NINE-BANDED ARMADILLO Dasypus novemcinctus FAUNA PARAGUAY

Dasypus novemcinctus Linnaeus 1758 Image Gallery
TAX: Class Mammalia; Subclass Theria; Infraclass Eutheria; Order Cingulata; Family Dasypodidae; Subfamily Dasypodinae (Myers et al 2006, Möller-Krull et al 2007). Seven species are recognised in this genus, three are present in Paraguay. Dasypus is derived from a Greek translation of the Aztec name "Azotochtli" which roughly means "tortoise-rabbit"; novemcinctus means "nine bands", in reference to the movable bands across the back. There are seven subspecies, the nominate subspecies D.n.novemcinctus Linnaeus 1758 is present in Paraguay (Gardner 2007). Examination of cranial characteristics is necessary to assign specimens to subspecies. Desmarest´s (1804) description was based on de Azara´s (1801) "Le Tatou Noir" and Buffon´s (1776) "Le Tatueté et Cachicame" Larrañaga´s (1923) description was based on de Azara´s (1802) "Negro". Synonyms adapted from McBee & Baker (1982) and Gardner (2007):
[Dasypus] novemcinctus Linnaeus 1758:51. Type locality "in America Meridionali" restricted to Pernambuco, Brazil by Cabrera (1958).
Tatus minor Fermin 1769:110. Unavailable.
Dasypus octocinctus Schreber 1774:pl.lxxiii. No locality given.
[Tatu] Novemcincta Blumenbach 1779:74. Name combination.
Dasypus longicaudatus Kerr 1792:112. Type locality "America".
Dasypus novemxinctus CW Peale & Palisot de Beuavois 1796:18. Incorrect spelling.
Dasypus longicaudatus Daudin in Lacépède 1802:173. Based on Buffon´s "Le Tatou á Longue Queue". Preoccupied.
lor[icatus] niger Desmarest 1804:28 Based on de Azara (1801) and Buffon (1776).
[Dasypus] serratus G.Fischer 1814:128. Type locality "Paraquaia, inprimis in provincia Buenos-Ayres (Boni Aëris)".
Dasypus decumanus Illiger 1815:108. Nomen nudum.
Dasypus decumanus Olfers 1818:219. Nomen nudum.
Dasypus niger Lichtenstein 1818:20. No type locality. Based on Dasypus novemcinctus Linnaeus (1758). Preoccupied.
Dasypus peba Desmarest 1822:360. Type localities "Le Brésil, le Guyane, le Paraguay... on ne trouve pas dans la Province de Buenos-Ayres"
Dasypus longicaudus Schinz 1824:253. Unavailable.
D[asypus]. longicaudus Wied-Neuwied 1826:531 Type Locality Rio Mucuri, Bahía, Brazil.
Tatusia peba Lesson 1827:311. Name combination.
Dasypus [(Cahcicamus)] novemcinctus McMurtrie 1831:163.
Dasypus uroceras Lund 1841:pl.12. Type locality "Rio das Velhas Floddal", Lagoa Santa, Minais Gerais, Brazil.
D[asypus] uroceros Burmeister 1848:199. Incorrect spelling.
Dasypus pepa Krauss 1862:19. Incorrect spelling.
D.[asypus]. longicaudatus W.Peters 1864:179. Incorrect spelling. Not D.longicaudatus Kerr (1792).
Hyperoambon peba W.Peters 1864:179. In part.
Dasypus fenestratus W.Peters 1864:180. Type locality "Costa Rica" restricted to San José, Costa Rica by Wetzel & Mondolfi (1979).
Dasypus Lundii Fitzinger 1871:340. Type locality "Brasilien".
Tatusia platycercus Hensel 1872:105. Type locality "Urwald von Rio Grande do Sul" Brazil.
Tatusia granadiana Gray 1873:14. Type locality "Concordia" Antioquia, Colombia.
Tatusia mexicana Gray 1873:14. Type locality Mexico.
Tatusia brevirostris Gray 1873:15. Type locality Rio de Janeiro, Brazil.
Tatusia leptorhynchus Gray 1873:15. Type locality Guatemala.
Tatusia boliviensis Gray 1873:16. Type locality Bolivia.
Tatusia leptocephala Gray 1873:16. Type locality "Brazils".
T[atusia]. leptorhinus Gray 1874:246. Incorrect spelling.
Tatusia novemcincta O.Thomas 1880:402. Name combination.
Tatusia longicaudatus JA Allen 1895:187. Name combination.
[Tatusia (Tatusia)] novem-cincta Trouessart 1898:1139. Name combination.
[Tatusia (Tatusia)] platycercus Trouessart 1898:1140. Name combination.
[Tatusia (Tatusia)] brevirostris Trouessart 1898:1140. Name combination.
[Tatusia (Tatusia)] leptocephala Trouessart 1898:1140. Name combination.
[Tatusia (Tatusia)] boliviensis Trouessart 1898:1140. Name combination.
[Tatusia (Tatusia)] granadiana Trouessart 1898:1140. Name combination.
[Tatus (Tatus)] novem-cinctus Trouessart 1905:814. Name combination.
[Tatus (Tatus)] platycercus Trouessart 1905:814. Name combination.
[Tatus (Tatus)] brevirostris Trouessart 1905:814. Name combination.
[Tatus (Tatus)] leptocephalus Trouessart 1905:814. Name combination.
[Tatus (Tatus)] boliviensis Trouessart 1905:814. Name combination.
[Tatus (Tatus)] granadiana Trouessart 1905:814. Name combination.
Tatua novemcincta W.Robinson & Lyon 1901:161. Name combination.
Tatusia novemcincta var. mexianae Hagman 1908:29. Type locality "Insel Mexiana" Pará, Brazil.
Dasypus boliviensis Grandidier & Neveu-Lemaire 1908:5. Type locality "Environs d´Uyuni" Potosi, Bolivia. Preoccupied.
D[asypus]. longi-cauda Larrañaga 1923:343. Type locality "Provincia Paracuarensi". Based on Azara (1802). Junior synonym and homonym of Wied-Neuwied (1826).
Dasypus mazzai Yepes 1933:226. Type locality "Tabacal, Departamento de Orán" Salta, Argentina.
D[asypus]. brevirostris Yepes 1933:230. Name combination.
ENG: Nine-banded Armadillo (Gardner 2007), Common Long-nosed Armadillo (Redford & Eisenberg 1992), Texas Armadillo (Long 2003).
ESP: Mulita grande (Redford & Eisenberg 1992, Diaz & Barquez 2002, Parera 2002), Armadillo mulita grande (Neris et al 2002), Tatu negro (Parera 2002). The Spanish name "Mulita" or "little mule" stems from the long, donkey-like ears present in members of this genus.
GUA: Tatu hu MPA (Neris et al 2002, Villalba & Yanosky 2000), Tatú-hu (Redford & Eisenberg 1992) Tatú-hú (Parera 2002), Tatu MPA (Villalba & Yanosky 2000), Chachú Ac (Villalba & Yanosky 2000). The Guaraní name is the most frequently utilised in Paraguay for this species, the Spanish names rarely being heard in everyday speech. Tatu hu means "black armadillo" in reference to the dark colouration of the carapace.
DES: "Long-nosed" Armadillos have a broad, depressed body, an obtusely-pointed rostrum, long, pointed ears (40-50% head length) and short legs. The carapace consists of two immobile plates, the scapular and pelvic shields each with 18-20 rows of ossified scutes and separated by 8 or 9 movable bands connected to each other by a fold of hairless skin. The vast majority of Paraguayan individuals possess 8 movable bands, despite the common name, and the average number of movable bands across the entire range is 8.3. The carapace is mostly blackish, hairless and with the scales of the anterior edge of the movable bands diagnostically pale yellowish-white in colour. Paler scales are also present on the lower halves of the scapular and pelvic plates, generally with broad blackish margins. Scutes on the movable bands are triangular in shape, but those on the main plates are rounded. The number of scutes present on the fourth movable band varies from 54 to 65, typically 60 (Díaz & Barquez 2002). The head is thin and triangular with a sloping forehead and long, mobile ears with rounded tips that are not separated by armour at the base. Head plate is generally pale and pinkish-yellow, often slightly darker and blacker on the crown. Scutes of the head plate are heavy and closely attached to the skull. The tail is long (equal to or greater than body length) with 12 to 15 rings of bicoloured scutes giving it a banded appearance along the basal 60% and narrowing towards the tip where it terminates in irregular scutes. There are four toes on the forefeet (characteristic of the Subfamily Dasypodinae), the middle two much the longest, and five on the hindfeet. The underside is naked and pinkish with only a light covering of coarse greyish hair sprouting from regularly-spaced papillae. Sexes are barely distinguishable, though males may be slightly hairier along the lateral part of the venter. Females have four nipples, two pectoral and two inguinal. Males lack a scrotum and the testes descend no further than the pelvis. Males have an average body temperature of 33.4ºC and females 31.3ºC, males also have a consistently faster heart rate. CR: Occipitonasal length 90mm (Diaz & Barquez 2002). Steeply descending frontal bone and an almost horizontal rostrum with a triangular tip in lateral profile. DF: Armadillos lack true teeth. "Long-nosed" armadillos have single-rooted, peg-like teeth that lack enamel. 7-9/7-9 = 28-36 - most often 8/8=32. CN: 2n=64, FN=80. Autosomes separable into five groups: two large pairs of metacentrics, four large pairs of acrocentrics, six pairs of medium metacentrics, fourteen pairs of medium acrocentrics and five pairs of small acrocentrics. The X chromosome is a large metacentric with a medial centromere. The Y chromosome is a small acrocentric.
TRA: Dasypus prints can be distinguished from those of other armadillos by their long, pointed toes with four toes on the forefoot and five on the hindfoot. However they generally leave the impression of only the two central toes on the forefeet (though sometimes the outer toe is also visible) and three central toes on the hindfeet. Given a full print, the hindfoot has a pointed heel with three long, somewhat pointed central toes and two, much shorter, outer toes set well back towards the heel. The forefoot has the inner toe much reduced and it rarely leaves an impression. FP: 4.8 x 3.5cm HP: 6 x 4.8cm. PA: 25cm. (Villalba & Yanosky 2000). Faeces resemble those of Tamandua tetradactyla, being composed largely of ant and termite remains, but they are generally more elongated and also contain soil and elements of vegetable matter. Typically no more than 2 or 3 pellets are dropped in one place. Faeces are usually deposited along runways in forested areas, making them difficult to locate (Taber 1945, Anacleto 2007).
MMT: The largest of the "long-nosed armadillos" in Paraguay. Males slightly larger than females. TL: 64.57 (60-100cm - generally in the range 60-85cm) Adult size is not reached until the animal is 3 to 4 years old (Redford & Eisenberg 1992); HB: 41.03cm (32.4-57.3cm); Height at Shoulder 155-230mm; TA: 30.18cm (21.1-45cm); FT: 87.5cm (5.9-11.4cm); EA: 4.32cm (3.3-5.9cm); WT: 3.91kg (2-9.83kg - generally between 3-6.5kg) Carapace accounts for c16% of body weight; WN: 85g. (Parera 2002, Nowak 1991, Emmons 1999, Ceresoli et al 2003, Redford & Eisenberg 1992, Diaz & Barquez 2002, McBee & Baker 1982). Hind foot and ear lengths can be used to determine fetal age (Rojas-Suárez & Maffei 2003).
50 days HF 7mm; EA 2mm; WT 2g.
70 days HF 13mm; EA 9mm; WT 16g.
90 days HF 20mm; EA 14mm; WT 36g.
120 days HF 27mm; EA 20mm; WT 64g.
SSP: This is much the largest of the "long-nosed armadillos" in Paraguay and can be immediately identified by the number of bands - generally eight or nine (usually 8) in this species and never more than seven in other Paraguayan Dasypus (Hamlett 1939). It can be further distinguished by the blackish colouration with yellowish-white triangular scales on the posterior edge of the movable bands. Of all the Dasypus this species has proportionately the longest ears (40-50% of head length) and tail (equal to or greater than body length). Dasypus hybridus is generally much paler in colouration, somewhat yellowish with proportionately shorter ears and 6 or 7 movable bands. The ears are shorter, typically 25-30% of head length, as is the tail (67-70% body length). It has a preference for open grassy areas and the two species are not usually found together. Using the fourth movable band as a standard, Hamlett (1939) noted that this species has a mean of 60 scutes whereas D.hybridus has a mean of 54 scutes. Dasypus septemcinctus is dark and blackish, lacking the pale edges to the movable bands which also number 6 or 7. It has similar length ears (40-50% head length) but a proportionately shorter tail (80-100% body length) than the Nine-banded. It has a mean of 48.4 scutes along the fourth movable band (Hamlett 1939). Both D.septemcinctus and D.hybridus have just 6 upper teeth in each jaw. Footprints are approximately 50% larger than those of D.septemcinctus.
DIS: This is the most widely distributed of all the armadillos, occurring from the USA south to Argentina, and is actually in the process of expanding its range to the north. It was recorded in Texas for the first time in 1880 and now extends east to Florida and north to central Oklahoma though cold its slowing its northward advance (MacDonald 1985). It is found throughout Central America and even some Caribbean Islands. West of the Andes it occurs south to northern Peru, but east of the Andes it reaches Uruguay and Argentina as far south as Provincia Entre Rios in the east of the country, though only to the Yungas in the west. In Paraguay it occurs throughout the country in all departments, though it is considerably more sparsely distributed in arid areas. Parera (2002) states that its presence in the Dry Chaco of Argentina requires confirmation, but Neris et al (2002) report multiple records of the species in the Dry Chaco of Paraguay - though they note that the species is more common in humid areas. The northern and southerly limits of the range are probably defined by cold, lack of insects and drought (Redford & Eisenberg 1992) whilst MacDonald (1985) also notes that competition with other armadillo species may also limit its southerly expansion. The subspecies found in Paraguay is D.n.novemcinctus, which is distributed through most of South America, east of the Andes from Colombia and Venezuela south to Argentina. The remaining subspecies are: D.n.mexicanus from the USA south to southern Mexico; D.n.davisi from the Balsas Basin to the mountains of Morelos in Mexico; D.n.hoplites from Trinidad and Tobago, Grenada, Barro Colorado Island and Costa Rica; D.n.aequatorialis west of the Andes in Ecuador and probably also Peru and Colombia; D.n.fenestratus from Oaxaca through Central America to Panama and possibly also across northern South America; D.n.mexianae is endemic to the delta of the Rio Amazonas, Pará, Brazil. The species has been successfully introduced to the Pacific island of Guam some time after 1959 (Long 2003).
HAB: This adaptable species is able to colonise a variety of habitats from humid Atlantic forest, through grasslands and agricultural areas, to the palm savanna of the Humid Chaco and drier chaco forest and cerradón. It is more common in warm, humid areas and seems to prefer some forest cover, being the most frequently encountered armadillo species in undisturbed humid forest habitats (ECOSARA Biodiversity Database). It reaches its greatest abundance in the Humid Chaco but is generally absent from open, grassy areas where other species of Dasypus are present.
ALI: Considered a generalist insectivore that exploits the food materials most easily available in any given habitat. Feeds largely on ants and termites which are plentiful in its Paraguayan range. Anacleto (2007) studied the diet via faecal samples during the dry season in the cerrado of Mato Grosso, Brazil and found Coleoptera to be the biggest single item in the diet by weight (71.3%) followed by ants (16.5%) and termites (12.2%) of the genera (in order of prominence as biomass) Nasutitermes, Velocitermes, Rhynchotermes, Cornitermes and Anhangatermes. The nests of the two most prominent termite species in this study were those that are the most easily opened, suggesting that the species excavates when foraging for termites. In the USA it also supplements the diet with carrion, fruit and small vertebrates, but these are less prominent in the diet of South American specimens. Parera (2002) also lists amphibians, reptiles, birds eggs and juvenile micro-mammals in the diet of Argentinean individuals and to a lesser extent fruit, roots and seeds. An analysis of stomach contents estimated the 93.3% of foods taken were animal in nature (McBee & Baker 1982). They begin to forage immediately on leaving the burrow after nightfall and studies suggest that there is a move from more forested to more open habitats as the night progresses (McBee & Baker 1982). Foraging animals are noisy and follow random paths, snorting constantly and investigating every potential source of food. Eyesight is poor and food is located by smell, with most food items taken from the soil surface or by using the claws to dig shallow, triangular feeding holes just below the surface. (Redford & Eisenberg 1992). Frequently they will rise up onto the hindlegs, using the tail for balance, and sniff the air. The nose is often pressed into the hole to maintain the scent trail and they may hold their breath for up to 6 minutes to avoid dust inhalation (MacDonald 1985). Nine-banded Armadillos re-use traditional feeding trails and foraging individuals are so preoccupied with the job in-hand that they may even collide with an immobile observer (Emmons 1999). Capable swimmers they may enter water to feed, using the forefeet to search for aquatic invertebrates in the substrate (Parera 2002). Adults have been seen using the forefeet to smash bird´s eggs and to hold down rabbit carcasses whilst ripping off pieces of flesh with the mouth (McBee & Baker 1982). Armadillos must drink water frequently and do so by lapping in the manner of a dog, leaving a film of their thick saliva over the water surface. Taber (1945) fed captive individuals on bone and fishmeal mixed with sour milk and mash. They also learned how to eat dead chicken, using the foreclaws to scratch meat from the bones and tugging with the mouth to loosen it. When the carcasses became infested with maggots the maggots were preferred to the chicken. Chicken eggs were eaten only when broken first. Watermelon and cantelopue were taken only when broken and only when other food was unavailable. Cereals were ignored, even when soaked in fish oil, but bread soaked in fish oil was accepted. Beets, potatoes, onions and turnips were never consumed despite being offered for over a month.
REP: Females are receptive once a year, but males produce sperm all year-round and the main breeding season is from August to November (Neris et al 2002). Parera (2002) states that in fact the females ovulate twice a year and that mating occurs in the summer-autumn season. In Izogog, Departamento Cordillera, Bolivia births were found to be highly seasonal and concentrated from October to December with just a single litter per year (Rojas-Suárez & Maffei 2003). Pairs form for a single breeding season, at which time they may share a burrow. Males are sometimes polygynous, but females always take a single mate. Because of the position of the genitalia mating occurs with the female on her back. (Mc Bee & Baker 1982). In common with the other "long-nosed armadillos" this species produces multiple, same sex offspring from a single egg - essentially clones of each other. Rojas-Suárez & Maffei (2003) found a greater proportion of males to females in 44 litters in Bolivia (males 26:14 females), and the proportion was sustained in their sampling of the adult population. Four young are almost always produced in this species, limited by the number of nipples available on the mother, though Parera (2002) notes that exceptionally litters of 2 to 8 have been recorded. In 300 examined litters there were only four cases of triplets, four cases of quintuplets and one set of twins (McBee & Baker 1982). Because of the polyembryonic breeding strategy there is almost always an even number of offspring. The gestation period is 8 to 9 months, a remarkably long pregnancy explainable by a 3 to 4 month embryonic diapause (delayed implantation). At 5 to 7 days the ovum forms a blastocyst and passes into the uterus where development ceases, becoming implanted several months later. Neris et al (2002) quotes the gestation period at 120 days, presumably not including the diapause. Newborns have soft, leathery skin which takes several weeks to harden and the eyes are open from the first day. They are able to walk within a few hours and join their mother on foraging expeditions after a couple of weeks (McBee & Baker 1982). Wild individuals are weaned in three months but the family may remain together for nine months. Females reach sexual maturity and ovulate for the first time at two years, males by the end of their first year. In captivity females have been known to bear young 13 to 24 months after capture and isolation from males. (Redford & Eisenberg 1992).
BEH: Activity Solitary and largely nocturnal, though occasionally active at dusk and during the day (P.Smith pers.obs.). Activity levels are likely determined by ambiental factors, particularly temperature and they may become more diurnal during the reproductive season (Parera 2002). Peaks of activity usually occur around 21-22-00h. They are not well-adapted to cold conditions and are able to lower the body temperature by only around 2.5ºC, resulting in periods of inactivity during cold spells. Rain does not affect activity unless accompanied by cold weather (Taber 1945). The lifespan is 12 to 15 years. Nesting Typically this species spends the day in self-dug burrows 0.5-6m long (mean 1.25m) which are usually located in forested areas close to a water source. Any one animal may have a mean of 4.5 to 8.5 burrows depending on locality (Taber 1945). Burrows are dug by wedging the nose and forefeet into the soil to loosen it, then scratching with the forefeet to position it below the abdomen. Using the forefeet and tail for balance, the hindfeet are brought forward in front of the pile of dirt, the back is arched and then suddenly thrusted straight so that the loose soil is propelled behind the animal. (McBee & Baker 1982). Burrows generally have multiple entrances 17-20cm in diameter and contain a grass nesting chamber c34cm in diameter which is used for reproduction, sleeping and refuge during colder periods. Average vertical depth to the nest chamber is 50cm, but an extreme of 151cm has been recorded. Few studies have investigated burrow orientation in this species, but what little data exists appears to suggest a random orientation, unlike those of savanna species which typically have the entrance to the burrow sheltered from the prevailing winds. It has been hypothesised that their preference for forested environments provides the necessary protection from prevailing winds, liberating them from constraints affecting burrow orientation (Platt et al 2004). Plant material is carried between the chin and forelegs, the animal zig-zagging backwards, at which time the tail may serve a tactile function. On arrival at the burrow the armadillo enters backwards, raking the plant material into the hole with the nose and feet. (McBee & Baker 1982). In Arkansas Taulman (1994) observed somewhat different nest-building behaviour. He noted that collecting material involved 3 to 5 scrapes lasting about 3 to 5 seconds and amassed a bundle of leaf litter c20cm in diameter. The armadillo made several collecting trips, each in different directions, and return trips followed the same direct path as the outward journey, with the animal hopping backwards to its burrow. Between collecting bouts (approximately once a minute) the animal paused to sniff the air before continuing with its task. The animal only turned to face the burrow upon arrival, when it used the forefeet and snout to push material into the hole. The material from the last trip (the fifth) was simply dumped at the burrow entrance, the hole apparently beingcompletely full with material, and the armadillo then walked away. Taber (1945) noted that nests of this species have no structure and rather are piles of leaves into which the animal buries itself. As a result occupied burrows often appear to have the entrance plugged with plant material, a fact that may serve a purpose for camouflage as well as providing insulation to prevent over-heating or heat loss. (Taulman 1994). Several individuals may use the same burrow, but these are likely members of the same family, though usually all of the same sex. Burrows are frequently used by other species. In areas that are subject to periodic flooding (eg Humid Chaco) nests are frequently built above ground to avoid drowning and resemble miniature haystacks (Nowak 1991). Range size and territorial behaviour Home ranges of males often overlap considerably without antagonism and may encompass the ranges of several females. The size of the range varies depending on the carrying capacity of the habitat, with figures of 3.4ha suggested for prime habitat and 15ha for suboptimal habitats having been suggested (Redford & Eisenberg 1992, Nowak 1991), with male ranges up to 50% larger than those of females (MacDonald 1985). This species possesses glands on the ears, eye-lids, anus and soles of the feet which play an advertising role warning of the presence of the individual in the area. Upon meeting they frequently sniff the anal regions and to further mark territories they defecate and urinate on prominent posts and along feeding trails (MacDonald 1985). The yellowish glandular secretions may also be used to deter predators, producing a nauseating effect if inhaled in quantity. Animals have been seen to feed in close proximity to each other without signs of aggression, though sick or injured animals have been reported to victims of mutilation or even cannibalism by conspecifics (Mc Bee & Baker 1982). Swimming and Bathing Nine-banded Armadillos are capable of swimming short distances by gasping to inflate the stomach and intestine with air for buoyancy - upon first entering the water they are almost completely submerged, but ride higher on the water surface the longer they swim (McBee & Baker 1982). Swimming armadillos fold the ears back against the head, kick all four legs in doggy fashion and periodically raise the nose to breathe before lowering it again (Taber 1945). Though by no means a common occurrence, the ability to hold the breath and accrue a considerable oxygen debt means they can also cross streams and small rivers by walking along the river bed (MacDonald 1985) - in fact they may even feed in the water (Parera 2002). Taulman (1994) described an individual seen bathing in a large puddle 15-28cm deep. The animal dipped its snout into the water and raised the head quickly so that water dripped backwards over the carapace, before systematically shaking in a front to back sequence lasting 1 to 2s. The animal then rolled onto its back and writhed briefly as if rubbing mud onto its dorsal surface. Each rolling sequence lasted approximately 4 seconds and it was repeated three times before the animal trotted away. Aggressive Behaviour Aggression is commonly observed during the breeding season. Male aggression likely assures exclusivity of mates, female aggression may be a result of competition for limited resources or to stimulate dispersal of the previous year´s young (McDonough 1994). Adults have been observed chasing juveniles at high speed for a period of up to 35 minutes, presumably to displace them (Breece & Dusi 1985). Fights are rare but generally take the form of one individual leaning back on the hind feet and clawing with the forefeet or clawing at the sides of another individual with the hind feet whilst rolling and flipping (McDonough 1994). Defensive Behaviour Surprised animals frequently explode into a run, crashing through the undergrowth towards their nest burrow. One animal at Itabó Itaipú Reserve, Departamento Alto Paraná was observed to run and collide with a tree when disturbed, stopping momentarily before continuing on (K.Atkinson pers.comm.). Animals frightened at close range may perform remarkable vertical jumps with arched back prior to fleeing or when pursued (Emmons 1999) and if overtaken may attempt to curl up to protect as much of the body as possible (Nowak 1991). When pursued into a burrow they may arch the back, digging the scales of the carapace into the tunnel roof and make it impossible to pull them out (McBee & Baker 1982). Enemies In Belize this species constitutes up to 50% of the diet of the Jaguar and remains have also been found in the nests of Harpy Eagles. Juveniles are likely taken by Lesser Grison, foxes, small cats and Black Tegu. Parasites Relatively few parasites have been documented for this species despite its extensive range. All ticks reported on armadillos are in the genus Amblyomma, with the species A.concolor and A.pseudoconcolor documented on this species. Fleas are rarely found but the following species have been reported: Echidnophaga gallinacea, Juxtapulex echidnophagoides, Polygenis roberti, P.occidentalis and Tunga travassosi. It is also known as a carrier of the protozoan Trypanosoma cruzi which is responsible for Chaga´s Disease, probably becoming infected by eating insect carriers. It is also susceptible to infection by Schistosoma haematobium but develops no symptoms and the eggs and parasites do not reach the urogenital system. Helminths (Ascarops sp., Aspidodera fasciata, Brachylaemus virginianus, Hamanniella sp., Mazzia mazzia, Oncicola canis, Oochoristica sp., Physocephalus sp., Travassosia carinii). Strongyloidea (Dasypostrongylus filamentosus, Delicata ransomi, D.uncinata, D.cameroni, Maciela macieli, M.flagellata, Moennigia moennigi, Pintonema intrusa, P.pulchra, P.pseudopulchra, P.pintoi, Pulchrostrongylus complexus). Fujita et al (1995) reported the following nematodes from two specimens in Paraguay: Ancylostoma sp., Aspidodera esperanzae (described as a new species in the same paper), Moeniggia complexus, M.pintoi and an unidentified species of Heterakinae. The two specimens were infected with 69 specimens of four species and 3 specimens of one species respectively. Navone (1990) recorded the nematode Aspidodera vazi (Aspidoderidae) and the cestode Mathevotaenia surinamensis (Anoplocephalidae) in the Paranaense and Chaco regions of Argentina, with Aspidodera fasciata (Aspidoderidae) additionally recorded in the Chaco region.
VOC: Though it does not generally vocalise, the Nine-banded Armadillo is one of the noisiest forest inhabitants given its snorting foraging style and clumsy form of ambulation with complete disregard for the noise created by snapping branches, dry leaves and moving undergrowth. A medium-sized animal charging through the undergrowth of humid forest at high speed at night is likely to be this species (Emmons 1999).
HUM: This species is one of the principal prey items of the Aché in Mbaracayú Biosphere Reserve who prize it for its white meat (Esquivel 2001). In the period 1980 to 1996 a total of 1500 individuals of this species were taken by the Aché in the reserve, representing 5,750kg of meat or 35.2% of the biomass that they consumed (Cartés 2007). The species has become more prominent in the diet of the Aché since 1994, when it represented 43.1% of the wild game in their diet compared to just 13.5% in 1980, a change likely related to deforestation and the reduction in numbers of larger prey species such as White-lipped Peccary which declined from 22% to 2% of the diet over the same period (Cartés 2007). In other areas Nine-banded Armadillos is one of the preferred wild game food items over much of its range with the meat being considered a substitute for chicken (Neris et al 2002) and the species is customarily cooked in its "shell" (McBee & Baker 1982). A campesino community in Departamento Caazapá took 66 individuals of this species over a four year period, making it the third most frequently hunted species and representing 21% of all animals taken, though this corresponded to only 5% of the total biomass (Cartés 2007). Fat is used as a remedy for the treatment of coughs, bronchitis, asthma, rheumatism, open wounds, appendicitis and concussion (Cartés 2007, Neris et al 2002), making it the second most utilised mammal species for medicinal purposes in Paraguay after the Capybara. The carapace may be used to fashion wallets as a cheap substitute for leather and the species is often taken as a pet (Neris et al 2002). With the reduction in populations of other myrmecophagous species, Nine-banded Armadillos probably play an important role in population control of ants and termites. Though easily tamed they do not adapt well to captivity (Nowak 1991). They have been falsely accused of attacking domestic poultry in some areas of their range (McBee & Baker 1982). Recently the species has been used in laboratory studies of in birth defects, multiple births, organ transplants and investigations of diseases such as leprosy, trichinosis and typhus (Nowak 1991). There are no documented cases of leprosy being passed from an armadillo to a human (McBee & Baker 1982). The species is considered a minor agricultural pest in the southern USA on account of its burrowing activities which affect fruit and agricultural plantations and apparently affect dykes, fences and undermine buildings (Long 2003).

CON: The Nine-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 secure in Paraguay but with the potential to become threatened in the future and give it the code N4. The species is not listed by CITES. The species is able to coexist with humans in rural areas because of its largely nocturnal habits and large litter size, but has declined over much of its range in eastern Paraguay as a result of destruction of optimal habitat. It remains most numerous in the Humid Chaco were large unpopulated areas of relatively undisturbed habitat allow the species to proliferate. The species is extremely susceptible to leprosy both under captive conditions and in the wild (Nowak 1991). Population density has been estimated at 0.36/km² in the cerrado of Brazil (suboptimal) to 50/km² in the coastal prairies of Texas and may be even higher in tropical forest areas (MacDonald 1985).
Citable Reference: Smith P (2007) FAUNA Paraguay Online Handbook of Paraguayan Fauna Mammal Species Account 8 Dasypus novemcinctus.
Last Updated: 26 June 2009.
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ACKNOWLEDGEMENTS
Many thanks to Mariella Superina for assisting with obtaining some of the references used in the construction of this species account.