SIX-BANDED ARMADILLO Euphractus sexcinctus FAUNA PARAGUAY

Euphractus sexcinctus (Linnaeus 1758) Image Gallery
TAX: Class Mammalia; Subclass Theria; Infraclass Eutheria; Order Cingulata; Family Dasypodidae; Subfamily Euphractinae (Myers et al 2006, Möller-Krull et al 2007). The genus Euphractus was defined by Wagler in 1830 and contains a single species. The genus name Euphractus was taken from the Greek meaning "true or good shell" and the species name sexcinctus means "six bands". Five subspecies were tentatively recognised by Gardner (2007), two of which are present in Paraguay - Euphractus sexcinctus flavimanus (Desmarest 1804) in the east and E.s.boliviae in the Chaco (O.Thomas 1907). Desmarest´s description of the subspecies E.s.flavimanus was based on de Azara´s (1801) "Le Tatou Poyou" and "L´´Encoubert" of Buffon (1763). Synonyms adapted from Gardner (2007):
Dasypus sexcinctus Linnaeus 1758:51. Type locality "America Meridionali" restricted to Pará, Brazil by O.Thomas (1907).
lor[icatus]. flavimanus Desmarest 1804:28. Type locality “Paraguay”, based on de Azara (1801) and Buffon (1763).
Dasypus flavipes G.Fischer 1814:122. Type locality “Paráguay”.
Dasypus gilvipes Illiger 1815:108. Nomen nudum.
Das[ypus]. gilvipes Lichtenstein 1818:215. Renaming of Dasypus gilvipes Illiger (1815).
T[atus]. gilvipes Olfers 1818:220. Type locality “Paraguay, Brasilien, Guiana”
Dasypus pilosus Olfers 1818:220. Nomen nudum.
Dasypus encoubert Desmarest 1822:370. Type locality “Le Paraguay”.
Tatus sexcinctus Schinz 1824:pl.113. Name combination.
D[asypus]. setosus Wied-Neuwied 1826:520. Type locality “in den Grossen Campos Geraes und den Angrän zenden Gegenden des Sertong” restricted to Bahía, Brazil by Ávila-Pires (1965).
Euphractus musetlinus Fitzinger 1871:259. In part. Type locality “Sud-und-Mittel-Amerika”.
Scleropleura bruneti Milne-Edwards 1872:1. Type locality “San Antonio … Ceará”, Brazil based on deformed specimen.
[Dasypus (Dasypus)] sexcinctus Trouessart 1898:1145. Name combination.
[Scleropleura] Bruneti Trouessart 1898:1141. Name combination.
[Dasypus] poyú Larrañaga 1923:243. Type locality implied as Uruguay. Based on Dasypus sexcinctus Gmelin (1788) (=Dasypus sexcinctus Linnaeus 1758) and de Azara (1802).
ENG: Six-banded Armadillo (Gardner 2007), Yellow Armadillo (Parera 2002), White-bristled Hairy Armadillo (Long 2003).
ESP: Armadillo de seis bandas (Neris et al 2002), Armadillo amarillo (Emmons 1999), Peludo (Emmons 1999), Gualacate (Parera 2002).
GUA: Tatu poju M (Villalba & Yanosky 2000), Poju AP (Villalba & Yanosky 2000), Krypurá Ac (Villalba & Yanosky 2000), Kry´y pura Ac (Esquivel 2001), Tatú poyú (Parera 2002), Tatú-podyu (Emmons 1999). "Poju" and its variations which feature in the Guaraní names refers to the needle-like claws of the forefeet.
DES: Predominately yellowish to reddish-brown in armour colour (usually yellowish), darker and somewhat blackish-brown on skin. The head plate is triangular with a straight posterior margin, coming to a blunt point just before the nose and is distinctly flattened on the upper part. It is relatively narrow, the width about 70-80% of its length. It extends as a spur behind the eye and there may be traces of scales in a semi-circle below the eye. Scales of the head plate are large and arranged in a well-defined pattern. Ears are medium-lenght, extending backwards to the second or third line of the scapular plate. They are well-separated by a distance greater than the ear-length across the top of the head. The carapace consists of two fixed plates, the scapular and pelvic plates, with 6 to 8 movable bands separating them. There is a single "nuchal" band between the head and scapular the plate. A sparse covering of long, stiff yellowish-tan hairs sprout from the skin between the bands. The tail is cylindrical and well-armoured with 2 to 4 bands of scales at the base. Two to four holes in the plates above the tail base are glandular openings responsible for the animals distinctive odour - this character is found only in Euphractus and Chaetophractus. The forefoot possesses five toes with robust claws, the third of which is the longest. Females have two pectoral nipples. Body temperature is 34ºC. CR: Broad and relatively long rostrum. Interorbital with a marked postorbital constriction. Brain case with obvious lateral markings, somewhat squarish and flattened. Zygomatic arch slender and elongate. Jugal never twice as high as the overlying anterior edge of the squamosal. Tympanic bulla present. External auditory meatus is ossified. Condylonasal Length: 114.5mm (109-125.5mm); Zygomatic Width: 68.6mm (61.7-75.4mm). (Diaz & Barquez 2002, Redford & Wetzel 1985). DF: Armadillos lack true teeth, but possess a series of "molariform" teeth that do not follow the standard mammal dental formula. They are particularly robust in this species. 9-10/10 = 38-40. First molariform located in the premaxillary as in Chaetophractus. There are nine pairs of maxillary teeth and 10 pairs of mandibular teeth in adults. CN: 2n=58, FN=102. (Gardner 2007).
TRA: The slightly rocking gait of this species leads to a distinctive print pattern in which the steps are almost perfectly aligned but the feet are slightly inward pointing towards the medial line. The tail is carried clear of the floor and does not usually leave an impression in the substrate. Despite the size of the animal the prints are surprisingly small, leaving somewhat rounded impressions, especially towards the tips of the digits. The hindfoot appears to have just three short digits and is similar in basic form to the forefoot, though slightly larger and with more rounded pads. FP: 2 x 1.7cm HP: 3 x 2.2cm. PA: 5cm. (Villalba & Yanosky 2000). Faeces typically with an irregular surface, measure 15 (+/-1.5mm) x 20mm (+/-1.3mm) and contain c62% soil and some plant material (18%). They are of fragile consistency and have a strong odour, presumably due to formic acid and decaying plant material. Weight 2.3g (+/-0.8mm). Typically found near areas of excavation and along trails (Anacleto 2007).
MMT: The largest of the "hairy armadillos" in Paraguay. TL: 61.64cm (40.1-95cm); HB: 39.57cm (34.1-49.5cm); TA: 22.02cm (11.9-30cm); FT: 8.35cm (7.5-9.2cm); EA: 3.52cm (2.4-4.7cm); WT: 4.32kg (2-6.5kg). Like other "hairy armadillos" they store fat and a captive female individual reached 11kg in weight. WN: 95-115g. (Parera 2002, Nowak 2001, Emmons 1999, Ceresoli et al 2003, Redford & Eisenberg 1992, Diaz & Barquez 2002, Redford & Wetzel 1985).
SSP: This is a large armadillo, second only in size to Priodontes maximus. Its covering of hair on the dorsum means that it is most likely to be confused with Chaetophractus villosus, C.vellerosus being considerably smaller. Note however that that species is generally darker and more reddish and hairier overall, especially ventrally, with conspicuous tufts of hair on the cheeks, legs and throat. Perversely the dark brown hairs of the dorsum of C.villosus may be harder to see against the dark carapace than the paler, yellowish-tan hairs of Euphractus. The number of bands is variable in this species and whilst C.villosus may possess 7 or 8 bands, it never has 6. Another useful character is the flattened head of this species, C.villosus shows a more rounded forehead and crown in profile. Behind the eye the head plate descends as a "spur" in Euphractus which may also show traces of a plate below the eye, both these features are absent in C.villosus which has a smooth and even edge to the plate behind the eye. Examination of the scales on the head plate shows a regular pattern of large scales in this species, that of C.villosus is a somewhat less-organised meleé of smaller scales, generally with a well-marked posterior "border". Beyond this border C.villosus shows at least two distinct dorsal bands prior to the scapular plate, only one is present in Euphractus. Note also that Chateophractus armadillos in Paraguay are confined to the Chaco and the cerrado belt of the northern Orient, whilst this species is less conservative in its habitat choice.
DIS: Widely distributed in eastern and central South America east of the Andes and south of Amazonia. It occurs from Pará, Brazil south through the caatinga and cerrado belt to Bolivia, Paraguay, Uruguay and northern Argentina (Misiones, Corrientes, Formosa, Chaco, Salta, Jujuy, Tucumán, northern Santa Fé and Santiago del Estero). There is an isolated population north of Amazonia in extreme southern Surinam and adjacent Brazil. In Paraguay the species occurs widely in both eastern Paraguay (E.s.flavimanus) and the Chaco (E.s.boliviae) and is absent only from extensively forested areas and urban zones. E.s.flavimanus is distributed through eastern Paraguay, Mato Grosso in Brazil, northeastern Argentina and Uruguay. E.s.boliviae occupies the Gran Chaco of Bolivia, western Paraguay and northern Argentina. The remaining subspecies are E.s.setosus in southeastern Brazil, E.s.tucumanus in the Argentinean Provincias of Tucumán and Catamarca, and E.s.sexcinctus occupying the northern part of the range and southern Surinam. The latter intergrades with flavimanus in Mato Grosso and setosus in southern Brazil. E.s.boliviae intergrades with tucumanus in the southernmost part of its range. The species was introduced into Central Chile, but does not appear to have become established (Long 2003).
HAB: Typical of open grassy and scrubby habitats, generally in dry areas, but also occurs in gallery forest in the Pantanal area and seasonally-flooded palm savanna in the Humid Chaco. In the Dry Chaco they are less common in densely-forested areas, preferring forest edge and scrub. In the cerrado belt they are found in campo limpio, campo sucio and sensu-strictu cerrado but are less frequent in cerradón - though Bonato et al (2008) reported that in the cerrado of Brazil they could find no clear preference between the patchwork of cerrado habitats and the species was equally common in grassland and forest. Habitat choice is apparently unaffected by fire, a study in the cerrado of Mato Grosso, Brazil finding that they utilised burnt areas as frequently as they do unburnt areas when foraging (Prada & Marinho-Filho 2004) - burning is of course a natural occurrence in the cerrado biome. Apparently less common at humid forest edge and probably absent from humid forest interior, but the expansion of the agricultural frontier is probably allowing them to colonise new areas. (Parera 2002, Nowak 2001, Emmons 1999, Neris et al 2002, Redford & Eisenberg 1992).
ALI: Omnivorous and able to exploit a wide variety of foodstuffs from fruits and plant matter to meat and even carrion. Eyesight is poor and smell is the primary method used for locating food. Six-banded Armadillos take large insects and exploit seasonally plentiful resources such as palm nuts and bromeliad fruits - at certain times of year plant material can compose a significant part of the diet. In Mato Grosso, Brazil, a study found the diet to consist of 90% plant matter, notably palm fruits (Acrocomia sp), pineapple (Ananas sp.) and figs (Ficus sp.), with the remaining 10% made up of beetles, crickets, ants, spiders, larvae, pupae and one amphibian. A study of stomachs of road kill individuals in São Paulo, Brazil found plant matter to make up just 33% of the diet, whereas insects (principally Formicidae and Scarabaeidae) made up more than 50% of the samples. Vertebrate remains included a Sigmodontid rodent, armadillo plates (likely from scavenging), pig skin, a snake and a small bird (Dalponte & Tavares-Filho 2004). One female roadkill individual from Goiás, Brazil had four Calomys sp. mice in its stomach, two of which were young (Bezerra et al 2001), while in the cerrado of Brazil Bonato (2002) found Oxymycterus sp., Clymomys sp., remains of a tanager (Thraupidae) and snakes amongst vertebrate remains. Bezerra et al (2001) suggested that predation on rodents during the dry season in the cerrado may be related to the scarcity of plant matter available at this time of year. Captive individuals have been seen to kill large rats, but their hunting technique was ineffectual and they were incapable of making a quick, clean kill (Redford & Eisenberg 1992). The flesh is removed by standing on the carcass and ripping off pieces with the mouth (Redford & Wetzel 1985). Captive individuals have also been observed attacking a Grey Brocker Deer fawn Mazama gouazoubira and a Rhea Rhea americana and attempting to drag them into their burrow (Dalponte & Tavares-Filho 2004). Though generally solitary, numbers may gather at large carcasses to feed on carrion and maggots (Nowak 1991). In common with other members of the subfamily Euphractinae they are able to store fat and this may be of assistance during seasonal food shortages (Redford & Wetzel 1985).
REP: Neris et al (2002) note the breeding season as "spring and summer" but that captive individuals have also engaged in breeding activities during autumn. Pregnant females have been found in central Brazil during September and October, and in Uruguay during January. Bonato et al (2008) considered breeding to be year round in the Brazilian cerrado whilst Cuéllar (in press) states that in the Bolivian Chaco there is a short concentrated breeding season at the end of the dry season with maximum fecundity in the first months of the wet season. Desbiez (2006) detailled chasing behaviour involving up to 8 individuals in a straight line and noted that local people in the Brazilian Pantanal consider this to be a form of mate competition, with several males chasing a female ready to breed. Such chases were seen to last over an hour and covering an area of no more than one hectare, the animals moving in an dout of denser vegetation. A female taken in Mato Grosso, Brazil was pregnant with two well-developed young in July. The gestation period is 60 to 64 days. Captive females have been seen to construct nests shortly before giving birth and typical litters contain one to three young and may include both sexes (Redford & Wetzel 1985). Meritt (1976 in Nowak 1991) states that twins frequently consist of one male and one female offspring. Newborns are poorly-developed and have a soft carapace. Females disturbed with their young may try to move or hide them and/or respond aggressively towards the intruder. Young displaced from the nest are returned by the mother. The eyes open after 22 to 25 days and by the end of the first month they have quadrupled in weight and are able to take solid food. Sexual maturity is reached at 9 months in captivity. (Redford & Eisenberg 1992, Parera 2002, Redford & Wetzel 1985).
BEH: General Behaviour Solitary and diurnal or nocturnal, tending to be nocturnal in areas where human population pressure is higher. Bonato et al (2008) reported that the species is nocturnal in the Brazilian cerrado, despite the lack of human predation in the area and suggested that this was because the species maintains body temperatures at a low temperature range. They are active, alert foragers and constantly on the move, maintaining a steady trot with a distinctly rocking motion, stopping occasionally to investigate potential sources of food. No difference in activity levels between seasons was noted by Bonato et al (2008) but there is some evidence to suggest that activity patterns are closely tied to rainfall and ambient temperature, perhaps because of their effect on food sources (Brooks 1995, Parera 2002). Adept diggers, they scratch the earth with the forefeet and uses the hindfeet to kick it clear of the burrow entrance (Nowak 1991). Caves typically have a single semi-circular entrance with the U-shaped roof corresponding to the convex dorsum of the animal being almost as high as they are wide - approximate dimensions being 20-22cm wide by 19-20cm high (Emmons 1999, Parera 2002). Carter & Encarnaçao (1983) found the mean dimensions of the burrow to be 21cm wide x 19 cm high at the entrance, narrowing to 20cm x 16cm 10cm inside the burrow, and the mean burrow angle to be 32.4º. Many burrows may be dug in a relatively small area and unlike many other armadillos burrows are frequently re-used, one male using a single burrow for 18 consecutive days (Carter & Encarnaçao 1983). In general burrows are only 1 or 2m deep, but open into a chamber that is wide enough for the animal to turn around (Nowak 1991). They are frequently constructed between tree roots (Parera 2002) and are dug in a direction so that the prevailing wind blows away from the entrance (Carter & Encarnaçao 1983). Minimum home range in Brazil was found to average 93.3ha (Nowak 1991, Parera 2002). Captive animals mark their cages with secretions from their pelvic scent gland and it would seems likely that burrows were marked by the same means in the wild state. (Emmons 1999, Redford & Eisenberg 2002). This species has a characteristic odour which is released from holes in the plates above the base of the tail (Nowak 1991). Defecation always takes place away from the burrow. A captive individual lived for 18 years and 10 months (Jones 1982). Aggressive Behaviour Female aggression is generally associated with lactation. Chases involving several individuals may have an aggressive function (Desbiez 2006). Defensive Behaviour Unlike other armadillos this species is reported to bite when handled (Emmons 1999, Redford & Eisenberg 2002) but despite handling numerous wild individuals of this species I have yet to encounter an individual which has attempted to bite when captured. Its main defence is to run for its burrow and if captured en route it will attempt to burrow rapidly into the ground even when held by the tail. Burrowing animals show remarkable strength and stubbornness and an animal which has begun to burrow can rarely be dislodged from its work unless it is in extremely soft soil. The animal will stop burrowing when pulling pressure is applied, digging in its feet and holding firm, but will immediately continue to burrow as soon as the pressure desists (P.Smith pers. obs.). Enemies Six-banded Armadillos figure in the diet of big cats such as Jaguar and Puma, but are probably also at risk from smaller cats and foxes. Juveniles may be taken by Lesser Grison which are small enough to enter burrows. Parasites Fujita et al (1995) reported the following nematodes from two specimens in Paraguay: Ascaris dasypodina, Aspidodera fasciata, A.esperanzae (described as a new specis in the same paper), A.scoleciformis, Cruzia tentaculata, Mazzia mazzia, Moeniggia complexus, Spirura guianensis, Trichohelix tuberculata and an unidentified species of Heterakinae. The two specimens were infected with 1504 specimens of six species and 97 specimens of four species respectively.
VOC: Generally quiet apart from the usual grunting noises produced by foraging armadillos. Juveniles are able to produce soft clicks and squeaks. (Redford & Wetzel 1985).
HUM: This species is frequently hunted for food by indigenous groups and rural populations on account of its large size and abundance (Cartés 2007, Emmons 1999). However it is not the preferred armadillo species for the table, in some areas having a reputation for being "unclean" on account of its habit of consuming carrion and it is even said to spread leprosy in some areas (Neris et al 2002). The meat is said to have a strong flavour (Redford 1994). Though it likely suffers from burning of grasslands to create pasture, the species may actually have benefited from agricultural activities opening up potential new areas for colonisation. However conflict arises when the crops are tubers such as manioc (mandioca) or sweet potato (batata), favoured foods for the armadillo, resulting in its persecution as a pest species. The presence of a large number of burrows in a small area creates a potential pitfall for horses and their riders (Nowak 1991). The fat of the animal may be used to cure respiratory illnesses and contusions (Neris et al 2002). In Argentina the tail may be used by indigenous groups to strike with flint to make sparks, as well as to carry fire-making tools (Redford & Wetzel 1985). In the Brazilian caatinga snuff is inhaled through the hollowed-out tail (Redford & Wetzel 1985).
CON: The Six-banded 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. This is generally one of the most commonly-encountered armadillo species in drier areas of the country and, despite hunting pressure in some areas, the population does not appear to be in decline. It occurs in a number of protected areas and its future seems secure. Biomass in the Brazilian Pantanal was estimated at18.8kg/km² about two-thirds of the overall armadillo biomass in the area (Redford & Wetzel 1985). In Mato Grosso, Brazil, the density was estimated in varying types of habitat with the following results 0.48/km² in cerrado, 2/km²in gallery forest, 0.59/km²in secondary forest and 2.9/km²in deciduous forest (Parera 2002). In the Brazilian cerrado the density was calculated at 0.14 individuals/ha (Bonato 2002, Bonato et al 2008) whereas in the Bolivian Chaco it was as low as 0.012 individuals/ha (Cuéllar in press). In São Paulo State this species made up 37% of all roadkills during a highway survey. However it does not appear to be unduly affected by burning within its habitat, Prada & Marinho-Filho (2004) considering direct mortality caused by fire to be negligible in the cerrado of Brazil.
Citable Reference: Smith P (2007) FAUNA Paraguay Online Handbook of Paraguayan Fauna Mammal Species Account 5 Euphractus sexcinctus.
Last Updated: 26 June 2009.
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Wied-Neuwied MP zu 1826 - Beiträage zue Naturgeschichte von Brasilien. Verzeichniss der Amphibien, Säugthiere un Vögel, welche auf einer Reise Zwischen dem 13ten dem 23sten Grade Südlicher Breite im Östlichen Brasilien Beobachtet Wurden II Abtheilung Mammalia, Säugthiere - Gr. HS priv. Landes Industrie Comptoirs, Weimar.

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