Nine banded armadillo

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Nine banded armadillo
Nine banded armadillo

Nine banded armadillo

Systematics
Order : Armored siderails (Cingulata)
without rank: Armadillos (Dasypoda)
Family : Dasypodidae
Subfamily : Dasypodinae
Genre : Long-nosed armadillos ( Dasypus )
Type : Nine banded armadillo
Scientific name
Dasypus novemcinctus
Linnaeus , 1758

The nine-banded armadillo ( Dasypus novemcinctus ) is a species of mammal belonging to the group of armadillos (Dasypoda). It is one of the best known and most widespread representatives of this group, and it is also the best researched. The occurrence of this armadillo is not limited to large parts of South America , as the only species still alive today it is also widespread in North America , where it was first observed in the 1850s. The habitats inhabited by the nine-banded armadillo are very diverse and mainly include forests and partly open landscapes, which must, however, be more or less well moisturized; It is also considered to be extremely adaptable, but it cannot cope with longer cold spells. The animals live solitary and dig underground burrows. They mate once a year. The usually four newborns usually represent four genetically identical young animals and remain with the mother for up to nine months. The armadillo is also very adaptable in terms of nutrition and eats mainly insects , but also smaller vertebrates . Due to the wide distribution, up to seven subspecies are distinguished. In relation to humans, the fact that the nine-banded armadillo carries pathogens of leprosy and Chagas disease and is so often used in medicine as a laboratory animal is significant . In some countries in its range, it is also considered a food resource. The nine-banded armadillo is not classified as endangered.

features

Habitus

Nine-banded armadillo, the armored head is clearly visible

The nine-banded armadillo reaches a head-trunk length of 32 to 57 cm, the tail is 21 to 45 cm long and is therefore usually about as long as the trunk or sometimes longer. The weight varies from 3 to a maximum of 9.8 kg, although 6.5 kg is rarely exceeded; Overall, males are slightly heavier than females. The armadillo has up to 10 cm long and narrow, triangular shaped head with a long, extended snout. The ears are up to 5.9 cm in length compared to the head, large, very mobile and have rounded ends. The typical head shield consists of large bone shields that lie close to the skeletal substructure. Like all armadillos, the nine-banded armadillo is characterized by its back armor, which comprises three separate sections: a fixed shoulder and pelvis section and a section consisting of 8 to 10, but usually 9, movable ligaments in between, which are connected by folds of skin. The solid armor parts each consist of about 18 to 20 rows of small, rounded bone platelets. However, due to the covering of the bone platelets with small horn shields, they do not always appear symmetrically arranged. The bone plates of the movable ligaments, on the other hand, are square shaped. They have a triangular pattern with a point pointing backwards. The fourth ligament has 54 to 65 such bony formations. In addition, the first two thirds of the long tail are surrounded by 12 to 15 rings of bones. The carapace is mostly brown in color, with the lower parts being a bit lighter, as are the moving bands in front. The head shield is also tinted lighter, while the ears are very dark in color. The sparsely hairy belly has a pink tint. The legs are short, the front feet have four toes and the rear feet have five toes, all of which have sharp claws. Those of the two middle forefoot toes are the longest. The rear foot reaches an average length of 8.7 cm. Females have four mammary glands . The animal also has a strongly fluctuating core body temperature of 30 to 36 ° C, with an average of 33.4 ° C in males than in females with 31.7 ° C.

Skull and skeletal features

skeleton

The skull is up to 9.9 cm long and a good 4.3 cm wide at the zygomatic arches . The bony part of the snout , the rostrum, is about 5.9 cm long, making it significantly longer and also narrow. The lower jaw can be 7.8 cm long. The structure of teeth differs from that of other mammals and does not follow the common dentition formula . The teeth are simply built and without tooth enamel , have only one root and have a pin-like shape. Both in the upper and in the lower jaw there are 7 to 9 teeth per half of the jaw, i.e. a total of 28 to 36, but on average there are 8 per arch, i.e. a total of 32. However, there seem to be clear differences within the individual subspecies of the nine-banded armadillo since only 7 teeth per mandibular arch were observed in populations from Nicaragua . The total length of the lower row of teeth is 2.6, that of the upper 2.4 cm. On the forelimbs, the ulna has a particularly large upper joint ( olecranon ) that, with a total bone length of 6.9 cm, reaches about 2.7 cm. Such large joint ends on the forelimbs are typical of animals with a burrowing way of life.

Sensory lines and vocalizations

The nine-banded armadillo has a rather poor sense of sight , but an extremely good sense of smell , which it mainly uses when foraging for food. Several loud calls are known. Digging and struggling animals emit a panting grunt, fearful a squeak. Low-frequency contact calls have also been detected during the mating season. In addition, the armadillo is a very noisy animal, which emits a constant snort or grunt when looking for food. Otherwise it also causes a lot of noise by breaking dry branches or rustling in the leaves and is considered to be one of the most noisy forest dwellers.

distribution

Entire distribution area

General

The distribution area of ​​the nine-banded armadillo extends from North to Central to South America and thus has the largest extension of all armadillo species. As the only representative of the armadillos, it can also be found in the USA , where it mainly inhabits the south and southeast. The southern limit of distribution is in northern Argentina and Uruguay . In western South America, the Andes plateaus represent a natural border, especially in Peru and Bolivia . In addition, individual populations can also be found on the Caribbean islands , namely Grenada and Trinidad and Tobago . The entire inhabited area covers 19.1 million square kilometers, but the actual occupied area is unknown.

The nine-banded armadillo lives from sea level up to heights of around 2000 m. It is very adaptable and tolerates numerous habitats . The armadillo is just as native in moist forests as it is in open grasslands, on agricultural land and in urban areas. In general, there is a preference for warm and humid climates with only a small proportion of frost days in the year. Preferably inhabited landscapes are riparian areas and wet forests, especially the Atlantic coastal forests ( Mata Atlântica ), but also swamps. However, it is particularly lacking in the very dry areas of the Gran Chaco with less than 380 mm of annual precipitation, which are inhabited by other armadillo species. This may also be the reason why the nine-banded armadillo is not found further south. The population density of the individual regions is very different. In the Cerrado region, for example, there is only one individual per three square kilometers, but in the coastal areas of Texas there are up to 50 animals per square kilometer. In protected areas of the Atlantic coastal forests, up to 24 individuals can occupy the same area.

Spread to North America

Distribution area in the USA with current distribution (red) with the representation of different distribution limits at different times, the red dots are new observations

The spread to North America, particularly the US, which has occurred more recently and is still ongoing, has been well studied. The first animals were observed in southernmost Texas on the Rio Grande in 1854, and the armadillo had already reached the Nueces River in the southeast of the US state in 1880. The westward expansion stopped around 1905 at the Pecos River in New Mexico , currently only isolated animals can be found west of the river. To the east, the nine-banded armadillo first appeared in Arkansas in 1921 . As early as 1919, several animals were released by marines in Florida , at the same time some specimens escaped from a zoo, both groups established the population in this state. In the further course of the eastern expansion of the range, the nine-banded armadillo appeared for the first time in Louisiana in 1925 , crossed the Mississippi in 1934, arrived in Alabama in 1939 and in Georgia in 1942 . Furthermore, the species expanded to the north, the first occurrence of the species is reported in 1936 in Oklahoma , 1943 in Kansas , 1947 in Missouri , 1966 in Colorado , 1970 in Nebraska , 1974 in South Carolina and finally in 1992 in Illinois . Currently, the distribution covers the entire southeastern United States, in the west the expansion is largely completed by the dry desert areas with violent onset of winter, but in the north it continues and proceeds at an average speed of 7.8 km per year in the Great Plains it is even 11 km, on the Gulf Coast it reached up to 17.2 km per year. However, expansion to the north is slowed down by the cold winters, as the nine-banded armadillo is hardly resistant to the cold. Another limiting factor is the less abundant food supply there in the cold season.

Why the nine-banded armadillo appeared so late in North America is unknown, as it has been found in Mexico much earlier, for several thousand years, and there are no climatic differences to the southern part of North America or geographic barriers. Possibly it has to do with the size and density of the population in Mexico or with overcoming a physiological inhibiting factor. The latter could only be explained with an evolutionary further development of the nine-banded armadillo.

Way of life

Territorial behavior

Nine-banded armadillo in Oklahoma

The nine-banded armadillo is the best studied and documented armadillo species, so much is known about its way of life. The animal lives solitary and is mostly nocturnal, but it is also sometimes visible during the day. Since it is less adapted to the cold, the activities in cooler phases are lower, and the body temperature is lowered by up to 2.5 ° C. The representatives of the armadillo species are not territorial, but they use action spaces ( home ranges ), in which they prefer to stay and whose size increases with the age of the individual. In optimal habitats they are usually between 1.2 and 3.4 hectares in size, in less suitable landscapes they can be up to 15 hectares. Differences between male and female animals were not observed. The action spaces of various individual animals, both males and females, can overlap without causing aggression among one another. Only in the rutting season are the areas of the males strictly separated. When individuals come into contact with one another, the genitals are usually sniffed first. During the daily foraging for food, a single animal covers up to 1.2 km, but the average distances are 80 to 180 m. In addition, the nine-banded armadillo is able to cross bodies of water, where it simply wanders through shallow, narrow bodies of water at the bottom, but crosses larger bodies of water by swimming; to do this, it pumps air into its digestive tract. Occasionally one sees a single animal actively bathing in a small body of water.

Entrance to the construction of a nine-banded armadillo

The nine-banded armadillo spends the day mostly in its own dug burrows, which it creates with its strong forefeet. On average, an animal has between 4 and 8 burrows in its action area. These are 0.5 to 6 m long and are mostly located in forests with well-moistened soils and near bodies of water and mostly on slopes. They reach about 50 cm obliquely into the subsoil, in extreme cases they are up to 150 cm deep. Each burrow has several entrances, 17 to 20 cm in diameter, which are sometimes covered with plants. The entrances are not oriented in a certain direction, in contrast to the burrows of armadillo species that live in the open country and therefore create them protected from the wind. The long corridor ends in a larger chamber around 34 cm in diameter, often padded with dried grass or leaves. The nine-banded armadillo fetched the material for the nest cushion in several phases from the vicinity. The temperature inside the building is often 3 to 4 ° cooler than outside. Sometimes you can find several animals in one den, but they usually belong to the same family and are always of the same sex. In some cases, the nine-banded armadillo shares its burrow with other species such as the common black iguana or skunks and opossums . In regions that are regularly flooded, the armadillo builds nests made of plant material on the ground that are about 20 by 34 cm in size and 22 cm high and have only one entrance.

nutrition

Nine-banded armadillo in Wolf River, Memphis, Tennessee
Erected nine-banded armadillo

The nine-banded armadillo is primarily a carnivore with a specialization in insects , preferring to eat beetles . Studies from North America show a proportion of 30 to 45% of beetles and their larvae in the diet, here mainly scarab beetles , ground beetles and click beetles . In addition, hymenoptera , grasshoppers and butterflies are represented with proportions of 4 to 15% each, and small vertebrates such as amphibians and reptiles were also consumed. The armadillo also occasionally plunders the egg clutches of the diamond turtle and the nests of the Virginia quail . It has also been observed that it eats meat from dead rabbits or birds , such as the American Robin. Fruits and berries, including those from ebony plants , play a rather subordinate role, but regionally plant-based food can also account for up to 10% of the nutritional requirement. Similar studies from the Cerrado region of Brazil showed a frequency of only 1.1% in beetles, which made up 71% of the biomass eaten. Ants were ingested more frequently with 32.6%, but the biomass proportion here was only 16.5%. However, the highest amount was consumed by termites with 66.3%, which, however, only consumed 12.2% as pure biomass. Members of the termite genera Velocitermes and Nasutitermes were eaten in particularly large numbers . Also in the Serra da Capivara National Park in northern Brazil, examinations of excrement residues and stomach contents have shown predominantly beetles, ants and termites, but also remains of iguanas . Various analyzed stomach contents in both the South and North American range contained snakes as food residues. After observations in the Santa Rosa National Park in Costa Rica , a nine-banded armadillo hunted a poisonous coral snake by jumping the back of the animal on the animal. It took about 30 minutes to completely eat the 1 m long snake. Overall, the nine-banded armadillo shows itself to be very opportunistic in its eating behavior.

The nine-banded armadillo uses 77 to 90% of its wakefulness to search for food. It paces back and forth nimbly, sniffs at all potential eating spots and gives off a constant snort of itself. An animal tends to use frequent paths and often moves from deeper forests to more open land. During the search, it drills numerous holes; in a marsh pine ecosystem in Georgia, an average of 267 holes 9 cm deep per hectare were found. Often it can also be observed that the animal sits upright, propping itself up with its tail, and sniffs in the air. In order to follow a scent trail, it holds its nose very close to the ground while searching for food. In addition, breathing stops for up to six minutes so that no dust is inhaled. When foraging for food, the nine-banded armadillo is so concentrated that it sometimes runs into obstacles.

Reproduction

Males are sexually mature at around twelve months, females at 18. Mating only takes place once a year, the rutting in females lasts a good four days. Due to the widespread distribution of the nine-banded armadillo, the mating season is very different from region to region and depends on the seasons . In North America it takes place mainly from June to November, in Bolivia from October to December and in Paraguay from August to November. It is only during this time that males and females form pairs. There are also dominance fights between the advertising males, some of which stand on their hind feet and are fought with the claws of the front feet, but also when an animal kicks backwards. Due to the anatomy of the shell, fertilization takes place with the female lying on her back.

The gestation period lasts about 8 to 9 months, the actual development in the womb only takes about 140 days on average, however, due to a diapause after fertilization and storage of the egg in the uterus , the embryo is delayed in development by up to 4 months. This means that the offspring can be born in a climatically favorable time. In cases of stress , however, the egg cell can be delayed implantation, which means that the birth does not take place until 32 months after the last fertilization. It was also observed that females gave birth twice in a row without any contact with males, which is also due to the delayed implantation.

Usually four young are born, but there have also been reports of two or eight offspring. Due to polyembryony , all young animals are genetically identical and the number of offspring is almost always an even number, and triplets were only rarely observed. A cub weighs around 85 to 100 g. It already resembles adult animals, but the carapace and claws are softer and only harden in the course of the first few weeks of life. Furthermore, the eyes are open and a newborn can walk after just a few hours. The suckling phase lasts about three months in total and the young animals initially gain about 11 g per day. They leave the nest for the first time after around 20 days and go looking for food with the mother, where they also drink water for the first time. However, the first insects are only eaten after around 70 days. After weaning, the family group stays together for about six more months until the young have grown up and leave the mother. Life expectancy is twelve to fifteen years.

The result of polyembryony is that the genetic diversity of a group in the immediate vicinity of a mother's nest is relatively low, which studies on several hundred individuals in the Yazoo National Wildlife Reserve in the US state of Mississippi have shown. The genetic variability only increases with increasing distance from the original nest. This is related to the brain drain after weaning. The increasing diversity in genetic structure is higher in females than in males. The scientists involved in the study offered two explanatory models for this phenomenon: on the one hand, this could lead to more frequent migration of the females and greater local loyalty ( philopathy ) of the males, on the other hand, the different mating successes of the males in polygynous reproduction could also be reflected here . An exclusive migration of female individuals from other areas of the range has so far only rarely been genetically proven.

Predator and enemy behavior

Jaguars are one of the greatest predators

The nine-banded armadillo usually flees into the next burrow when danger approaches and blocks the entrance like a cork with its pelvic armor; it also arches its back and gets caught in the surrounding earth with the movable straps, so that it can hardly be pulled out of its burrow. When an animal runs, it is very quick and has reportedly been able to outrun hunting dogs . In addition, the nine-banded armadillo produces a secretion that causes nausea in potential predators. An attentive animal sniffs while standing on its hind legs, while a frightened one jumps into the air with all fours and bends its armor. The most important predators are the jaguar and the puma , but the pressure that both predators exert on the populations varies from region to region. In Belize was in 54% of 183 surveyed Kotresten the jaguar, the nine-banded armadillo be shown in the Mata Atlantica -Küstenwäldern of southeastern Brazil, there were still 15%. In the Brazilian state of São Paulo , 8.3% of the feces from the puma and 3.7% from the jaguar consisted of remains of the armadillo. In North America, however, it hardly seems to belong to the prey spectrum of the puma, just as little to that of the coyote , since no evidence could be provided in 566 examined faeces. In some cases the ocelot has also been reported as a predator. Young animals can also fall victim to the harpy or the black and white tegu .

Parasites

Despite the wide distribution of the nine-banded armadillo, only a few parasites have been identified to infect it. Common external ones include ticks of the genus Amblyomma . Fleas are among other Tunga - and polygenic species represented. Internal parasites include predominantly nematodes , such as Aspidodera and Moeniggia . The armadillo is particularly important as a carrier of Mycobacterium leprae , which can also cause leprosy in humans. However, the level of the transmission risk is unclear. In Louisiana , where the disease has been endemic since the mid-19th century, the nine-banded armadillo didn't emerge until the 1920s. Furthermore, leprosy is much more widespread in the USA alone than the armadillo species as a whole, but a 2011 study was able to prove the transmission path via the animal species. The disease itself only became indigenous with the colonization of America by the Europeans and did not occur before that. Likewise, Trypanosoma cruzi demonstrated in nine-banded armadillo. This single cell causes Chagas disease , which is common in South America .

Systematics

Internal systematics of the armadillos according to Gibb et al. 2015
  Dasypoda  
  Dasypodidae  

 Dasypus kappleri


   


 Dasypus septemcinctus


   

 Dasypus hybridus



   


 Dasypus mazzai


   

 Dasypus sabanicola



   

 Dasypus novemcinctus


   

 Dasypus pilosus






  Chlamyphoridae  
  Euphractinae  

 Euphractus


   

 Chaetophractus


   

 Zaedyus




   
  Chlamyphorinae  

 Chlamyphorus


   

 Calyptophractus



  Tolypeutinae  

 Priodontes


   

 Tolypeutes


   

 Cabassous







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The nine-banded armadillo is one of seven species that today form the genus of the long-nosed armadillos ( Dasypus ). The long-nosed armadillos , in turn, belong to the group of armadillos (Dasypoda) and form a family of their own , the Dasypodidae . This also includes numerous extinct genera, such as Stegotherium and Propraopus . The former is largely known from the Miocene and comprised several species, while the latter, on the other hand, comes from the Pleistocene and also appeared with several species. According to molecular genetic studies, the Dasypodidae separated from the line of other armadillos as early as the Middle Eocene around 45 million years ago. Today these are placed in the family of the Chlamyphoridae , which includes all other recent genera.

Together with the seven -banded armadillo ( Dasypus septemcinctus ), the southern seven -banded armadillo ( Dasypus hybridus ), the savannah ( Dasypus sabanicola ) and the Yungas armadillo ( Dasypus mazzai ), the nine-banded armadillo forms a closely related group, that of the Dasypus subgenus listened to. According to molecular genetic studies from 2015, it is again closely related to the Yungas and savanna armadillos. The Kappler armadillo ( Dasypus kappleri ), on the other hand , is referred to its own subgenus, Hyperoambon , as is the fur armadillo ( Dasypus pilosus ) in Cryptophractus . Another result of the more recent molecular genetic studies is that a greater diversification of the Dasypus subgenus began about 5 million years ago, which corresponds to the transition from the Miocene to the Pliocene .

A total of seven subspecies are distinguished today:

  • D. n. Aequatorialis Lönnberg , 1913; South America west of the Andes
  • D. n. Davisi Russell , 1953; Central America from the Río Balsas to Morelos
  • D. n. Fenestratus Peters , 1864; Central America from Oaxaca to Panama
  • D. n. Hoplites Allen , 1911; Caribbean Islands and Costa Rica
  • D. n. Mexianae Hagmann , 1908; South America in the mouth of the Amazon
  • D. n. Mexicanus Peters , 1864; North America and Central America to southern Mexico
  • D. n. Novemcinctus Linnaeus , 1758; South America east of the Andes to Argentina

According to studies, the populations in North America have comparatively little genetic variability, in contrast to those in South America. This is explained by only a small group of founders that reached North America around 160 years ago. Genetic analyzes of around 150 animals from Mexico showed that they belong to two different clades that probably already differentiated themselves in the Upper Pleistocene in South America. The two groups are distributed more in the eastern and in the western part of the country with the Sierra Madre Oriental as a possible natural barrier. As a result, North America was probably reached twice independently of different populations of the nine-banded armadillo.

It is possible that a population from French Guiana represents an independent species, as it differs more strongly according to the more recent genetic studies from 2015 and probably split off from the line that led to today's nine-banded armadillos about 3 million years ago would have. This is also confirmed by anatomical studies of the skull from 2018. These also showed that the nine-banded armadillo can be divided into three to four morphological types in its rest of the area of ​​distribution. According to the authors of the analysis, these types could form their own taxonomic units, which, however, do not correspond to the previously defined subspecies.

Linnaeus , 1775

Few fossil finds of the nine banded armadillo are known, the species is almost entirely restricted to the Holocene . From the Medford Cave in Florida come individual remains in the form of bone platelets and limb remains, which, according to genetic studies, can be regarded as the remains of the nine-banded armadillo. Since the age of the finds is dated to about 10,000 to 12,000 years and thus belongs to the end of the Pleistocene, the presence of this species is very early in North America. There are also only a few fossils from South America, mostly bone shields, from the Gruta dos Moura and Gruta do Urso caves in the Brazilian state of Tocantins and from the Gruta do Urso Fóssil in the Ubajara National Park in the Brazilian state of Ceará . The temporal range of the finds includes the end of the Pleistocene and the early Holocene. The majority of the fossil finds of long-nosed armadillos of the Pleistocene is related to Dasypus bellus , the "beautiful armadillo", which is considered to be closely related to the nine-banded armadillo, but with a total length of around 1.2 m and a weight of over 15 kg had significantly larger dimensions. This close relationship was contradicted in a study from 2008 in which Dasypus bellus is related to the Kappler armadillo, but phylogenetic studies in 2015 confirm the close relationship with the nine-banded armadillo.

The first mention of the nine-banded armadillo in Europe was based on the Spanish conquistador Francisco Hernández de Córdoba , who discovered the Yucatán peninsula in 1517 and used the local Aztec word azotochtli for the animal in his writings . Azotochtli means something like "turtle hare" and refers to the appearance of the animal. The first description of the nine-banded armadillo was in 1758 by Linnaeus , where he specified "in America Meridionali" as a type occurrence. He differentiated the armadillo from the seven-banded armadillo ( Dasypus septemcinctus ) on the basis of the number of movable ligaments, although he assumed India as the distribution area for the latter . Linnaeus translated the generic name Dasypus from the word Azotochtli into the Greek language , as he found the latter to be unsuitable for Latinization and thus for use in the binary nomenclature that he had introduced. It is Dasypus as tautonyme form of Azotochtli to look at and for this reason has the same meaning. The species name novemcinctus refers to the number of freely moving ligaments.

Nine-banded armadillo and man

Nine-banded armadillo in Silver River State Park, Florida

meaning

As the only armadillo species, the nine-banded armadillo has been able to expand its range dramatically since the arrival of Europeans in North America. The relationship between humans and these animals is ambivalent: on the one hand, they are valued for their meat and as insect eaters. On the other hand, they are persecuted because they occasionally dig up fields and eat bird eggs, but these only make up a fraction of their diet. These animals play an important role in medical research: Since the armadillo can carry both the leprosy bacterium Mycobacterium leprae and the parasite Trypanosoma cruzi, which causes Chagas disease , treatment methods and vaccines are tested on them.

Threat and protection

No major threats to the nine-banded armadillo are known, but it is hunted within its entire range. It is the main food of the Aché Indian people in Paraguay , who, according to a study, shot a total of 1500 individuals of this armadillo between 1980 and 1996, which corresponded to a total weight of 5.7 t and accounted for about 32% of their required food. In other regions it is a popular game dish, some of which is served in a tank. In North America, nine-banded armadillos are often killed in car accidents, and their habit of jumping up when frightened is their undoing here. It is the third most common vehicle-killed mammal species in the southern Great Plains . Overall, the nine-banded armadillo is widespread and is classified by the IUCN as “ least concern ”, with an increasing population size. It occurs in numerous protected areas within the entire range.

literature

  • Karen McBee, Robert J. Baker: Dasypus novemcinctus. In: Mammalian Species. 162, 1982, pp. 1-9.
  • CM McDonough and WJ Laughry: Dasypodidae (Long-nosed armadillos). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 30–47 (pp. 44–45) ISBN 978-84-16728-08-4 .

Individual evidence

  1. ↑ a b c d e f g h i j k l m n o p q Paul Smith: Nine-banded armadillo Dasypus novemcinctus Linnaeus 1758. In: Mammals of Paraguay. 8, 2008, pp. 1-30.
  2. ↑ a b c d e f g h i j Karen McBee, Robert J. Baker: Dasypus novemcinctus. In: Mammalian Species. 162, 1982, pp. 1-9.
  3. ↑ a b c d e f g h i j C. M. McDonough and WJ Laughry: Dasypodidae (Long-nosed armadillos). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 30–47 (pp. 44–45) ISBN 978-84-16728-08-4 .
  4. ^ Hugh H. Genoways, Robert M. Timm: The Xenarthrans of Nicaragua. In: Mastozoologia Neotropical. 10 (2), 2003, pp. 231-253.
  5. ^ SF Vizcaíno, N. Milne: Structure and function in armadillo limbs (Mammalia: Xenarthra: Dasypodidae). In: Journal of Zoology. 257, 2002, pp. 117-127.
  6. ↑ Agustín M. Abba, M. Superina: Dasypus novemcinctus. In: Edentata. 11 (2), 2010, p. 160.
  7. ↑ a b c d e f Mariella Superina: Biology and keeping of armadillos (Dasypodidae). University of Zurich, 2000, OCLC 632874842 .
  8. ^ Edward Drinker Cope: On the zoological position of Texas. In: Bulletin of the United States National Museum. 17, 1880, pp. 1-51.
  9. ↑ Jennifer K. Frey, James N. Stuart: Nine-Banded Armadillo (Dasypus novemcinctus) Records in New Mexico, USA. In: Edentata. 8-10, 2009, pp. 54-55.
  10. ^ S. Erich Eichler, Timothy J. Gaudin: New records of the nine-banded armadillo, Dasypus novemcinctus, in southeast Tennessee, and their implications. In: Edentata. 12, 2011, pp. 7-13.
  11. ↑ Maria Clara Arteaga, Eduardo Martins Venticinque: Influence of topography on the location and density of armadillo burrows (Dasypodidae: Xenarthra) in the central Amazon, Brazil. In: Mammalian Biology. 73, 2008, pp. 262-266.
  12. ↑ James N. Layne: Above-ground nests of the nine-banded armadillo in Florida. In: Florida Field Naturalist. 12, 1984, pp. 58-61.
  13. ↑ Kent H. Redford: Dietary specialization and variation in two mammalian myrmecophages (variation in mammalian myrmecophagy). In: Revista Chilena de Historia Natural. 59, 1986, pp. 201-208.
  14. ↑ Teresa Cristina da Silveira Anacleto: Food Habits of Four Armadillo Species in the Cerrado Area, Mato Grosso, Brazil. In: Zoological Studies. 46, 2007, pp. 529-537.
  15. ↑ Vanderson Corrêa Vaz, Ricardo Tadeu Santori, Ana Maria Jansen, Ana Cláudia Delciellos, Paulo Sérgio D'Andrea: Notes on food habits of armadillos (Cingulata, Dasypodidae) and anteaters (Pilosa, Myrmecophagidae) at Serra da Capivara National Park (Piauí State , Brazil). In: Edentata. 13, 2012, pp. 84-89.
  16. ↑ Eduardo Carrillo and Todd K. Fuller: Predation of a Central American coral snake (Micrurus nigrocinctus) by a nine-banded armadillo (Dasypus novemcinctus) in Santa Rosa National Park, Costa Rica. In: Edentata. 19, 2018, pp. 67-69.
  17. ↑ Samuel M. Simkin, William K. Michener: Faunal Soil Disturbance Regime of a Longleaf Pine Ecosystem. In: Southeastern Naturalist. 4 (1), 2005, pp. 133-152.
  18. ^ Clara R. Rojas-Suárez, Leonardo Maffei: Reproducción de Dasypus novemcinctus en el Izozog, Santa Cruz, Bolivia. In: Edentata. 5, 2003, pp. 47-54.
  19. ↑ Mariella Superina, WJ Loughry: Life on the Half Shell: Consequences of a carapace in the Evolution of Armadillos (Xenarthra: Cingulata). In: Journal of Mammal Evolution. 19, 2012, pp. 217-224.
  20. ↑ Silvia D. Frutos, Ronald A. van den Bussche: Genetic diversity and gene flow in Nine-banded armadillos in Paraguay. In: Journal of Mammalogy. 83 (3), 2002, pp. 815-823.
  21. ↑ Loren A. Binns, WJ Loughry, Colleen M. McDonough, Corey Devin Anderson: Spatial genetic structure within a population of nine-banded armadillos in western Mississippi. In: Journal of Mammalogy. 101 (1), 2020, pp. 143-150, doi: 10.1093 / jmammal / gyz174 .
  22. ^ AR Rabinowitz, BG Nottingham: Ecology and Behavior of the Jaguar Panthera onca in Belize, Central America. In: Journal of Zoology. 210, 1986, pp. 149-159.
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  24. ↑ Rogério Martins, Juliana Quadros, Marcello Mazzolli: Hábito Alimentar e Interferência Antrópica na Atividade de Marcação Territorial do Puma concolor e Leopardus pardalis (Carnivora: Felidae) e Outros Carnívoros na Estação Ecológica de Juréia, São Paulo, Brasil, Brazil. In: Revista Brasileira de Zoologia. 25, 2008, pp. 427-435.
  25. ^ AA Guglielmone, A. Estrada-Peña, CA Luciani, AJ Mangold, JE Kerans: Hosts and distribution of Amblyomma auricularium (Conil 1878) and Amblyomma pseudoconcolor Aragão, 1908 (Acari: Ixodidae). In: Experimental and Applied Acarology. 29, 2003, pp. 131-139.
  26. ↑ Richard W. Truman, Pushpendra Singh, Rahul Sharma, Philippe Busso, Jacques Rougemont, Alberto Paniz-Mondolfi, Adamandia Kapopoulou, Sylvain Brisse, David M. Scollard, Thomas P. Gillis, Stewart T. Cole: Probable Zoonotic Leprosy in the Southern United States. In: The New England Journal of Medicine. 364 (17), 2011, pp. 1626-1633.
  27. ↑ a b c d Gillian C. Gibb, Fabien L. Condamine, Melanie Kuch, Jacob Enk, Nadia Moraes-Barros, Mariella Superina, Hendrik N. Poinar, Frédéric Delsuc: Shotgun Mitogenomics Provides a Reference Phylogenetic Framework and Timescale for Living Xenarthrans. In: Molecular Biology and Evolution. 33 (3), 2015, pp. 621–642.
  28. ↑ Timothy J. Gaudin, John R. Wible: The phylogeny of living and extinct armadillos (Mammalia, Xenarthra, Cingulata): a craniodental analysis. In: Matthew T. Carrano, Timothy J. Gaudin, Richard W. Blob, John R. Wible (Eds.): Amniote Paleobiology: Phylogenetic and Functional Perspectives on the Evolution of Mammals, Birds and Reptiles. University of Chicago Press, Chicago 2006, pp. 153-198.
  29. ↑ Laureano Raúl González Ruiz, Gustavo Juan Scillato-Yané: A new Stegotheriini (Mammalia, Xenarthra, Dasypodidae) from the “Notohippidian” (early Miocene) of Patagonia, Argentina. In: New Yearbook of Geology and Paleontology, Treatises. 252 (1), 2009, pp. 81-90.
  30. ↑ a b Ascanio D. Rincón, Richard S. White, H. Gregory Mcdonald: Late Pleistocene Cingulates (Mammalia: Xenarthra) from Mene De Inciarte Tar Pits, Sierra De Perijá, Western Venezuela. In: Journal of Vertebrate Paleontology. 28 (1), 2008, pp. 197-207.
  31. ↑ a b Maren Möller-Krull, Frédéric Delsuc, Gennady Churakov, Claudia Marker, Mariella Superina, Jürgen Brosius, Emmanuel JP Douzery, Jürgen Schmitz: Retroposed Elements and Their Flanking Regions Resolve the Evolutionary History of Xenarthran Mammals (Armadillos, Anteaters and Sloths) . In: Molecular Biology and Evolution. 24, 2007, pp. 2573-2582.
  32. ↑ a b Frédéric Delsuc, Mariella Superina, Marie-Ka Tilak, Emmanuel JP Douzery, Alexandre Hassanin: Molecular phylogenetics unveils the ancient evolutionary origins of the enigmatic fairy armadillos. In: Molecular Phylogenetics and Evolution. 62, 2012, pp. 673-680.
  33. ↑ Dorothée Huchon, Frédéric Delsuc, François M. Catzeflis, Emmanuel JP Douzery: Armadillos exhibit less genetic polymorphism in North America than in South America: nuclear and mitochondrial data confirm a founder effect in Dasypus novemcinctus (Xenarthra). In: Molecular Ecology. 8 (10), 1999, pp. 1743-1748.
  34. ^ Maria Clara Arteaga, Daniel Piñero, Luis E. Eguiarte, Jaime Gasca, Rodrigo A. Medellín: Genetic structure and diversity of the Nine-banded armadillo in Mexico. In: Journal of Mammalogy. 93 (2), 2012, pp. 547-559.
  35. Jump up ↑ Lionel Hautier, Guillaume Billet, Benoit de Thoisy and Frédéric Delsuc: Beyond the carapace: skull shape variation and morphological systematics of long-nosed armadillos (genus Dasypus). In: PeerJ. 5, 2017, p. E3650, doi: 10.7717 / peerj.3650 .
  36. ↑ Beth Shapiro, Russell W. Graham, Brandon Leets: A revised evolutionary history of armadillos (Dasypus) in North America based on ancient mitochondrial DNA. In: Boreas. 44 (1), 2015, pp. 14-23, doi: 10.1111 / bor.12094 .
  37. ↑ Esteban Soibelzon, LS Avilla, M. Castro: The cingulates (Mammalia: Xenarthra) from the late Quaternary of northern Brazil: Fossil records, paleoclimates and displacements in America. In: Quaternary International. 377, 2015, pp. 118–125, doi: 10.1016 / j.quaint.2015.02.052 .
  38. ↑ Paulo V. Oliveira, Ana Maria Ribeiro, Édison V. Oliveira, Maria Somália S. Viana: The Dasypodidae (Mammalia, Xenarthra) from the Urso Fóssil Cave (Quaternary), Parque Nacional de Ubajara, State of Ceará, Brazil: paleoecological and taxonomic aspects. In: Anais da Academia Brasileira de Ciências. 86 (1), 2014, pp. 147–158.
  39. ^ H. Gregory McDonald: Paleoecology of extinct Xenarthans and the Great American Biotic Interchange. In: Bulletin of the Florida Museum of Natural History. 45 (4), 2005, pp. 313-333.
  40. ↑ Mariela C. Castro, Ana Maria Ribeiro, Jorge Ferigolo, Max C. Langer: Redescription of Dasypus punctatus Lund, 1840 and considerations on the genus Propraopus Ameghino, 1881 (Xenarthra, Cingulata). In: Journal of Vertebrate Paleontology. 33 (2), 2013, pp. 434-447.
  41. ↑ Mariela C. Castro, Martín R. Ciancio, Victor Pacheco, Rodolfo M. salas-Gismondi, J. Enrique Bostelmann, Alfredo A. Carlini: Reassessment of the hairy long-nosed armadillo “Dasypus” pilosus (Xenarthra, Dasypodidae) and revalidation of the genus Cryptophractus Fitzinger, 1856. In: Zootaxa. 3947 (1), 2015, pp. 30-48, doi: 10.11646 / zootaxa.3947.1.2 .
  42. ↑ JM Anderson, K. Benirschke: The armadillo, Dasypus novemcinctus, in experimental biology. In: Laboratory Animal Care. 16, 1966, pp. 202-216.
  43. ↑ Oldfield Thomas: The mammals of the tenth edition of Linnaeus; an attempt to fix the types of the genera and the exact bases and localities of the species. In: Proceedings of the Zoological Society of London. 1911, pp. 120-158.
  44. ^ Agustín M. Abba, Mariella Superina: Dasypus novemcinctus. In: IUCN 2012: IUCN Red List of Threatened Species. Version 2012.2. last accessed on March 27, 2013.

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