GREATER NAKED-TAILED ARMADILLO Cabassous tatouay FAUNA PARAGUAY

Cabassous tatouay (Desmarest 1804) Image Gallery
TAX: Class Mammalia; Subclass Theria; Infraclass Eutheria; Order Cingulata; Family Dasypodidae; Subfamily Tolypeutinae, Tribe Priodontini (Myers et al 2006). The genus Cabassous was defined by McMurtrie (1831) and contains four species, two of which are present in Paraguay. The genus was reviewed by Wetzel (1980). This species is monotypic (González 2001). The species name tatouay is an adaptation of the widely-used Tupi/Guaraní name Tatu ai. Desmarest (1804) described the species based on de Azara´s (1801) "Tatou Tatouay". Larrañaga´s (1923) description was based on de Azara´s (1802) "Tatuaí". Cabrera (1958) lists more synonyms, though these are based on misidentifications of other authors who considered C.tatouay a junior synonym of C.uninctus (Gardner 2007). Formerly placed in the Euphractinae, Möller-Krull et al (2007) provided DNA evidence that demonstrated their position within the Tolypeutinae. Synonyms adapted from Wetzel (1980) and Gardner (2007):
Loricatus tatouay Desmarest 1804:28. No type locality given, but restricted to Paraguay by Cabrera (1958) who considered the description to be based on de Azara (1801).
[Dasypus] dasycercus G.Fischer 1814:124. Type locality "Paraquaia".
[Dasypus] gynmurus Illiger 1815:108. Nomen nudum.
T[atus] gymnurus Olfers 1818:220. Type locality Paraguay. Based on de Azara (1801).
Tatusia tatouay Lesson 1827:311. Name combination.
Dasypus gymnurus Rengger 1830:290. Name combination.
Xenurus nudicaudis Lund 1839:231. Nomen nudum.
Dasypus 12-cinctus Burmeister 1854:282. Variant spelling of Dasypus duodecimcinctus Schreber (1774), wrongly applied.
Xenurus tatouay P.Gervais 1855:254. Name combination.
Xenurus unicinctus Gray 1865:254. In part. Not Dasypus unicinctus Linnaeus (1758).
Xenurus gymnurus Fitzinger 1871:242. Name combination.
Tatoua unicinctus Miller 1899:2. Not Dasypus unicinctus Linnaeus.
Xenurus duodecimcinctus Winge 1915:32. Not Dasypus duodecimcinctus Schreber (1774).
D[asypus] nudi-cauda Larrañaga 1923:343. Type locality "Provincia Paracuarensi". Based on de Azara (1802).
Cabassous tatouay Cabrera 1958:219. First use of current name.
Cabassous gymnurus Ximénez, Langguth & Praderi 1972:14. Name combination.
Cabassous duodecimcinctus Paula-Couto 1973:267. Name combination.
ENG: Greater Naked-tailed Armadillo (Redford & Eisenberg 1992), Spiny Armadillo (Cimardi 1996).
ESP: Tatu de rabo molle (González 2001), Armadillo de Cola Pelada Grande, Armadillo grande (Esquivel 2001).
GUA: Tatu ai (Emmons 1999, Esquivel 2001).
DES: Essentially similar to miniature versions of Giant Armadillo. Cabassous armadillos have a short, broad snout and small eyes. The cephalic shield has an average of 48.3 scutes +/-3.7. Dorsal plates are arranged in transverse rows along the body and there are 12-14 movable bands (mean 12.8 +/-0.6) - more than in any other Paraguayan armadillo. The third movable band has a mean of 31 scutes +/-1.7 and the fourth a mean of 30.8 scutes +/-1.6. There are two "nuchal" bands between the head and scapular plates and only isolated scales on the cheek. Scutes of the movable bands are poorly differentiated from the rest of the dorsal scutes and are of a similar size and shape. The scapular and pelvic shields extend almost to the base of the limbs. The first complete band of the scapular shield has a mean of 21.8 scutes +/-5.5 and the last a mean of 29 scutes +/-1.5. The first complete band of the pelvic shield has a mean of 29.1 scutes +/-1.4 and the last a mean of 8 scutes +/-1.3. Colour ranges from reddish-brown to blackish, but they are frequently stained by soils and they may be yellower laterally. There are no hairs present on the dorsal surface, though the lateral hairs may be fairly long. Ventrally greyish almost naked and only sparsely haired. The tail is fairly long and slender, armoured with thin, widely-spaced plates. This species is best identified by its long, funnel-shaped ears, well-separated from each other and extending well above the top of the head. Ears are scaled on the posterior side. Both feet have five pale claws, those of the forefeet being particularly long, especially the central (third) one which is greatly elongated and sickle-shaped. (Wetzel 1980). Core body temperature ranges from 32-34ºC (Zajic & Myers 2006). DF: Teeth are peg-like. Dental formula 9/8=34. There are no teeth present in the premaxillary bone. Maxillary teeth transversely compressed. Upper Tooth Row Length 38.3mm (+/-2.1); Lower Tooth Row Length 35mm (+/-1.9). Dimensions of Maxillary (Upper) Teeth (Length x Width): 4th=3.6 (+/-0.25) x 2.8 (+/-0.26), 5th=3.7 (+/-0.2) x 3.1 (+/-0.21), 6th=3.6 (+/-0.14) x 3.3 (+/-0.2), 7th=3.5 (+/-0.28) x 3.2 (+/-0.24); Dimensions of Mandibular (Lower) Teeth (Length x Width): 5th=3.8 (+/-0.29) x 3.2 (+/-0.21), 6th=3.7 (+/-0.17) x 3.2 (+/-0.25), 7th=3.3 (+/-0.2) x 2.8 (+/-0.32). (Wetzel 1980). CR: Mandible narrow and straight with condyloid process of slightly greater height than coronoid process. Tympanic rings present rather than bullae. Palate more elongate than other members of the genus with medial posterior margin extending posteriorly beyond a line between the anterior margins of the zygomatic rami of the squamosals. Ratio of Palatal length to Upper tooth row 1.81 +/-0.09. Ratio of Rostral Length to Postrostral Length 0.94 (+/-0.05). Condylonasal Length 109.9mm (+/-4.5); Rostral Length 53mm (+/-2.8); Palatal Length 68.2mm (+/-3.2); Postrostral Length 56.4mm (+/-2.4); Palatal Width 17.1mm (+/-1); Anterior Rostral Width 17mm (+/-1.6); Interlacrimal Width 44.4mm (+/-2.6); Interorbital Width 33.8mm (+/-1.2); Zygomatic Width 56.3mm (+/-2.8); Mastoidal Width 49.7mm (+/-3.2); Cranial Height 42.4mm (+/-2.2). (Wetzel 1980). CF: 2n=50, FN=71. Autosomal complement consists of 4 large metacentrics, 6 smaller metacentrics and submetacentrics and 14 teleocentrics. X is a small submetacentric and Y minute and likely acrocentric. (Gardner 2007).
TRA: This species walks supported by the claws of the forefeet while the entire sole of the hindfoot comes into contact with the substrate. The most obvious characteristic of the track of this species is the sickle-shaped central claw on the forefeet which is greatly elongated and curves inwards. Typically only the three longest claws leave impressions on both feet, the two central toes of the forefeet being notably longer and more curved than the others. The impression of the hindfoot usually shows the three narrow, tapering central toes which are not curved and have normal-sized claws. In soft soil the entire foot leaves an impression, the hindfoot being elongated with the innermost and outermost toes greatly reduced and set well-back from the longer central toes. Faeces are pelleted and composed of insect remains and soil (Zajic & Myers 2006).
MMT: The larger of the "naked-tailed armadillos" in Paraguay and indeed the largest member of the genus. TL: 63.68cm (56-80cm); HB: 45.78cm (36-49cm); TA: 17.9cm (15-20cm) González (2001) gives the tail measurements as 24-32cm, considerably longer than other published sources; FT: 8.22cm (8-8.6cm); EA: 4.17cm (4-4.4cm); WT: 5.35kg (3.4-6.4kg). The weight range of 7-12kg given in Gonzélez (2001) seems at odds with all other published sources. (Wetzel 1980, Redford & Eisenberg 1992, González 2001).
SSP: Cabassous armadillos are essentially smaller versions of Priodontes maximus, but size alone immediately rules out confusion with that species - no other armadillo even approaches Priodontes in size. They can be further identified by the lack of armoured plates on the tail (the tail is "naked") and the greatly enlarged claw on the forefeet. Highly-fossorial in behaviour these armadillos dig to escape danger - they do not try to run away. As a result of these distinctive features in Paraguay this species is only likely to be confused with the congeneric Cabassous chacoensis. That is a smaller species (approximately 50% smaller than this species) and crucially has much shorter ears with an unusual fleshy expansion on the anterior margins. The long funnel-shaped ears of this species extend well above the top of the head and back beyond the first complete scapular band when folded backwards - quite different in length and form to the extremely short ears of C.chacoensis. The two appear to be allopatric with this species confined to eastern Paraguay and C.chacoensis only recorded in the Chaco. Note that this species has consistently greater scale counts and cranial measurements than C.chacoensis for all standard measurements used in the description. The Southern Naked-tailed Armadillo Cabassous unicinctus squamicaudis has been hypothetically reported for Paraguay in the past, though no evidence of the species occurrence in the country has been forthcoming. It is intermediate in size between the two documented species, but closer to C.tatouay in general proportions. Externally the two species are best separated on measurements and scale counts (particulary of the cephalic shield) and a comparison table is presented below taken from Wetzel (1980):
Number of movable bands: C.tatouay 12.8 (+/-0.6) C.unicinctus squamicaudis 12 (+/-0.4).
Number of head scutes: C.tatouay 48.3 (+/-3.7) C.unicinctus squamicaudis 54 (+/-5.5).
Number of scutes on 3rd movable band: C.tatouay 31 (+/-1.7) C.unicinctus squamicaudis 28 (+/-1.3).
Number of scutes on 4th movable band: C.tatouay 30.8 (+/-1.6) C.unicinctus squamicaudis 27.4 (+/-1.3).
Number of scutes on 1st scapular band: C.tatouay 21.8 (+/-5.5) C.unicinctus squamicaudis 20.1 (+/-1.9).
Number of scutes on last scapular band: C.tatouay 29 (+/-1.5) C.unicinctus squamicaudis 26.3 (+/-1.7).
Number of scutes on 1st pelvic band: C.tatouay 29.1 (+/-1.4) C.unicinctus squamicaudis 24.4 (+/-1.6).
Number of scutes on last pelvic band: C.tatouay 8 (+/-1.3) C.unicinctus squamicaudis 6.6 (+/-1).
Head and Body Length (excluding tail) : C.tatouay >36cm C.unicinctus squamicaudis <35cm.
Ear Length: C.tatouay >40mm C.unicinctus squamicaudis <30mm.
Ratio of Palatal length to Upper tooth row: C.tatouay 1.81 (+/-0.09) C.unicinctus squamicaudis 1.62 (+/-0.05).
Ratio of Rostral Length to Postrostral Length: C.tatouay 0.94 (+/-0.05) C.unic. squamicaudis 0.86 (+/-0.06).
DIS: Occurs from south-central Brazil (Pará, northern Goiás and southern Bahía) south to Uruguay. It is apparently widespread in eastern Paraguay with records from all Departments. It appears to be absent from most of Ñeembucú Department where the habitat and climate are reminiscent of the Humid Chaco. In Argentina it is found in Corrientes and Misiones Provinces, and perhaps as far as south as Entre Rios Province in that country. In Uruguay it has been recorded in Lavalleja and Cerro Largo Departments, with unconfirmed reports from Rivera and Treinta y Tres. Its fossorial habits means that it is likely under-recorded.
HAB: Occurs in open, grassy habitats generally with loose, sandy soils that are ideal for digging, but its primary habitat appears to be forest. Esquivel (2001) notes that the species is found in Atlantic forest areas at the Mbaracayú Forest in Paraguay as well as in the extensive cerrado area. Their is little data available for the presence of the species in Atlantic Forest in Paraguay. In Uruguay it is found in grasslands with bushes and patches of forest (González 2001).
ALI: They feed largely on ants and termites which are extracted from their nests, either by burrowing into mounds from the surface, or by approaching from below via the tunnels. The long, extensile tongue is used to lick up prey and the sickle-shaped mid-claw of the forefeet is used to cut through small roots as the animal digs (Nowak 1991). In the process other invertebrates may be incidentally ingested, as well as soil. They possess a keen sense of smell which is used in the location of food (Zajic & Myers 2006).
REP: A single young is born (González 2001).
BEH: General Behaviour Almost nothing is known of the behaviour of this species other than the fact that it is highly fossorial and rarely observed. They are primarily nocturnal, with activity beginning after dark and are apparently solitary. At Serra da Canastra NP in Brazil Carter & Encarnaçao (1983) found the mean dimensions of the burrow to be 21cm wide x 15 cm high at the entrance, narrowing slightly to 21cm x 14cm 10cm inside the burrow, finding no significant difference in burrow width from those of Six-banded Armadillo but a significant difference in burrow height. The mean burrow angle was calculated to be 47.7º. Burrows are dug so that the prevailing wind blows away from the entrance. Holes have a single entrance and are used only once. On rare occasions they may spend more than 24 hours in a single burrow but burrows are never re-used. Carter & Encarnaçao (1983) only found burrows to be dug into active termite mounds. Defensive Behaviour Being essentially slow-moving animals their main form of defence is to dig rapidly into soft soil, and they are capable of completely burying themselves within a few seconds. However some may attempt to escape by running, and some individuals have been recorded to immerse themselves in water during pursuit (Grzimek 2003). The dorsal plate extends well down the side of the animal but would provide little defence against large predators.
VOC: Handled males may give a pig-like grunting noise, but females are generally silent.
HUM: This species was recorded amongst the prey items of the indigenous Aché in Mbaracayú Biosphere Reserve, though only 24 individuals were taken in the period 1980 to 1996, representing just 130kg of meat or 0.8% of the biomass that they consumed, suggesting that it is not one of their principal targets, or perhaps, given the ease with which it may be captured, reflecting the scarcity with which the species is encountered (Cartés 2007).
CON: The Greater Naked-tailed Armadillo is considered Lowest Risk, least concern by the IUCN, click here to see their latest assessment of the species. The Centro de Datos de Conservación in Paraguay consider the species to be persecuted by humans in Paraguay and give it the code N3. The species is not listed by CITES. Though widespread, the fossorial behaviour of this species means that it is rarely detected. It is able to tolerate some degree of habitat disturbance, but the transformation of large tracts of eastern Paraguay into agricultural land is not conducive to the continued presence of this species and it is presumably in decline in many areas in Departamentos Alto Paraná and Itapúa or else has suffered from regional extinctions. Of course the fossorial habits of this species are damaging to agriculture and it may be persecuted for that reason. Clearing of habitat by fire is also likely to take its toll, the species being ill-equipped to escape. It is hunted opportunistically for food along with other armadillo species, but hunting pressure is unlikely to be the major cause of population decline. It is present in several protected areas in Paraguay.
Citable Reference: Smith P (2008) FAUNA Paraguay Online Handbook of Paraguayan Fauna Mammal Species Account 13 Cabassous tatouay.
Last Updated: 30 January 2009.
References:
Azara F de 1801 - Essais sur l´Histoire Naturelle des Quadrupèdes de la Province du Paraguay - Charles Pougens, Paris.
Burmeister H 1854 Systematische Uebersicht der Thiere Brasiliens: Welche während einer Reise durch die Provinzen von Rio de Janeiro und Minas Geraës Gesammlt oder beobachtet wurden Vol 1 - G.Reimer, Berlin.
Cabrera A 1958 - Catálogo de los Mamíferos de América del Sur - Revista Museo Aregntino de Ciencias Naturales Bernadino Rivadavia Zoology 4: p1-307.
Carter TS, Encarnaçao CD 1983- Characteristics and Use of Burrows by Four Species of Armadillos in Brazil - Journal of Mammalogy 64: p103-108.
Cartés JL 2007 - Patrones de Uso de los Mamíferos del Paraguay: Importancia Sociocultural y Económica p167-186 in Biodiversidad del Paraguay: Una Aproximación a sus Realidades - Fundación Moises Bertoni, Asunción.
Cimardi AV 1996 - Mamíferos de Santa Catarina - FATMA, Florianópolis.
de Azara F 1801 - Essais sur l´Histoire Naturelle des Quadrupèdes de la Province du Paraguay Vol 2 - Pougens, Paris.
Desmarest AG 1804 - Tableau Méthodique des Mamifères in Nouveau Dictionnaire d´Histoire Naturelle Vol 24 - Chez Deterville, Paris.
Díaz MM, Barquez RM 2002 - Los Mamíferos del Jujuy, Argentina - LOLA, Buenos Aires.
Emmons LH 1999 - Mamíferos de los Bosques Húmedos de América Tropical - Editorial FAN, Santa Cruz.
Esquivel E 2001 - Mamíferos de la Reserva Natural del Bosque Mbaracayú, Paraguay - Fundación Moises Bertoni, Asunción.
Fischer G 1814 - Zoognosia Tabulis Synopticus Illustrata Vol 3 - NS Vsevolozsky, Moscow.
Fitzinger LJ 1871 - Die Natürliche Familie der Gürtelthiere (Dasypodes) - Abth. I. Sitz. Math. Natur. Kais. Akad. Wiss. Wien 64: p209-276.
Gardner AL 2007 - Mammals of South America Vol 1: Marsupials, Xenarthrans, Shrews and Bats - University of Chicago Press, Chicago.
Gervais P 1855 - Histoire Naturelle des Mammifères Vol 2 - L.Curmer, Paris.
González EM 2001 - Guía de Campo de los Mamíferos de Uruguay - Vida Silvestre, Montevideo.
Gray JE 1865 - Revision of the Genera and Species of Entomophagous Edentata, Founded on the Examination of the Specimens in the British Museum - Proceedings of Zoological Society oif London 1865: p359-386.
Grzimek B 2003 - Grzimek´s Animal Life Encyclopedia: Mammals - Thomson Gale, Germany.
Illiger K 1815 - Ueberblick der Säugthiere nach ihrer Vertheilung über die Welttheile - Abhandl. König. Akad. Wiss. Berlin 1804-1811: p39-159.
Larrañaga A 1923 - Escritos - Instituto Històrico y Geogràfico del Uruguay 2: p1-512.
Lesson RP 1827 - Manuel de Mammalogie ou Histoire Naturelle des Mammifères - Roret, Paris.
Lund PW 1839 - Blik paa Brasiliens Dyreverden för Sidste Jordomvaeltning - K. Danske Videnske. Selskabs Nat. Math. Afhandl. 8:p27-144.
McMurtrie H 1831 - The ANimal Kingodm Arranged in Conformity with its Organization by the Baron Cuvier ... The Crustacea, Arachnida and Insecta by PA Latreille .... - G & C & H Carvill, New York.
Miller GS 1899 - Notes on the Naked-tailed Armadillos - Proceedings of Biological Society of Washington 13: p1-8.
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.
Myers P, Espinosa R, Parr CS, Jones T, Hammond GS, Dewey A 2006 - The Animal Diversity Web (online). Accessed December 2007.
Neris N, Colman F, Ovelar E, Sukigara N, Ishii N 2002 - Guía de Mamíferos Medianos y Grandes del Paraguay: Distribución, Tendencia Poblacional y Utilización - SEAM, Asunción.
Nowak RM 1991 - Walker´s Mammals of the World 5th Ed Volume 1 - Johns Hopkins, Baltimore.
Olfers I 1818 - Bemerkungen zu Illiger´s Ueberblick der Säugthiere nach ihrer Vertheilung über die Welttheile p292-237 in Neue Bibliothek der Wichtigsten Reiseschreibungen... Vol 15 Heft 2 - Weimar.
Paula-Couto C 1973 - Edentados Fósseis de Sao Paulo - An Acad. Brasileira Ciênc. 45:p261-275.
Redford KH 1994 - The Edentates of the Cerrado - Edentata 1: p4-10.
Redford KH, Eisenberg JF 1992 - Mammals of the Neotropics: Volume 2 The Southern Cone - University of Chigaco Press, Chicago.
Rengger JR 1830 - Naturgeschichte der Saeugethiere von Paraguay - Schweighauserschen, Basel.
Schreber JCD von 1774 - Die Säugthiere in Abbildungen nach der Natur mit Beschreibungen - Wolfgang Wallther, Erlangen.
Wetzel RM 1980 - Revision of the Naked-tailed Armadillos Genus Cabassous McMurtrie - Annals of the Carnegie Museum of Natural History 49: p323-357.
Winge H 1915 - Jordfunde og Nulevende Gumlere (Edentata) fra Lagoa Santa, Minas Geraes, Brasilien - E. Museo Lundii, Copenhagen 3:p1-321.
Ximénez A, Langguth A, Praderi R 1972 - Lista Sistemática de los Mamìferos del Uruguay - Ann. Museo Nacional de Historia Natural de Montevideo Serie 2a 7:p1-49.
Zajic L, Myers P 2006 - Cabassous tatouay (On-line), Animal Diversity Web. Accessed July 24, 2008.

ACKNOWLEDGEMENTS
Many thanks to Mariella Superina for assisting with obtaining some of the references used in the construction of this species account.