Designed by Paul Smith 2006. This website is copyrighted by law.
Material contained herewith may not be used without the prior written permission of FAUNA Paraguay.
Photographs on this web-site were taken by Paul Smith, Hemme Batjes, Regis Nossent,
Alberto Esquivel, Arne Lesterhuis, José Luis Cartes, Rebecca Zarza and Hugo del Castillo and are used with their permission.
Material contained herewith may not be used without the prior written permission of FAUNA Paraguay.
Photographs on this web-site were taken by Paul Smith, Hemme Batjes, Regis Nossent,
Alberto Esquivel, Arne Lesterhuis, José Luis Cartes, Rebecca Zarza and Hugo del Castillo and are used with their permission.
Click on the species links for access to the image gallery.
Click on the HBook icon on the right hand side for access to pdfs of the FAUNA Paraguay Handbook of the Mammals of Paraguay Volume 2 Xenarthra.
See below for a key to the Paraguayan species.
FAMILY DASYPODIDAE - ARMADILLOS
General characteristics: Twelve species recorded in Paraguay. The back is rounded and armoured with a carapace covered with horny scales and with bony dermal scutes forming movable bands across the midsection. Both the forepart and the hind part of the carapace are immovable. The underside of the animal is hairy and lacks armour. Armadillos run with rapid movements of their short, strong legs. They have five toes on the hindfoot and three to five toes on the forefoot, each marked with strong claws. Forelimb structure is specialised for digging, exceptionally so in some species. Hindlimb structure is associated with weight bearing and measurement indices are correlated with body size. Most species are excellent diggers, constructing species-specific caves with a rounded roof in which they sleep and raise their young. Armadillos are generally solitary, approachable and tame, with poor vision but an excellent sense of smell. The Guaraní common name is most often used for the species in Paraguay, the Spanish names being understood but rarely used in everyday speech. The Guaraní word for armadillo, "tatu" is also used in vulgar speech in reference to female genitalia! The family is ancient, dating from the late Paleocene of South America.
Dental characteristics: The long mandibles are lined with a varying number of semicircular "molariform" teeth which grow continuously through life. These teeth lack enamel. Dental formula is variable and canines and incisors are lacking.
Skeletal characteristics: More fossorial species showing a greater length of the olecronon process of the ulna and more robust humerus.
Taxonomy: According to Möller-Krull et al (2007) the Dasypodidae are split into four subfamilies - Dasypodinae, Chlamyphorinae, Tolypeutinae and Euphractinae.
Subfamily Dasypodinae Gray, 1821
One genus and three species present in Paraguay. Characteristics are discussed under the generic account below.
Dasypus Linnaeus, 1758: Long-nosed Armadillos
There are seven species in this genus, two present in Paraguay. Synonyms adapted from Gardner (2007).
Synonyms:
Dasypus Linnaeus 1758:50. Type species Dasypus novemcinctus Linnaeus (1758) by tautonomy.
Tatus Fermin 1769:110. Name unavailable.
Tatu Frisch 1775:Table. Name unavailable.
Tatu Blumenbach 1779:94. Type species Tatu novemcinctus (=Dasypus novemcinctus Linnaeus 1758) by monotypy.
Cataphractus Storr 1780:40. Proposed for all armadillos known at the time.
Loricatus Desmarest 1804:28. In part. Type species Loricatus niger Desmarest (1804) (=Dasypus novemcinctus Linnaeus 1758).
Tatus Olfers 1818:220. In part. Incorrect spelling.
Tatusia Lesson 1827:309. In part. Type species Dasypus peba Desmarest (1822) (=Dasypus novemcinctus Linnaeus 1758).
Cachicamus McMurtie 1831:163. Type species Dasypus novemcinctus Linnaeus (1758).
Cachicama P.Gervais in I.Geoffroy St.-Hilaire 1835:53. Invalid emendation.
Zonoplites Gloger 1841:114. Name proposed for armadillos with four toes on forefeet and two middle toes longer than outer toes.
Praopus Burmeister 1854:295. Type species Dasypus longicaudis Wied-Neuwied (1826) by monotypy.
Cryptophractus Fitzinger 1856:123. Type species Cryptophractus pilosus Fitzinger (1856) by monotypy.
Hyperoambon W.Peters 1864:180. Type species Dasypus pentadactylus W.Peters (1864). Valid as a subgenus.
Muletia Gray 1874:244. Type species Dasypus septemcinctus Gray (1874) (=Loricatus hybridus Desmarest 1804, not Dasypus septemcinctus Linnaeus 1758) by monotypy.
Tatua W.Robinson & Lyon 1901:161. Incorrect spelling.
Mulletia Yepes 1928:506. Incorrect spelling.
Mulieta Talmage & Buchanan 1954:80. Incorrect spelling.
General characteristics: Small to large armadillos that are easily recognized by their long, naked, tubular ears, set close together on the head, long snout (55% or more of head length) and long, pointed tail (<50% of head and body length). The carapace is smooth, high and rounded, the denticles placed in irregular circles. The proximal two-thirds of the tail is ringed, each ring formed by two or more rows of scutes. There are four toes on the forefeet (digits 2 and 3 the longest) and five on the hindfeet (digit 3 the longest). Diet consists mainly of insects, but somewhat omnivorous, also taking small vertebrates and fruit. The earlies fossils date from the Pliocene of South America.
Cranial characteristics: Palate extends way beyond the last molar. Brain case is high and rounded, becoming thinner towards rear. Tympanic ring represents an otherwise unossified bulla.
Dental characteristics: Dental formula 7-9/7-9 = 28-36
Dasypus hybridus - Southern Long-nosed Armadillo
Dasypus novemcinctus - Nine-banded Armadillo
Subfamily Euphractinae Winge, 1823
General characteristics: Three species in Paraguay in two genera. These are small to medium-sized armadillos with a broad head, short snout and short ears. Conspicuous body hair on the carapace, sides and venter. The cephalic shield is broad and forms a ledge over the orbit. Scutes along the ventral margin of the carapace are rounded anteriorly and pointed posteriorly. The forefoot possesses four toes, claws on digits II and III being longest.
Cranial characteristics: Auditory bullae and external auditory meatus are ossified.
Chaetophractus Fitzinger, 1871: Hairy Armadillos
There are three species in this genus, two present in Paraguay. Synonyms adapted from Gardner (2007).
Synonyms:
Loricatus Desmarest 1804:28. In part.
Dasypus Illiger 1870:70. In part. Not Dasypus Linnaeus (1758).
Tatus Olfers 1816:220. In part. Incorrect spelling of Tatu Blumenbach (1779).
Tatusia Lesson 1827:309. In part.
Euphractus Burmeister 1861:427. In part. Not Euphractus Wagler (1830).
Chaetophractus Fitzinger 1871:268. Type species Dasypus villosus (=Loricatus villosus Desmarest 1804) by designation.
Dasyphractus Fitzinger 1871:264. Type species Dasyphractus brevirostris by monotypy.
Choetophractus Trouessart 1898:1146. Incorrect spelling.
General characteristics: Two species of small to medium-sized armadillos. Head is armed with a dermal shield that reaches to the nose. The medium-length ears are well-separated with rounded ends. The dorsal scutes are arranged in a linear fashion and covered in sparse, stiff hairs, with six to eight flexible bands across the centre. The legs and underparts are densely haired. Armadillos in this genus are omnivorous, with some suggestion of seasonal diet changes. The genus is known from the Pliocene of South America.
Cranial characteristics: Marked post-orbital constriction. Broad zygomatic arch. Brain case expanded and flattened dorsolaterally. Palatine terminates in a V-shape.
Dental characteristics: Dental formula 9/10 = 36
Chaetophractus vellerosus - Lesser Hairy Armadillo
Chaetophractus villosus - Greater Hairy Armadillo
Euphractus Wagler, 1830: Six-banded Armadillo
This is a monotypic genus. Synonyms adapted from Gardner (2007).
Synonyms:
Dasypus Linnaeus 1758:50. In part.
Loricatus Desmarest 1804:28. In part.
Tatus Olfers 1818:220. In part. Incorrect spelling.
Euphractus Wagler 1830:36. Type species Dasypus sexcinctus Linnaeus (1758) by designation.
Encoubertus McMurtie 1831:163. In part. Proposed as a subgenus of Dasypus Linnaeus (1758).
Pseudotroctes Gloger 1841:113. Type species Dasypus setosus Wied-Neuwied (1826) by monotypy.
Scleropleura Milne-Edwards 1871:178. Type species Scleropleura bruneti Milne-Edwards (1871) by monotypy.
Scelopleura Trouessart 1898:1141. Incorrect spelling.
General characteristics: One large species. The carapace is flattened and with a fine layer of stiff pale hairs protruding from between the dorsal bands. The head is triangular in shape and the ears are short. Forefeet with five digits, the third being the longest. Two to four openings for scent glands present in the mid-dorsum of the pelvic shield. An active burrower, digging for food and to escape enemies. Omnivorous, in addition to insects taking fruit, small vertebrates and even carrion. The genus is known from the mid-Pleistocene of South America.
Dental characteristics: Dental formula 9/10 = 36
Euphractus sexcinctus - Six-banded Armadillo
Subfamily Tolypeutinae Gray, 1865
General characteristics: This subfamily contains the most specialised fossorial armadillos and the most powerful diggers, though notably the genus Tolypeutes is non-fossorial. Three genera and four species present in Paraguay. The subfamily is split into two tribes, the Priodontini (Cabassous and Priodontes) and the Tolypeutini (Tolypeutes).
Tribe Priodontini Gray, 1873
General characteristics: Small to extremely large armadillos with a blunt, rounded snout, thick fleshy ears and relatively large eyes. The carapace is domed and separated from the cephalic plate by three rows of nuchal scutes. Scapular and pelvic shields are separate by 11 to 14 movable bands. Body hair is sparse. Five toes on the forefoot with digit III armed with a scythe-shaped claw.
Cranial characteristics: Frontal bones have a dome-like swelling. Only the tympanic ring of each bulla is ossified.
Cabassous McMurtrie, 1831: Naked-tailed Armadillos
Four species in this genus, three present in Paraguay. Synonyms adapted from Gardner (2007):
Synonyms:
Dasypus Linnaeus 1758:50. In part.
Tatus Olfers 1818:220. In part. Incorrect spelling of Tatu Blumenbach (1779).
Xenurus Wagler 1830:36. Type species Dasypus gymnurus Wied-Neuwied (1826) (=Xenurus squamicaudis Lund 1845) by monotypy. Preoccupied.
Tatusia Lesson 1827:309. In part.
Cabassous McMurtrie 1831:164. Type species Dasypus unicincttus Linnaeus (1758) by monotypy. Proposed as a subgenus of Dasypus Linnaeus (1758).
Arizostus Gloger 1841:114. Type species Dasypus gymnurus (=Tatus gymnurus Olfers 1818) by monotypy.
Tatoua Gray 1865:378. Type species Dasypus unicincttus Linnaeus (1758) by monotypy. Proposed as a subgenus of Dasypus Linnaeus (1758).
Ziphila Gray 1873:22. Type species Ziphilus lugubris Gray (1873) by monotypy.
Lysiurus Ameghino 1891:254. Proposed as replacement name for Xenurus Wagler (1830).
Cabassus Trouessart 1905:820. Incorrect spelling.
Cabassus Neveu-Lemaire & Grandidier 1911:103. Incorrect spelling.
General characteristics: Three species of medium-sized armadillo characterized by the fairly long, thin tail lacking well-developed plates (hence “naked-tailed”). The snout is very short and broad with a blunt end and the nostrils are guarded by a fine line of bristles. Ears funnel-shaped and capable of folding flat, closing the external auditory meatus. All feet possess five toes. Digits three, four and five of the foreefeet are broad and scythe-shaped and the claw on the first digit is long and slender. Hindfoot with blunt claws, the third being the longest. Carapace ovoid, dome-shaped and quite flexible with 11 to 14 movable bands. Dorsal plates are in transverse rows across the length of the body. These are nocturnal, solitary animals that give birth to a single young. The diet consists largely of ants and termites, these armadillos burrow frequently and often forage underground. The genus is known from the mid-Pleistocene of Brazil.
Cranial characteristics: Frontal bone with dome-like expansion. Rostrum short and tympanic ring present. Elongated infraorbital canal. Distance from lacrimal foramen and anterior opening of infraorbital foramen one-quarter to one-third rostral length. Mandible slender with articular process higher than coronoid process.
Dental characteristics: Dental formula 7-10/8-9 =30-38. Teeth peg-shaped and vary from circular to oviod in form on either axis.
Skeletal characteristics: 7 cervical vertebrae, 12 (or 12-13) thoracic, 3 (or 3-4) lumbar, 10 (or 9-11) sacral and 18 (or 15-20) caudal.
Cabassous chacoensis - Chaco Naked-tailed Armadillo
Cabassous tatouay - Greater Naked-tailed Armadillo
Cabassous unicinctus - Southern Naked-tailed Armadillo
Priodontes F. Cuvier, 1825: Giant Armadillo
This is a monotypic genus. Synonyms adapted from Gardner (2007).
Synonyms:
Dasypus Kerr 1792:112. In part. Not Linnaeus (1758).
Loricatus Desmarest 1804:28. In part.
Tatus Olfers 1818:220. In part. Incorrect spelling of Tatu Blumenbach (1779).
Priodontes F.Cuvier 1825:527. Type species Dasypus gigas G.Cuvier (1817) by monotypy.
Cheloniscus Wagler 1830:35. Type species Dasypus gigas G.Cuvier (1817) by monotypy.
Priodon McMurtrie 1831:164. Type species Dasypus gigas G.Cuvier (1817) by monotypy. Proposed as a subgenus of Dasypus Linnaeus (1758). Preoccupied by Priodon Quay & Gaimard (1824).
Polygomphius Gloger 1841:114. Type species Dasypus gigas G.Cuvier (1817) by monotypy.
Prionodon Gray 1843:xxvii. Nomen nudum.
Priodonta Gray 1843:xxvii. Incorrect spelling.
Prionodontes Schinz 1845:312. Incorrect spelling.
Prionodos Gray 1865:374. Introduced as a replacement for Priodontes F.Cuvier (1825).
General characteristics: A single species of huge size, much the largest member of its family. The snout is rounded and blunt-ended. Ears and eyes are large. Carapace is divided into bands of which the central 11 to 13 are flexible. The tail is covered with tightly-packed plates that are not arranged in rows. Feeds almost exclusively on ants and termites, destroying their nests with the long, sharply hooked claws of the forefeet (third digit being especially long). This species digs habitually, creating diagnostic large burrows which are occupied for a single night. Such burrows are often dug into termite mounds and a large number may be clumped into a single small area.
Dental characteristics: Highly variable with as many as 20 in a single tooth row, many of which are lost as the animal ages.
Priodontes maximus - Giant Armadillo
Tribe Tolypeutini Gray, 1865
Consists of a single genus of which one species is present in Paraguay. Characteristics are discussed under the generic account below.
Tolypeutes Illiger, 1811: Three-banded Armadillo
Two species, one present in Paraguay. Synonyms adapted from Gardner (2007)
Synonyms:
Tolypeutes Illiger 1811:111. Type species Dasypus tricinctus Linnaeus (1758) by designation (Yespes 1928).
Matacus Rafinesque 1815:57. Nomen nudum.
Tolypeutis Olfers 1818:221. Incorrect spelling.
Tatusia Lesson 1827:309. In part.
Apara McMurtrie1831:163. Type species Dasypus tricinctus Linnaeus (1758) by monotypy. Proposed as a subgenus of Dasypus Linnaeus (1758) .
Sphaerocormus Fitzinger 1871:376. Type species Tolypeutes conurus I. Geoffroy St.Hilaire (1847) by monotypy.
Cheloniscus Gray 1873:23. Type species Cheloniscus tricinctus Gray (1873) (=Dasypus tricinctus Linnaeus (1758) by monotypy. Preoccupied.
Tolypentes Matschie 1894:62. Incorrect spelling.
Tolypoïdes Grandidier & Neveu-Lemaire 1905:370. Type species Tolypoïdes bicinctus Grandidier & Neveu-Lemaire (1905) by monotypy.
Tolypuetes Talmage & Buchanan 1954:73. Incorrect spelling.
General characteristics: A single species of small armadillo possessing a uniquely short, stumpy tail covered in tubercles. The ears are fairly large and rounded and the carapace is high, rounded and almost without hairs. Pectoral and pelvic shields are convex and extremely hard separated by 2-4 (usually 3) movable bands. Head shield is is elongated, flattened and with rounded lateral and posterior margins. Body hair is conspicuous below the carapace. In the Paraguayan species there are 4 toes on the forefeet and five toes on the hindfeet with digits II to IV unified and bearing hoof-like claws. Feeds almost exclusively on ants, termites and soft-bodied invertebrates. When threatened it takes flight rather than burrowing as do other armadillos, characteristically rolling into a ball when captured. The earliest record of the genus is from the Pliocene of South America.
Tolypeutes matacus - Southern Three-banded Armadillo
Subfamily Chlamyphorinae Bonaparte 1850
General characteristics: This tribe contains the most fossorial of all armadillos. The ears are small and eyes tiny. Head shield is reduced to a thin flexible plate which is separated from the rounded rump plate. The sides of the body and ventral region is covered with long, thick whitish hair. Forefoot possesses five digits with disproportionately large claws - the species is adapted for "fast digging" in soft soils as opposed to "power digging" in hard substrates. One genus and one species present in Paraguay.
Calyptophractus Fitzinger, 1871: Fairy Armadillos
This is a monotypic genus. Synonyms adapted from Gardner (2007).
Synonyms:
Chlamyphorus Burmeister 1863:167. Not Chlamyphorus Harlan (1825).
Burmeisteria Gray 1865:381. Type species Burmeisteria retusa (=Chlamyphorus retusus Burmeister 1863) by monotypy. Preoccupied by Burmeisteria Salter (1865).
Calyptophractus Fitzinger 1871:388. Type species Chlamyphorus retusus Burmeister (1863) by monotypy.
General characteristics: A single, distinctive species of small armadillo. Head shield is broad and carapace is thin and flexible, attached along the sides of the body. The tail is extremely short, flattened dorsoventrally and with a pointed tip. The rump plate is rounded, abruptly flattened and acts as a “plug” when burrowing. The genus is known from the Pleistocene of South America.
Cranial characteristics: External auditory meatus is long, extending forwards above the zygomatic arch to the posterior border of the orbit.
Calyptophractus retusus - Chaco Fairy Armadillo
Key to Adults of the Paraguayan Species of Armadillos
1a Carapace complete on head and dorsum, with movable bands on latter. Total length >25cm.........................2
1b Carapace incomplete, reduced to small dorsal plates on head and along spine. Heavily furred. Total length tiny <16cm...........................................................................................................................Calyptophractus retusus
2a Forefeet with one or more greatly enlarged, scythe-like foreclaws....................................................................3
2b Forefeet lacking greatly enlarged, scythe-like foreclaws...................................................................................6
3a Enormous, total length >140cm, body length >70cm.........................................................Priodontes maximus
3b Total length <80cm, body length <50cm.........................................................................................................4
4a Three movable bands across dorsal surface. Curls into ball for defence - does not dig. Tail short, stumpy and covered in bony tubercles..........................................................................................................Tolypeutes matacus
4b >10 movable bands across dorsal surface. Digs rapidly to escape persecution. Tail long, thin and "naked" - lacking well-developed plates................................................................................................................Cabassous 5
5a Ears long and funnel-shaped extending above the head. Distributed in eastern Paraguay............................ 10
5b Ears short and rounded do not extend above the head. Fleshy expansion on anterior margin of ear. Distributed in the Chaco...........................................................................................................Cabassous chacoensis
6a Stiff dorsal hairs present on carapace. Ears well-separated at base. Snout short and blunt..............................8
6b Carapace steeply rounded and lacking dorsal hairs. Ears joined at base. Snout long and tubular........Dasypus 7
7a Total length >60cm. Tail equal to or greater than head and body length. 8 or 9 movable bands with distinct pale edges. Mean of 60 scutes along 4th movable band..........................................................Dasypus novemcinctus
7b Total length <50cm. Tail equal to or shorter than head and body length. 6 or 7 movable bands .......................................................................................................................................................Dasypus hybridus
8a Total length <40cm. Ears long, projecting well above head and extending to the second row of scapular scutes when laid back. Carapace somewhat rounded. Lacks holes in pelvic shield.............Chaetophractus vellerosus
8b Total length >40cm. Ears short and rounded. Carapace flattened dorsally. Holes in pelvic shield................9
9a Scutes typically dark reddish brown, guard hairs dark. 7 or 8 movable bands (never 6) between scapular and pelvic shields, at least 2 movable neck bands. Scutes of head shield small and disorganised .Chaetophractus villosus
9b Scutes typically pale yellow-brown, guard hairs pale. 6 to 8 movable bands between scapular and pelvic shields and one movable neck band. Scutes of head shield large and well-organised...............Euphractus sexcinctus
10a Large (<36cm Head body length) with shield colouration usually reddish. Mean number of cephalic scutes 48.3 (+/-3.7). Ear length >40mm. Number of scutes on 4th movable band 30.8 (+/-1.6). Usually in red soil areas throughout eastern Paraguay. ............................................................................................ Cabassous tatouay
10b Medium-sized (<35cm Head body length) with shield colouration usually dark brownish or blackish. Mean number of cephalic scutes 54 (+/-5.5). Ear length <30mm. Number of scutes on 4th movable band 27.4 (+/-1.3). Usually in sandy soil areas in the cerrado zone of north-eastern Paraguay. .................. Cabassous unicinctus
Online Resources
Edentate Specialist Group - Publishes the online journal Edentata dealing with armadillos, sloths and anteaters.
REFERENCES
Diaz MM & Barquez RM 2002 - Los Mamíferos de Jujuy, Argentina - LOLA, Buenos Aires
Emmons LH & Feer F 1999 - Mamíferos de los Bosques Húmedos de América Tropical - FAN Bolivia
Gardner AL 2007 - Mammals of South America Volume 1: Marsupials, Xenarthrans, Shrews and Bats - University of Chicago Press.
Möller-Krull M, Delsuc F, Churakov G, Marker C, Superina M, Brosius J, Douzery EJP, Schmitz J 2007 - Retroposed Elements and Their Flanking Regions Resolve the Evolutionary History of Xenarthran Mammals (Armadillos, Anteaters and Sloths) - Molecular Biology and Evolution 24: p2573-2582.
Redford K & Eisenberg JF 1992 - Mammals of the Neotropics Vol 2: The Southern Cone Chile, Argentina, Uruguay and Paraguay - University of Chicago Press.
Smith P. Owen R, Atkinson K, del Castillo H, Northcote-Smith E 2011 - First Records of Southern Naked-tailed Armadillo Cabassous unicinctus (Linnaeus, 1758) (Xenarthra: Dasypodidae) in Paraguay - Edentata 12.
Vizcaíno SF, Milne N 2002 - Structure and Function in Armadillo Limbs (Mammalia: Xenarthran: Dasypodidae) - Journal of Zoological Society of London 257: p117-127.
Wetzel RM 1980 - Revision of the Naked-tailed Armadillos Genus Cabassous McMurtie - Annals of the Carnegie Museum of Natural History 49: p323-357.
Click on the HBook icon on the right hand side for access to pdfs of the FAUNA Paraguay Handbook of the Mammals of Paraguay Volume 2 Xenarthra.
See below for a key to the Paraguayan species.
FAMILY DASYPODIDAE - ARMADILLOS
General characteristics: Twelve species recorded in Paraguay. The back is rounded and armoured with a carapace covered with horny scales and with bony dermal scutes forming movable bands across the midsection. Both the forepart and the hind part of the carapace are immovable. The underside of the animal is hairy and lacks armour. Armadillos run with rapid movements of their short, strong legs. They have five toes on the hindfoot and three to five toes on the forefoot, each marked with strong claws. Forelimb structure is specialised for digging, exceptionally so in some species. Hindlimb structure is associated with weight bearing and measurement indices are correlated with body size. Most species are excellent diggers, constructing species-specific caves with a rounded roof in which they sleep and raise their young. Armadillos are generally solitary, approachable and tame, with poor vision but an excellent sense of smell. The Guaraní common name is most often used for the species in Paraguay, the Spanish names being understood but rarely used in everyday speech. The Guaraní word for armadillo, "tatu" is also used in vulgar speech in reference to female genitalia! The family is ancient, dating from the late Paleocene of South America.
Dental characteristics: The long mandibles are lined with a varying number of semicircular "molariform" teeth which grow continuously through life. These teeth lack enamel. Dental formula is variable and canines and incisors are lacking.
Skeletal characteristics: More fossorial species showing a greater length of the olecronon process of the ulna and more robust humerus.
Taxonomy: According to Möller-Krull et al (2007) the Dasypodidae are split into four subfamilies - Dasypodinae, Chlamyphorinae, Tolypeutinae and Euphractinae.
Subfamily Dasypodinae Gray, 1821
One genus and three species present in Paraguay. Characteristics are discussed under the generic account below.
Dasypus Linnaeus, 1758: Long-nosed Armadillos
There are seven species in this genus, two present in Paraguay. Synonyms adapted from Gardner (2007).
Synonyms:
Dasypus Linnaeus 1758:50. Type species Dasypus novemcinctus Linnaeus (1758) by tautonomy.
Tatus Fermin 1769:110. Name unavailable.
Tatu Frisch 1775:Table. Name unavailable.
Tatu Blumenbach 1779:94. Type species Tatu novemcinctus (=Dasypus novemcinctus Linnaeus 1758) by monotypy.
Cataphractus Storr 1780:40. Proposed for all armadillos known at the time.
Loricatus Desmarest 1804:28. In part. Type species Loricatus niger Desmarest (1804) (=Dasypus novemcinctus Linnaeus 1758).
Tatus Olfers 1818:220. In part. Incorrect spelling.
Tatusia Lesson 1827:309. In part. Type species Dasypus peba Desmarest (1822) (=Dasypus novemcinctus Linnaeus 1758).
Cachicamus McMurtie 1831:163. Type species Dasypus novemcinctus Linnaeus (1758).
Cachicama P.Gervais in I.Geoffroy St.-Hilaire 1835:53. Invalid emendation.
Zonoplites Gloger 1841:114. Name proposed for armadillos with four toes on forefeet and two middle toes longer than outer toes.
Praopus Burmeister 1854:295. Type species Dasypus longicaudis Wied-Neuwied (1826) by monotypy.
Cryptophractus Fitzinger 1856:123. Type species Cryptophractus pilosus Fitzinger (1856) by monotypy.
Hyperoambon W.Peters 1864:180. Type species Dasypus pentadactylus W.Peters (1864). Valid as a subgenus.
Muletia Gray 1874:244. Type species Dasypus septemcinctus Gray (1874) (=Loricatus hybridus Desmarest 1804, not Dasypus septemcinctus Linnaeus 1758) by monotypy.
Tatua W.Robinson & Lyon 1901:161. Incorrect spelling.
Mulletia Yepes 1928:506. Incorrect spelling.
Mulieta Talmage & Buchanan 1954:80. Incorrect spelling.
General characteristics: Small to large armadillos that are easily recognized by their long, naked, tubular ears, set close together on the head, long snout (55% or more of head length) and long, pointed tail (<50% of head and body length). The carapace is smooth, high and rounded, the denticles placed in irregular circles. The proximal two-thirds of the tail is ringed, each ring formed by two or more rows of scutes. There are four toes on the forefeet (digits 2 and 3 the longest) and five on the hindfeet (digit 3 the longest). Diet consists mainly of insects, but somewhat omnivorous, also taking small vertebrates and fruit. The earlies fossils date from the Pliocene of South America.
Cranial characteristics: Palate extends way beyond the last molar. Brain case is high and rounded, becoming thinner towards rear. Tympanic ring represents an otherwise unossified bulla.
Dental characteristics: Dental formula 7-9/7-9 = 28-36
Dasypus hybridus - Southern Long-nosed Armadillo
Dasypus novemcinctus - Nine-banded Armadillo
Subfamily Euphractinae Winge, 1823
General characteristics: Three species in Paraguay in two genera. These are small to medium-sized armadillos with a broad head, short snout and short ears. Conspicuous body hair on the carapace, sides and venter. The cephalic shield is broad and forms a ledge over the orbit. Scutes along the ventral margin of the carapace are rounded anteriorly and pointed posteriorly. The forefoot possesses four toes, claws on digits II and III being longest.
Cranial characteristics: Auditory bullae and external auditory meatus are ossified.
Chaetophractus Fitzinger, 1871: Hairy Armadillos
There are three species in this genus, two present in Paraguay. Synonyms adapted from Gardner (2007).
Synonyms:
Loricatus Desmarest 1804:28. In part.
Dasypus Illiger 1870:70. In part. Not Dasypus Linnaeus (1758).
Tatus Olfers 1816:220. In part. Incorrect spelling of Tatu Blumenbach (1779).
Tatusia Lesson 1827:309. In part.
Euphractus Burmeister 1861:427. In part. Not Euphractus Wagler (1830).
Chaetophractus Fitzinger 1871:268. Type species Dasypus villosus (=Loricatus villosus Desmarest 1804) by designation.
Dasyphractus Fitzinger 1871:264. Type species Dasyphractus brevirostris by monotypy.
Choetophractus Trouessart 1898:1146. Incorrect spelling.
General characteristics: Two species of small to medium-sized armadillos. Head is armed with a dermal shield that reaches to the nose. The medium-length ears are well-separated with rounded ends. The dorsal scutes are arranged in a linear fashion and covered in sparse, stiff hairs, with six to eight flexible bands across the centre. The legs and underparts are densely haired. Armadillos in this genus are omnivorous, with some suggestion of seasonal diet changes. The genus is known from the Pliocene of South America.
Cranial characteristics: Marked post-orbital constriction. Broad zygomatic arch. Brain case expanded and flattened dorsolaterally. Palatine terminates in a V-shape.
Dental characteristics: Dental formula 9/10 = 36
Chaetophractus vellerosus - Lesser Hairy Armadillo
Chaetophractus villosus - Greater Hairy Armadillo
Euphractus Wagler, 1830: Six-banded Armadillo
This is a monotypic genus. Synonyms adapted from Gardner (2007).
Synonyms:
Dasypus Linnaeus 1758:50. In part.
Loricatus Desmarest 1804:28. In part.
Tatus Olfers 1818:220. In part. Incorrect spelling.
Euphractus Wagler 1830:36. Type species Dasypus sexcinctus Linnaeus (1758) by designation.
Encoubertus McMurtie 1831:163. In part. Proposed as a subgenus of Dasypus Linnaeus (1758).
Pseudotroctes Gloger 1841:113. Type species Dasypus setosus Wied-Neuwied (1826) by monotypy.
Scleropleura Milne-Edwards 1871:178. Type species Scleropleura bruneti Milne-Edwards (1871) by monotypy.
Scelopleura Trouessart 1898:1141. Incorrect spelling.
General characteristics: One large species. The carapace is flattened and with a fine layer of stiff pale hairs protruding from between the dorsal bands. The head is triangular in shape and the ears are short. Forefeet with five digits, the third being the longest. Two to four openings for scent glands present in the mid-dorsum of the pelvic shield. An active burrower, digging for food and to escape enemies. Omnivorous, in addition to insects taking fruit, small vertebrates and even carrion. The genus is known from the mid-Pleistocene of South America.
Dental characteristics: Dental formula 9/10 = 36
Euphractus sexcinctus - Six-banded Armadillo
Subfamily Tolypeutinae Gray, 1865
General characteristics: This subfamily contains the most specialised fossorial armadillos and the most powerful diggers, though notably the genus Tolypeutes is non-fossorial. Three genera and four species present in Paraguay. The subfamily is split into two tribes, the Priodontini (Cabassous and Priodontes) and the Tolypeutini (Tolypeutes).
Tribe Priodontini Gray, 1873
General characteristics: Small to extremely large armadillos with a blunt, rounded snout, thick fleshy ears and relatively large eyes. The carapace is domed and separated from the cephalic plate by three rows of nuchal scutes. Scapular and pelvic shields are separate by 11 to 14 movable bands. Body hair is sparse. Five toes on the forefoot with digit III armed with a scythe-shaped claw.
Cranial characteristics: Frontal bones have a dome-like swelling. Only the tympanic ring of each bulla is ossified.
Cabassous McMurtrie, 1831: Naked-tailed Armadillos
Four species in this genus, three present in Paraguay. Synonyms adapted from Gardner (2007):
Synonyms:
Dasypus Linnaeus 1758:50. In part.
Tatus Olfers 1818:220. In part. Incorrect spelling of Tatu Blumenbach (1779).
Xenurus Wagler 1830:36. Type species Dasypus gymnurus Wied-Neuwied (1826) (=Xenurus squamicaudis Lund 1845) by monotypy. Preoccupied.
Tatusia Lesson 1827:309. In part.
Cabassous McMurtrie 1831:164. Type species Dasypus unicincttus Linnaeus (1758) by monotypy. Proposed as a subgenus of Dasypus Linnaeus (1758).
Arizostus Gloger 1841:114. Type species Dasypus gymnurus (=Tatus gymnurus Olfers 1818) by monotypy.
Tatoua Gray 1865:378. Type species Dasypus unicincttus Linnaeus (1758) by monotypy. Proposed as a subgenus of Dasypus Linnaeus (1758).
Ziphila Gray 1873:22. Type species Ziphilus lugubris Gray (1873) by monotypy.
Lysiurus Ameghino 1891:254. Proposed as replacement name for Xenurus Wagler (1830).
Cabassus Trouessart 1905:820. Incorrect spelling.
Cabassus Neveu-Lemaire & Grandidier 1911:103. Incorrect spelling.
General characteristics: Three species of medium-sized armadillo characterized by the fairly long, thin tail lacking well-developed plates (hence “naked-tailed”). The snout is very short and broad with a blunt end and the nostrils are guarded by a fine line of bristles. Ears funnel-shaped and capable of folding flat, closing the external auditory meatus. All feet possess five toes. Digits three, four and five of the foreefeet are broad and scythe-shaped and the claw on the first digit is long and slender. Hindfoot with blunt claws, the third being the longest. Carapace ovoid, dome-shaped and quite flexible with 11 to 14 movable bands. Dorsal plates are in transverse rows across the length of the body. These are nocturnal, solitary animals that give birth to a single young. The diet consists largely of ants and termites, these armadillos burrow frequently and often forage underground. The genus is known from the mid-Pleistocene of Brazil.
Cranial characteristics: Frontal bone with dome-like expansion. Rostrum short and tympanic ring present. Elongated infraorbital canal. Distance from lacrimal foramen and anterior opening of infraorbital foramen one-quarter to one-third rostral length. Mandible slender with articular process higher than coronoid process.
Dental characteristics: Dental formula 7-10/8-9 =30-38. Teeth peg-shaped and vary from circular to oviod in form on either axis.
Skeletal characteristics: 7 cervical vertebrae, 12 (or 12-13) thoracic, 3 (or 3-4) lumbar, 10 (or 9-11) sacral and 18 (or 15-20) caudal.
Cabassous chacoensis - Chaco Naked-tailed Armadillo
Cabassous tatouay - Greater Naked-tailed Armadillo
Cabassous unicinctus - Southern Naked-tailed Armadillo
Priodontes F. Cuvier, 1825: Giant Armadillo
This is a monotypic genus. Synonyms adapted from Gardner (2007).
Synonyms:
Dasypus Kerr 1792:112. In part. Not Linnaeus (1758).
Loricatus Desmarest 1804:28. In part.
Tatus Olfers 1818:220. In part. Incorrect spelling of Tatu Blumenbach (1779).
Priodontes F.Cuvier 1825:527. Type species Dasypus gigas G.Cuvier (1817) by monotypy.
Cheloniscus Wagler 1830:35. Type species Dasypus gigas G.Cuvier (1817) by monotypy.
Priodon McMurtrie 1831:164. Type species Dasypus gigas G.Cuvier (1817) by monotypy. Proposed as a subgenus of Dasypus Linnaeus (1758). Preoccupied by Priodon Quay & Gaimard (1824).
Polygomphius Gloger 1841:114. Type species Dasypus gigas G.Cuvier (1817) by monotypy.
Prionodon Gray 1843:xxvii. Nomen nudum.
Priodonta Gray 1843:xxvii. Incorrect spelling.
Prionodontes Schinz 1845:312. Incorrect spelling.
Prionodos Gray 1865:374. Introduced as a replacement for Priodontes F.Cuvier (1825).
General characteristics: A single species of huge size, much the largest member of its family. The snout is rounded and blunt-ended. Ears and eyes are large. Carapace is divided into bands of which the central 11 to 13 are flexible. The tail is covered with tightly-packed plates that are not arranged in rows. Feeds almost exclusively on ants and termites, destroying their nests with the long, sharply hooked claws of the forefeet (third digit being especially long). This species digs habitually, creating diagnostic large burrows which are occupied for a single night. Such burrows are often dug into termite mounds and a large number may be clumped into a single small area.
Dental characteristics: Highly variable with as many as 20 in a single tooth row, many of which are lost as the animal ages.
Priodontes maximus - Giant Armadillo
Tribe Tolypeutini Gray, 1865
Consists of a single genus of which one species is present in Paraguay. Characteristics are discussed under the generic account below.
Tolypeutes Illiger, 1811: Three-banded Armadillo
Two species, one present in Paraguay. Synonyms adapted from Gardner (2007)
Synonyms:
Tolypeutes Illiger 1811:111. Type species Dasypus tricinctus Linnaeus (1758) by designation (Yespes 1928).
Matacus Rafinesque 1815:57. Nomen nudum.
Tolypeutis Olfers 1818:221. Incorrect spelling.
Tatusia Lesson 1827:309. In part.
Apara McMurtrie1831:163. Type species Dasypus tricinctus Linnaeus (1758) by monotypy. Proposed as a subgenus of Dasypus Linnaeus (1758) .
Sphaerocormus Fitzinger 1871:376. Type species Tolypeutes conurus I. Geoffroy St.Hilaire (1847) by monotypy.
Cheloniscus Gray 1873:23. Type species Cheloniscus tricinctus Gray (1873) (=Dasypus tricinctus Linnaeus (1758) by monotypy. Preoccupied.
Tolypentes Matschie 1894:62. Incorrect spelling.
Tolypoïdes Grandidier & Neveu-Lemaire 1905:370. Type species Tolypoïdes bicinctus Grandidier & Neveu-Lemaire (1905) by monotypy.
Tolypuetes Talmage & Buchanan 1954:73. Incorrect spelling.
General characteristics: A single species of small armadillo possessing a uniquely short, stumpy tail covered in tubercles. The ears are fairly large and rounded and the carapace is high, rounded and almost without hairs. Pectoral and pelvic shields are convex and extremely hard separated by 2-4 (usually 3) movable bands. Head shield is is elongated, flattened and with rounded lateral and posterior margins. Body hair is conspicuous below the carapace. In the Paraguayan species there are 4 toes on the forefeet and five toes on the hindfeet with digits II to IV unified and bearing hoof-like claws. Feeds almost exclusively on ants, termites and soft-bodied invertebrates. When threatened it takes flight rather than burrowing as do other armadillos, characteristically rolling into a ball when captured. The earliest record of the genus is from the Pliocene of South America.
Tolypeutes matacus - Southern Three-banded Armadillo
Subfamily Chlamyphorinae Bonaparte 1850
General characteristics: This tribe contains the most fossorial of all armadillos. The ears are small and eyes tiny. Head shield is reduced to a thin flexible plate which is separated from the rounded rump plate. The sides of the body and ventral region is covered with long, thick whitish hair. Forefoot possesses five digits with disproportionately large claws - the species is adapted for "fast digging" in soft soils as opposed to "power digging" in hard substrates. One genus and one species present in Paraguay.
Calyptophractus Fitzinger, 1871: Fairy Armadillos
This is a monotypic genus. Synonyms adapted from Gardner (2007).
Synonyms:
Chlamyphorus Burmeister 1863:167. Not Chlamyphorus Harlan (1825).
Burmeisteria Gray 1865:381. Type species Burmeisteria retusa (=Chlamyphorus retusus Burmeister 1863) by monotypy. Preoccupied by Burmeisteria Salter (1865).
Calyptophractus Fitzinger 1871:388. Type species Chlamyphorus retusus Burmeister (1863) by monotypy.
General characteristics: A single, distinctive species of small armadillo. Head shield is broad and carapace is thin and flexible, attached along the sides of the body. The tail is extremely short, flattened dorsoventrally and with a pointed tip. The rump plate is rounded, abruptly flattened and acts as a “plug” when burrowing. The genus is known from the Pleistocene of South America.
Cranial characteristics: External auditory meatus is long, extending forwards above the zygomatic arch to the posterior border of the orbit.
Calyptophractus retusus - Chaco Fairy Armadillo
Key to Adults of the Paraguayan Species of Armadillos
1a Carapace complete on head and dorsum, with movable bands on latter. Total length >25cm.........................2
1b Carapace incomplete, reduced to small dorsal plates on head and along spine. Heavily furred. Total length tiny <16cm...........................................................................................................................Calyptophractus retusus
2a Forefeet with one or more greatly enlarged, scythe-like foreclaws....................................................................3
2b Forefeet lacking greatly enlarged, scythe-like foreclaws...................................................................................6
3a Enormous, total length >140cm, body length >70cm.........................................................Priodontes maximus
3b Total length <80cm, body length <50cm.........................................................................................................4
4a Three movable bands across dorsal surface. Curls into ball for defence - does not dig. Tail short, stumpy and covered in bony tubercles..........................................................................................................Tolypeutes matacus
4b >10 movable bands across dorsal surface. Digs rapidly to escape persecution. Tail long, thin and "naked" - lacking well-developed plates................................................................................................................Cabassous 5
5a Ears long and funnel-shaped extending above the head. Distributed in eastern Paraguay............................ 10
5b Ears short and rounded do not extend above the head. Fleshy expansion on anterior margin of ear. Distributed in the Chaco...........................................................................................................Cabassous chacoensis
6a Stiff dorsal hairs present on carapace. Ears well-separated at base. Snout short and blunt..............................8
6b Carapace steeply rounded and lacking dorsal hairs. Ears joined at base. Snout long and tubular........Dasypus 7
7a Total length >60cm. Tail equal to or greater than head and body length. 8 or 9 movable bands with distinct pale edges. Mean of 60 scutes along 4th movable band..........................................................Dasypus novemcinctus
7b Total length <50cm. Tail equal to or shorter than head and body length. 6 or 7 movable bands .......................................................................................................................................................Dasypus hybridus
8a Total length <40cm. Ears long, projecting well above head and extending to the second row of scapular scutes when laid back. Carapace somewhat rounded. Lacks holes in pelvic shield.............Chaetophractus vellerosus
8b Total length >40cm. Ears short and rounded. Carapace flattened dorsally. Holes in pelvic shield................9
9a Scutes typically dark reddish brown, guard hairs dark. 7 or 8 movable bands (never 6) between scapular and pelvic shields, at least 2 movable neck bands. Scutes of head shield small and disorganised .Chaetophractus villosus
9b Scutes typically pale yellow-brown, guard hairs pale. 6 to 8 movable bands between scapular and pelvic shields and one movable neck band. Scutes of head shield large and well-organised...............Euphractus sexcinctus
10a Large (<36cm Head body length) with shield colouration usually reddish. Mean number of cephalic scutes 48.3 (+/-3.7). Ear length >40mm. Number of scutes on 4th movable band 30.8 (+/-1.6). Usually in red soil areas throughout eastern Paraguay. ............................................................................................ Cabassous tatouay
10b Medium-sized (<35cm Head body length) with shield colouration usually dark brownish or blackish. Mean number of cephalic scutes 54 (+/-5.5). Ear length <30mm. Number of scutes on 4th movable band 27.4 (+/-1.3). Usually in sandy soil areas in the cerrado zone of north-eastern Paraguay. .................. Cabassous unicinctus
Online Resources
Edentate Specialist Group - Publishes the online journal Edentata dealing with armadillos, sloths and anteaters.
REFERENCES
Diaz MM & Barquez RM 2002 - Los Mamíferos de Jujuy, Argentina - LOLA, Buenos Aires
Emmons LH & Feer F 1999 - Mamíferos de los Bosques Húmedos de América Tropical - FAN Bolivia
Gardner AL 2007 - Mammals of South America Volume 1: Marsupials, Xenarthrans, Shrews and Bats - University of Chicago Press.
Möller-Krull M, Delsuc F, Churakov G, Marker C, Superina M, Brosius J, Douzery EJP, Schmitz J 2007 - Retroposed Elements and Their Flanking Regions Resolve the Evolutionary History of Xenarthran Mammals (Armadillos, Anteaters and Sloths) - Molecular Biology and Evolution 24: p2573-2582.
Redford K & Eisenberg JF 1992 - Mammals of the Neotropics Vol 2: The Southern Cone Chile, Argentina, Uruguay and Paraguay - University of Chicago Press.
Smith P. Owen R, Atkinson K, del Castillo H, Northcote-Smith E 2011 - First Records of Southern Naked-tailed Armadillo Cabassous unicinctus (Linnaeus, 1758) (Xenarthra: Dasypodidae) in Paraguay - Edentata 12.
Vizcaíno SF, Milne N 2002 - Structure and Function in Armadillo Limbs (Mammalia: Xenarthran: Dasypodidae) - Journal of Zoological Society of London 257: p117-127.
Wetzel RM 1980 - Revision of the Naked-tailed Armadillos Genus Cabassous McMurtie - Annals of the Carnegie Museum of Natural History 49: p323-357.
