Cingulata, part of the superorder

Xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. Ex ...

, is an order of

armored Armour (British English) or armor (American English; see spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, especially direct contact weapons or projectiles during combat Co ...

New World The term ''New World'' is often used to mean the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxford University Press, p. 3 ...

placental Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsup ...

mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...

s. Dasypodids and chlamyphorids, the

armadillo Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, along wi ...

s, are the only surviving

families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...

in the order. Two groups of cingulates much larger than extant armadillos (maximum body mass of 45 kg (100 lb) in the case of the giant armadillo) existed until recently: pampatheriids, which reached weights of up to 200 kg (440 lb) and chlamyphorid

glyptodont Glyptodonts are an extinct subfamily of large, heavily armoured armadillos. They arose in South America around 48 million years ago and spread to southern North America after the continents became connected several million years ago. The best-kn ...

s, which attained masses of 2,000 kg (4,400 lb) or more. The cingulate order originated in

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

during the

Paleocene epoch The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''palaió ...

about 66 to 56 million years ago, and due to the continent's former isolation remained confined to it during most of the

Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...

. However, the formation of a

land bridge In biogeography, a land bridge is an isthmus or wider land connection between otherwise separate areas, over which animals and plants are able to cross and Colonisation (biology), colonize new lands. A land bridge can be created by marine regre ...

allowed members of all three families to migrate to southern North America during the

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58 or early

Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...

as part of the

Great American Interchange The Great American Biotic Interchange (commonly abbreviated as GABI), also known as the Great American Interchange and the Great American Faunal Interchange, was an important late Cenozoic paleozoogeographic biotic interchange event in which lan ...

. After surviving for tens of millions of years, all of the pampatheriids and giant

s apparently died out during the

Quaternary extinction event The Quaternary period (from 2.588 ± 0.005 million years ago to the present) has seen the extinctions of numerous predominantly megafaunal species, which have resulted in a collapse in faunal density and diversity and the extinction of key ecolog ...

at the beginning of the

Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...

, along with much of the rest of the regional

megafauna In terrestrial zoology, the megafauna (from Greek μέγας ''megas'' "large" and New Latin ''fauna'' "animal life") comprises the large or giant animals of an area, habitat, or geological period, extinct and/or extant. The most common threshold ...

, shortly after the colonization of the Americas by Paleo-Indians.

Description

Armadillos have dorsal armor that is formed by

osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...

s, plates of dermal bone covered in relatively small, overlapping

keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...

ized epidermal scales called "

scute A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterior po ...

s". Most species have rigid shields over the shoulders and hips, with three to nine bands separated by flexible skin covering the back and flanks. Pampatheres also had shells that were flexible due to three movable lateral bands of osteoderms. The osteoderms of pampatheres were each covered by a single scute, unlike those of armadillos, which have more than one. Glyptodonts, on the other hand, had rigid, turtle-like shells of fused osteoderms. Both groups have or had a cap of armor atop their heads. Glyptodonts also had heavily armored tails; some, such as ''

Doedicurus ''Doedicurus'', or ''Dædicurus'', is an extinct genus of glyptodont from South America containing one species, ''D. clavicaudatus''. Glyptodonts are a member of the family Chlamyphoridae, which also includes some modern armadillo species, an ...

'', had mace-like clubs at the ends of their tails, similar to those of

ankylosaur Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...

s, evidently used for defensive or

agonistic An agonist is a chemical that activates a receptor to produce a biological response. Receptors are cellular proteins whose activation causes the cell to modify what it is currently doing. In contrast, an antagonist blocks the action of the agon ...

purposes. Most armadillos eat insects and other invertebrates; some are more

omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutri ...

and may also eat small vertebrates and vegetable matter. Pampatheres are thought to have been specialized for

grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to roam around and consume wild vegetations in order to convert the otherwise indigestible (by human gut) cellulose within grass and other ...

, and isotopic analysis indicates the diet of glyptodonts was dominated by C4

grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...

es.

Euphractinae Euphractinae is an armadillo subfamily in the family Chlamyphoridae. Euphractinae are known for having a well developed osteoderm that has large cavities filled with adipose tissue, and more hair follicles with well developed sebaceous glands i ...

is unique for speciations towards

carnivory A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, adipose tissue, f ...

, culminating in the macropredatory genus ''

Macroeuphractus ''Macroeuphractus'' is a genus of extinct armadillos from the Late Miocene to Late Pliocene of South America. The genus is noted for its large size, with ''Macroeuphractus outesi'' being the largest non- pampathere or glyptodont armadillo disc ...

''.

Classification

The taxonomic table below follows the results of a

phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

published by Delsuc ''et al.'', 2016. While glyptodonts have traditionally been considered stem-group cingulates outside the group that contains modern armadillos, this 2016 study conducted an analysis of ''

mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...

and found that it was, in fact, nested within the modern armadillos as the

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...

of a

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

consisting of Chlamyphorinae and Tolypeutinae. Order Cingulata * Family † Protobradyidae Ameghino 1902 ** Genus †'' Protobradys'' Ameghino 1902 * Family † Peltephilidae Ameghino 1894 ** Genus †'' Anantiosodon'' Ameghino 1891 ** Genus †'' Epipeltephilus'' Ameghino 1904 ** Genus †'' Parapeltecoelus'' Bordas 1938 ** Genus †'' Peltecoelus'' Ameghino 1902 ** Genus †'' Peltephilus'' Ameghino 1887 (Horned armadillo) ** Genus †'' Ronwolffia'' Shockey 2017 * Family † Paleopeltidae Ameghino 1895 ** Genus †'' Palaeopeltis'' Ameghino 1895 * Family †

Pampatheriidae Pampatheriidae ("Pampas beasts") is an extinct family of large plantigrade armored armadillos related to extant armadillos in the order Cingulata. However, pampatheriids have existed as a separate lineage since at least the middle Eocene Muster ...

** Genus ? †''

Machlydotherium ''Machlydotherium'' is an extinct genus of cingulate of uncertain systematic affinities, perhaps belonging to the Pampatheriidae. It lived from the Middle Eocene to the Early Oligocene, and its fossilized remains were found in South America. ...

'' Ameghino 1902 ** Genus †''

Holmesina ''Holmesina'' is a genus of pampathere, an extinct group of armadillo-like creatures that were distantly related to extant armadillos. Like armadillos, and unlike the other extinct branch of megafaunal cingulates, the glyptodonts, the shell wa ...

'' Simpson 1930 ** Genus †''

Kraglievichia ''Kraglievichia'' is an extinct genus of cingulate belonging to the family Pampatheriidae. It lived from the Late Miocene to the Early Pliocene, and its fossilized remains were discovered in South America. Description This animal looked like ...

'' Castellanos 1927 ** Genus †''

'' ** Genus †''

Pampatherium ''Pampatherium'' is an extinct genus of xenarthran that lived in the Americas during the Pleistocene. Some species went extinct right at the Pleistocene-Holocene border. Distribution ''Pampatherium humboldtii'' and ''P. typum'' lived in South Ame ...

'' Ameghino 1875 ex Gervais & Ameghino 1880 ** Genus †''

Scirrotherium ''Scirrotherium'' is an extinct genus of pampatheres, a family of herbivorous cingulates, related to the similar but smaller modern armadillos, and with the now extinct glyptodonts, well-known from their shell-like armor. Its scientific name ...

'' Edmund & Theodor 1997 ** Genus †''

Tonnicinctus ''Tonnicinctus'' is an extinct species of pampatheriid that lived in Argentina during the Pleistocene and Holocene. Description ''Tonnicinctus'' inhabited cool grassland regions. It was a medium-sized pampatheriid, smaller than its better-know ...

'' Góis et al. 2015 ** Genus †''

Vassallia ''Vassallia'' is an extinct genus of cingulate belonging to the family Pampatheriidae. It lived between the Middle Oligocene and the Early Pliocene in what is now South America. Description This animal must have been very similar in shape a ...

'' Castellanos 1927 'Plaina''_Castellanos_1937.html" ;"title="Plaina.html" ;"title="'Plaina">'Plaina'' Castellanos 1937">Plaina.html" ;"title="'Plaina">'Plaina'' Castellanos 1937** Genus †''Yuruatherium'' Ciancio et al. 2012 * Family †Pachyarmatheriidae Fernicola et al. 2018 ** Genus †''

Neoglyptatelus ''Neoglyptatelus'' is an extinct genus of xenarthran, belonging to the order Cingulata. It lived from the Middle to the Late Miocene, and its fossilized remains are found in South America. Description This animal was roughly similar with mod ...

'' Carlini, Vizcaíno & Scillato-Yané 1997 ** Genus †''

Pachyarmatherium ''Pachyarmatherium'' is a genus of extinct large armadillo-like cingulates found in North and South America from the Pliocene and Pleistocene epochs, related to the extant armadillos and the extinct pampatheres and glyptodonts. It was presen ...

'' Downing & White 1995 * Family

Dasypodidae Dasypodidae is a family of mostly extinct genera of armadillos. One genus, '' Dasypus'', is extant, with at least seven living species. __TOC__ Classification Below is a taxonomy of armadillos in this family. Family Dasypodidae *† Genus ...

(long-nosed armadillos) ** Genus †'' Acantharodeia'' ** Genus †'' Amblytatus'' ** Genus †'' Archaeutatus'' ** Genus †''

Astegotherium ''Astegotherium'' is an extinct genus of xenarthran, belonging to the family Dasypodidae. It lived from the Early to the Middle Eocene, and its fossilized remains are found in Argentina. Description This genus is only known from the dermal pla ...

'' ** Genus †''

'' ** Genus †'' Barrancatatus'' ** Genus †'' Chasicotatus'' ** Genus †''

Chorobates The chorobates, described by Vitruvius in Book VIII of the '' De architectura'', was used to measure horizontal planes and was especially important in the construction of aqueducts. Similar to modern spirit levels, the chorobates consisted of ...

'' ** Genus †'' Coelutaetus'' ** Genus †'' Eocoleophorus'' ** Genus †'' Epipeltecoelus'' ** Genus †''

Eutatus ''Eutatus'' is an extinct genus of large armadillos of the family (biology), family Chlamyphoridae. It was endemic to South America from the Miocene, Early Miocene to Late Pleistocene, living from 17.5 Annum, Ma-11,000 years ago, with possible su ...

'' ** Genus †'' Hemiutaetus'' ** Genus †'' Isutaetus'' ** Genus †'' Lumbreratherium'' ** Genus †'' Macrochorobates'' ** Genus †'' Mazzoniphractus'' ** Genus †'' Meteutatus'' ** Genus †'' Pedrolypeutes'' ** Genus †'' Prodasypus'' ** Genus †''

Proeutatus ''Proeutatus'' is an extinct genus of xenarthran, belonging to the order Cingulata. It lived during the Early Miocene, and its fossilized remains were discovered in South America. Description This animal was vaguely similar to the extant six-b ...

'' ** Genus †'' Prostegotherium'' ** Genus †'' Pucatherium'' ** Genus †'' Punatherium'' ** Genus †''

Stegotherium ''Stegotherium'' is an extinct genus of long-nosed armadillo, belonging to the Dasypodidae family alongside the nine-banded armadillo. It is currently the only genus recognized as a member of the tribe Stegotheriini. It lived during the Early Mio ...

'' ** Genus †'' Stenotatus'' ** Genus †''

Utaetus ''Utaetus'' is an extinct genus of mammal in the order Cingulata, related to the modern armadillos. The genus contains a single species, ''Utaetus buccatus''. It lived in the Late Paleocene to Late Eocene (about 60 to 36 million years ago) and i ...

'' ** Genus †'' Vetelia'' ** Subfamily

Dasypodinae Dasypodidae is a family of mostly extinct genera of armadillos. One genus, '' Dasypus'', is extant, with at least seven living species. __TOC__ Classification Below is a taxonomy of armadillos in this family. Family Dasypodidae *† Genu ...

*** Genus †'' Anadasypus'' *** Genus '' Dasypus'' *** Genus †''

Nanoastegotherium ''Nanoastegotherium'' is an extinct genus of cingulate, belonging to the family Dasypodidae, which includes the modern nine-banded armadillos. The name of the genus means "small ''Astegotherium''", referring to its small size, smaller than the m ...

'' *** Genus †'' Parastegosimpsonia'' *** Genus †'' Pliodasypus'' *** Genus †'' Propraopus'' *** Genus †'' Riostegotherium'' *** Genus †'' Stegosimpsonia'' * Family

Chlamyphoridae Chlamyphoridae is a family of cingulate mammals. While glyptodonts have traditionally been considered stem-group cingulates outside the group that contains modern armadillos, there had been speculation that the extant family Dasypodidae could b ...

: glyptodonts and other armadillos ** Subfamily

Chlamyphorinae Chlamyphorinae is a subfamily of South American armadillos in the family Chlamyphoridae. Members of this subfamily, the fairy armadillos, are largely fossorial and have reduced eyes and robust forearms with large claws for digging. __TOC__ Ta ...

: fairy armadillos *** Genus '' Calyptophractus'' *** Genus '' Chlamyphorus'' ** Subfamily

: hairy, six-banded and pichi armadillos *** Genus '' Chaetophractus'' *** Genus †'' Doellotatus'' *** Genus ''

Euphractus The six-banded armadillo (''Euphractus sexcinctus''), also known as the yellow armadillo, is an armadillo found in South America. The sole extant member of its genus, it was first described by Swedish zoologist Carl Linnaeus in 1758. The six- ...

'' *** Genus †''

'' ***Genus †'' Peltephilus'' *** Genus †''

Proeuphractus ''Proeuphractus'' is an extinct genus of xenarthran, related to the modern armadillos. It lived from the Early to the Late Miocene, and its fossilized remains were discovered in South America. Description This animal was quite similar to the ...

'' *** Genus †'' Paleuphractus'' *** Genus ''

Zaedyus The pichi (''Zaedyus pichiy''), dwarf armadillo or pygmy armadillo is an armadillo native to Argentina. It is the only living member of the genus ''Zaedyus'', and the only armadillo to hibernate. Description Pichis are relatively small armadi ...

'' ** Subfamily †

Glyptodontinae Glyptodonts are an extinct subfamily of large, heavily armoured armadillos. They arose in South America around 48 million years ago and spread to southern North America after the continents became connected several million years ago. The best-kn ...

: glyptodonts *** Genus †''

'' *** Genus †''

Glyptodon ''Glyptodon'' (from Greek for 'grooved or carved tooth': γλυπτός 'sculptured' and ὀδοντ-, ὀδούς 'tooth') is a genus of glyptodont (an extinct group of large, herbivorous armadillos) that lived from the Pleistocene, around 2.5 ...

'' *** Genus †'' Glyptotherium'' *** Genus †''

Hoplophorus ''Hoplophorus'' is an extinct genus of glyptodont, a subfamily of armadillos . The only confidently known species was ''H. euphractus'', found in Pleistocene deposits in Brazil, though fossils possibly from another species are known from Bolivia. ...

'' *** Genus †''

Panochthus ''Panochthus'' is an extinct genus of glyptodont, which lived in the Gran Chaco- Pampean region of Argentina ( Lujan, Yupoí and Agua Blanca Formations), Brazil ( Jandaíra Formation), Bolivia (Tarija and Ñuapua Formations), Paraguay and Urug ...

'' *** Genus †''

Parapropalaehoplophorus ''Parapropalaehoplophorus septentrionalis'' was a comparatively small (compared to ''Glyptodon'') species of glyptodont, extinct relatives of the modern armadillo. The mammal, identified in 2007 from the fossilized remains of a specimen found in 2 ...

'' *** Genus †'' Plaxhaplous'' ** Subfamily

Tolypeutinae Tolypeutinae is a subfamily of armadillos in the family Chlamyphoridae, consisting of the giant, three-banded and naked-tailed armadillos. __TOC__ Taxonomy It contains the following genera: *''Cabassous'' *''Kuntinaru'' *''Priodontes'' *''To ...

: giant, three-banded and naked-tailed armadillos *** Genus ''

Cabassous ''Cabassous'' is a genus of South and Central American armadillos. The name is the Latinised form of the Kalini word for "armadillo". Cladogram of living ''Cabassous'' The genus contains the following four species In biology, a species ...

'' *** Genus †''

Kuntinaru ''Kuntinaru'' is an extinct genus of tolypeutine chlamyphorid armadillo which existed in Bolivia, during the Late Oligocene ( Deseadan age). It is known from the holotype MNHN-SAL 1024, skull missing the apex of the rostrum and the paratype M ...

'' *** Genus ''

Priodontes The giant armadillo (''Priodontes maximus''), colloquially ''tatu-canastra'', ''tatou'', ''ocarro'' or ''tatú carreta'', is the largest living species of armadillo (although their extinct relatives, the glyptodonts, were much larger). It lives ...

'' *** Genus ''

Tolypeutes The genus ''Tolypeutes'' contains the two species of three-banded armadillos. They are restricted to open and semi-open habitats in South America. Of the several armadillo genera, only ''Tolypeutes'' rely heavily on their armor for protection. ...

References

{{Taxonbar, from=Q1242326 Xenarthrans Mammal orders Taxa named by Johann Karl Wilhelm Illiger Extant Thanetian first appearances