2016.

Dasypoda is also a higher-ranking homonym to the genus of the trouser bees ( Dasypoda ), which was introduced by Pierre André Latreille in 1802. The establishment of Dasypoda as an umbrella term for the armadillos met with criticism later due to the changeable usage in the past. Instead, the name Dasypodoidea was proposed, which in turn goes back to Gray.

etymology

The German trivial name "armadillo" refers to the movable straps or belts between the two fixed armor parts. The term armadillo used in the English- speaking world is derived from the Spanish word armado and means "little armored man". In the Spanish-speaking countries, especially in South America, the Guaraní word tatu is mainly used as a name for the armadillo.

Tribal history

Origin and Adaptive Radiation

Bone platelets from Riostegotherium , the oldest known representative of the armadillos

Overall, the tribal history of the armadillos has been little researched and there are still numerous gaps. More extensive finds are mainly from the Eocene , Miocene and Upper Pleistocene . The predominant fossil material is limited to the remains of the armor, skull and skeletal remains are much rarer. Overall, representatives of the group appear comparatively seldom in the fossil record , this also applies to many of today's armadillo species, which were only detected sporadically in the geological past.

The earliest armadillos show some significant differences from today's species. A striking feature is the different armor design. The back armor often only consisted of a fixed part, which was formed in the pelvic area, while the entire front section was made up of movable bands. Some forms also had an armor made only of movable bands. The armadillos' armor of today, with a fixed front and rear section separated by movable straps, was only created in the course of tribal history. If preserved in fossil form, enamel was still formed in the teeth of some early forms . This decreased in the course of evolutionary history. Today, an extremely thin and quickly chewed off enamel layer is only found in some long-nosed armadillos . The teeth were not typically shaped like nails, as is known today, but sometimes had a flat, rather lobate chewing surface. This is considered to be native to the secondary articulated animals and is also found in the Glyptodontidae , the Pampatheriidae and some of the extinct large ground sloths.

The first armadillos became fossilized in the Upper Paleocene around 58 million years ago. The findings from a crevice filling of the Itaboraí Formation near São José de Itaboraí in Itaboraí in southeastern Brazil , which include some bone platelets and a little postcranial skeletal material, are particularly significant . These are now part of the genus Riostegotherium , which belongs to the systematic proximity of the long-nosed armadillos ( Dasypus ).

Eocene

Bone platelets from Pucatherium (A – O) and Punatherium (P – Q)

In the Lower Eocene, the armadillos initially stayed in central and southern South America, which at that time was under the influence of tropical climates . From that phase mainly representatives of the Astegotheriini are proven. In what is now the Patagonia of Argentina, there are two sites, Laguna Fría and La Barda, on the central reaches of the Chubut River . Here mainly Prostegotherium and Stegosimpsonia occur, two rather small representatives of the early armadillos. Members of the Euphractinae are possibly first recorded in the Middle Eocene. Gran Barranca in central Patagonia is of outstanding importance here . The extremely complex site contained, among other things, remains of previously rare representatives such as Mazzoniphractus and Orthutaetus , but also more frequently documented forms such as Meteutatus , Parutaetus and Utaetus . The latter has numerous remains of the body skeleton, which can be used to reconstruct an animal weighing around 2 to 3 kg that still had teeth with enamel. Both Utaetus and Meteutatus , Parutaetus were also found in the Guabirotuba formation in the Brazilian state of Paraná, which is the same as the lower sections of Gran Barranca . There, several atypical osteoderms also refer to Proeocoleophorus , whose relationship is unclear.

In the Lumbrera Formation in north-western Argentina, which dates to the Lower and Middle Eocene, the armadillo forms known from Patagonia are largely missing. Instead, a skull find from Lumbreratherium, which is very rare for the early armadillos, comes from there . Its dentition was still heterodontic with a front canine-like tooth, which was separated from the rest of the molar-like teeth by a large gap ( diastema ). The rock unit also contained a partial skeleton of Pucatherium , which consists of the body armor and parts of the musculoskeletal system embedded in it. In contrast to Utaetus , whose armor had a firmly overgrown section on the basin, this consisted of flexible ligaments in both Lumbrerartherium and Pucatherium . Pucatherium was also described in the region from the Casa Grande , the Quebrada de los Colorados and the Geste formation . The latter also produced Punatherium, among other things . As a rule, however, only bone platelets have survived from the localities mentioned. In terms of time, they can be set a little later than the Lumbrera formation.

At that time, the first spread to parts of South America further north. Several species have been found near Santa Rosa on the Río Yurúa in eastern Peru, such as Yuruatherium and Parastegosimpsonia , which date to the late Eocene, the latter belongs to the Astegotheriini, the former cannot yet be precisely assigned.

Oligocene

Cooling climatic conditions in the transition from the Eocene to the Oligocene led to a significant increase in Euphractine armadillos, which obviously got along better with the prevailing temperatures. In the early part of the time phase, two genera appeared in Termas del Flaco in central Chile , Parutaetus and Meteutatus , which were detected there on the basis of the armor plates of the back shields, but also through numerous skull and postcranial skeletal material, including an almost complete armor. The subsequent succession in Gran Barranca in central Patagonia again shows a high diversity of armadillos. This includes Sadypus , Meteutatus and Barrancatatus from the Eutatini group as well as Parutaetus and Archaeutatus from the Euphractini group. From the Upper Oligocene, Kuntinaru has come down to us, a basal member of the Tolypeutinae without special group assignment, which could be described with the help of a skull from the Salla Beds of Bolivia near La Paz and is around 26 million years old. With Eocoleophorus an early representative of the Dasypodini was discovered in the Tabauté basin of Brazil, so that at that time three of the four main lines of the armadillos are tangible.

Miocene

Remnants of armor from Stegotherium

The very fossil-rich Santa Cruz Formation in the southernmost part of South America is of great importance for the Lower and Middle Miocene . The prevailing landscapes there at that time were subject to a humid subtropical climate and consisted of a mixture of open savannahs and partly closed forests, which gave rise to a richly shaped armadillo fauna. Stegotherium from the group of the Stegotheriini and their only representative, which has one of its earliest records here, but possibly already occurred in the Oligocene, is documented quite frequently . Among other things, it has been found in several sites near Lago Argentino , but has also been documented from the coastal areas along the Atlantic . The distinctly elongated snout and the teeth, which are significantly reduced in size, distinguish Stegotherium as a highly specialized form that mainly used ants and termites as a food source. It is the armadillos representative that has so far been most clearly adapted to this diet. The rock unit also contained an excellently preserved skeleton of Prozaedyus , which was about the size of today's dwarf armadillo. Also Proeutatus and Stenotatus are known here, the former was up to 15 kg while. All three representatives belong to the Euphractinae, with Prozaedyus representing the Euphractini, but the other two representing the Eutatini. It should be emphasized that there was an exchange within the Euphractinae at the beginning of the Miocene, as almost all older forms disappeared and were replaced by new ones.

The armadillos from the Santa Cruz Formation also form part of the fauna of other rock units of the same age in South America, such as the Chucal Formation in northern Chile and the Río Zeballos Formation in southern Chile. They are then still proven to the Middle Miocene, where finds come from the Cerdas Beds in southern Bolivia. However, some forms of the early Miocene survived until more recent times. This applies, for example, to Prozaedyus , which is documented with a very small relative from the Loma-de-Las-Tapias formation in north-western Argentina. This originated in the beginning of the Upper Miocene. The almost complete skull that was found shows an omnivore with its short snout. In the further course of the Middle Miocene, Pedrolypeutes appeared in the fossil discovery area of La Venta on the central reaches of the Río Magdalena in Colombia, a forerunner of the spherical armadillos and thus a member of the Tolypeutinae again after a gap of 12 to 14 million years. Other armadillos at the site are documented by Nanoastegotherium from the Astegotheriini group and Anadasypus from the Dasypodini group.

The late Middle and Upper Miocene is characterized primarily by the occurrence of numerous new members of the Eutatini and Euphractini. So here Macroeuphractus , Proeuphractus and Chasicotatus appear . All three genera have been identified from the Conglomerado osifero , an extremely fossil-rich deposit in northeastern Argentina. From the latter form, individual bone platelets also come from the Salicas formation in northwest Argentina, as do osteoderms from Macrochorobates . On Chasicotatus turn comes, among others, an almost complete carapace from the Arroyo Chasicó lineup in the Argentine province of Buenos Aires back, the same reference barg with Chlamydophractus the only previous evidence of fossil Gürtelmulls, from the next tank fragments a lower jaw and various elements of the The musculoskeletal system. There is here one of the oldest evidence of parasitism of fleas on armadillos. These are traces of feeding by the insects that nested in the armor of the animals living at the time. Most lines of the early armadillos largely died out towards the end of the Miocene or during the course of the following Pliocene .

Plio and Pleistocene

Skeleton of Eutatus

In the Pliocene , the Isthmus of Panama closed , creating a land bridge to North America . Then the Great American Fauna Exchange began. Initially, in the Pliocene, Macroeuphractus, one of the largest armadillo representatives, appeared, which reached the size of today's domestic pig . This has been proven , among other things, from northern Argentina near Buenos Aires . In addition, the first evidence of today's armadillos is known from this time. Such was Tolypeutes with the extinct species Tolypeutes pampaeus in the Pampas region until well into the outgoing Pleistocene spread in the same region developed Chaetophractus with the big hairy armadillo . This genus only reached its southern appearance today very late. Dasypus also originated in the Pliocene, with the northern part of South America being assumed to be the region of origin at that time due to the distribution of numerous closely related genera. The fossil species Dasypus bellus emerged first, which was found in North America at the end of the Pliocene and is documented quite frequently there, for example in Florida , but died out again at the end of the Pleistocene . This fate also met numerous other armadillo representatives in South America such as Propraopus , a close relative of Dasypus , or Eutatus from the line of the Eutatini, a large animal that reached the dimensions of today's giant armadillo . Dasypus did not return to North America until the 19th century with the nine-banded armadillo .

Armadillos and humans

Fuleco , 2014 World Cup mascot

Armadillos are often hunted in South America for their tasty meat , which, in addition to the destruction of their habitat by humans , has already led to the threatening decimation of some species. In contrast, the nine-banded armadillo , for example, reproduces constantly and continues to spread to North America . In some places the animals are resettled for pest control or kept as pets . Some species are considered a land scourge; They cause damage by digging for food or creating earthworks in agricultural areas or cause injuries in large grazing animals when they break into the burrows. The tanks of the dead animals are sometimes sold to tourists as bizarre baskets . Traditional musical instruments such as the charangos also consist of armor parts from some types of armadillos; however, their export from the countries of origin Bolivia and Peru is prohibited.

According to the IUCN , just under half of all species in the population are not endangered. The fur armadillo ( Dasypus pilosus ), the giant armadillo ( Priodontes maximus ) and the northern spherical armadillo ( Tolypeutes tricinctus ) are particularly endangered . Three other species - the girdle gullet ( Chlamyphorus retusus ), the Burmeister girdle gullet ( Calyptophractus truncatus ) and the Yungas armadillo ( Dasypus mazzai ) - cannot be classified in terms of their population threat due to a lack of data.

In addition to humans, armadillos are one of the few groups of mammals that can carry the bacterium that causes leprosy . A connection between their unusually low body temperature and the multiplication of mycobacteria is considered likely. This makes them indispensable when researching vaccines and new antibiotic combinations. But it also means that people who eat armadillos are at risk of contracting the disease. At least some of the leprosy cases in the United States are attributed to contact with armadillos. Furthermore, some armadillo species are also carriers of the single cell Trypanosoma cruzi , which causes Chagas disease , which is common in South America and can also occur in humans.

The armadillo Fuleco was the mascot of the 2014 World Cup in Brazil. The name is made up of the Portuguese words futebol ("football") and ecologia ("ecology") and should symbolize the protection of armadillos. A related call by scientists to better preserve local and supra-regional biodiversity has largely faded away.

literature

  • Alfred L. Gardner (Ed.): Mammals of South America, Volume 1: Marsupials, Xenarthrans, Shrews, and Bats. University of Chicago Press, 2008, pp. 128-157, ISBN 0-226-28240-6
  • CM McDonough and WJ Laughry: Dasypodidae (Long-nosed armadillos). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 30–47 ISBN 978-84-16728-08-4
  • Ronald M. Nowak: Walker's Mammals of the World . Johns Hopkins University Press, 1999, ISBN 0-8018-5789-9
  • Mariella Superina and Agustín Manuel Abba: Chlamyphoridae (Chlamyphorid armadillos). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 48–71 ISBN 978-84-16728-08-4
  • Sergio F. Vizcaíno and WJ Loughry (Eds.): The Biology of the Xenarthra. University Press of Florida, 2008, pp. 1-370, ISBN 978-0-8130-3165-1

Individual evidence

  1. ↑ a b c d e f g h i j k l C. M. McDonough and WJ Laughry: Dasypodidae (Long-nosed armadillos). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 30–47 ISBN 978-84-16728-08-4
  2. ↑ a b c d e f g h i j k l Mariella Superina and Agustín Manuel Abba: Chlamyphoridae (Chlamyphorid armadillos). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 48–71 ISBN 978-84-16728-08-4
  3. ↑ a b c Sergio E Vizcaíno and Gerardo De Iuliis: Evidence for advanced carnivory in fossil armadillos (Mammalia: Xenarthra: Dasypodidae). In: Paleobiology , 29 (1), 2003, pp. 123-138
  4. ↑ a b Darin A. Croft and Velizar Simeonovski: Horned armadillos and rafting monkeys. The fascinating fossil mammals of South America. Indiana University Press, 2016, pp. 1–304 (pp. 56–57 and 208–209)
  5. ↑ a b c Sergio F. Vizcaíno, M. Susana Bargo, Richard F. Kay and Nick Milne: The armadillos (Mammalia, Xenarthra, Dasypodidae) of the Santa Cruz Formation (early-middle Miocene): An approach to their paleobiology. In: Palaeogeography, Palaeoclimatology, Palaeoecology 237, 2006, pp. 255-269
  6. ↑ a b c d e f Paul Smith: The Xenarthra famalies Myrmecophagidae and Dasypodidae. Fauna Paraguay Handbook of the Mammals of Paraguay 2012, pp. 1-35
  7. ↑ a b c d e f g h i j Mariella Superina: Biology and keeping of armadillos (Dasypodidae). University of Zurich, 2000, pp. 1–248
  8. ^ A b Sergio F. Vizcaíno: The teeth of the “toothless”: novelties and key innovations in the evolution of xenarthrans (Mammalia, Xenarthra). In: Paleobiology 35 (3), 2009; Pp. 343-366
  9. ↑ Gerardo De Iuliis, María S. Bargo and Sergio F. Vizcaíno: Variation in skull morphology and mastication in the fossil giant armadillos Pampatherium spp. and allied genera (Mammalia: Xenarthra: Pampatheriidae), with comments on their systematics and distribution. In: Journal of Vertebrate Paleontology 20 (4), 2000, pp. 743-754
  10. ^ A b Daniela C. Kalthoff: Microstructure of Dental Hard Tissues in Fossil and Recent Xenarthrans (Mammalia: Folivora and Cingulata). In: Journal of Morphology 272, 2011, pp. 641-661
  11. ↑ Sergio F. Vizcaíno and María S. Bargo: Los armadillos (Mammalia, Dasypodidae) de La Toma (Partido de Coronel Pringles) y otros sitios arqueológicos de la provincia de Buenos Aires. Consideraciones paleoambientales. In: Ameghiniana 30, 1993, pp. 435-443
  12. ↑ a b c Alfredo A. Carlini, Martín Ricardo Ciancio, John J. Flynn, Gustavo J. Scillato ‐ Yané and André R. Wyss: The phylogenetic and biostratigraphic significance of new armadillos (Mammalia, Xenarthra, Dasypodidae, Euphractinae) from the Tinguirirican (early oligocene) of Chile. In: Journal of Systematic Palaeontology 7 (4), 2009, pp. 489-503
  13. ^ Matthew K. Vickaryous and Brian K. Hall: Osteoderm Morphology and Development in the Nine-Banded Armadillo, Dasypus novemcinctus (Mammalia, Xenarthra, Cingulata). In: Journal of Morphology 267 (11), 2006, pp. 1273-1283
  14. ^ Robert V. Hill: Comparative Anatomy and Histology of Xenarthran Osteoderms. In: Journal of Morphology 267, 2006, pp. 1441-1460
  15. ↑ CM Krmpotic, MR Ciancio, C. Barbeito, RC Mario and AA Carlini: Osteodermmorphology in recent and fossil euphractine xenarthrans. In: Acta Zoologica (Stockholm) 90, 2009, pp. 339-351
  16. ↑ CM Krmpotic, MR Ciancio, AA Carlini, MC Castro, AC Scarano and CG Barbeito: Comparative histology and ontogenetic change in the carapace of armadillos (Mammalia: Dasypodidae). In: Zoomorphology 134, 2015, pp. 601-616
  17. ↑ MR Ciancio, CM Krmpotic, AC Scarano and MB Epele: Internal Morphology of Osteoderms of Extinct Armadillos and Its Relationship with Environmental Conditions. In: Journal of Mammalian Evolution 26 (1), 2019, pp. 71-83, doi: 10.1007 / s10914-017-9404-y
  18. ↑ Irene H. Chen, James H. Kiang, Victor Correa, Maria I. Lopez, Po-Yu Chen, Joanna McKittrick and Marc A. Meyers: Armadillo armor: Mechanical testing and micro-structural evaluation. In: Journal of the Mechanical Behavior of Biomedical Materials 4 (5), 2011, pp. 713-722
  19. ↑ Cecilia M. Krmpotic, Fernando C. Galliari, Claudio G. Barbeito and Alfredo A. Carlini: Development of the integument of Dasypus hybridus and Chaetophractus vellerosus, and asynchronous events with respect to the postcranium. In: Mammalian Biology 77, 2012, pp. 314-326
  20. ↑ a b c d e f g Mariella Superina and WJ Loughry: Life on the Half-Shell: Consequences of a Carapace in the Evolution of Armadillos (Xenarthra: Cingulata). In: Journal of Mammalian Evolution 19, 2012, pp. 217-224
  21. ↑ Timothy J. Gaudin: The morphology of the xenarthrous vertebrae (Mammalia: Xenarthra). In: Fieldiana 1505 (41), 1999, pp. 1-38
  22. ↑ Fernando C. Galliari, Alfredo A. Carlini, Marcelo R. Sánchez-Villagra: Evolution of the axial skeleton in armadillos (Mammalia, Dasypodidae). In: Zeitschrift für Mammalskunde 75, 2010, pp. 326–333
  23. ^ A b c S. F. Vizcaíno and N. Milne: Structure and function in armadillo limbs (Mammalia: Xenarthra: Dasypodidae). In: Journal of Zoology 257, 2002, pp. 117-127
  24. ^ N. Milne, SF Vizcaíno and JC Fernicola. A 3D geometric morphometric analysis of digging ability in the extant and fossil cingulate humerus. In: Journal of Zoology 278 (1), 2009, pp. 48-56
  25. ^ Sergio F. Vizcaíno, Richard A. Fariña and Gerardo V. Mazzetta: Ulnar dimensions and fossoriality in armadillos. In: Acta Theriologica 44 (3), 199, pp. 309-320
  26. ↑ Nick Milne, Nestor Toledo and Sergio F. Vizcaíno: Allometric and Group Differences in the Xenarthran Femur. In: Journal of Mammalian Evolution 19, 2012, pp. 199-208
  27. ↑ M. Ciuccio, p Estecondo and EB Casanave: Scanning Electron Microscopy Study of the dorsal surface of the Tongue in Zaedyus pichiy (Mammalia, Xenarthra, Dasypodidae). In: International Journal of Morphology 26 (1), 2008. pp. 13-18
  28. ↑ Silvia Estecondo, Stella Maris Codón and Emma Beatriz Casanave: Histological Study of the Salivary Glands in Zaedyus pichiy (Mammalia, Xenarthra, Dasypodidae). In: International Journal of Morphology 23 (1), 2005. pp. 19-24
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