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Overview

Brief Summary

Description

As their common name suggests, this species is the largest of the living anteaters and is instantly recognised around the world. They are strange-looking animals, with the body roughly divided into three equal regions: the long nose and head, the body and the tail (3). Beneath the skin of the long nose is a bone tube formed by the fusion of the upper and lower jaw (3). The nose houses an impressively long, sticky tongue, which can measure up to 50 centimetres in length (3). They are protected from the bites of soldier ants and termites by their rubbery skin and very long hairs, which can measure up to 45 centimetres in length (3). A black stripe runs from beneath the snout to the mid-torso and is banded by white or cream (5). The front feet bear huge claws and giant anteaters walk on their knuckles with their claws folded up into their palms for protection (5). Male and female anteaters look so alike that females can only be identified when they are accompanied by their offspring (3). The name of the giant anteater in Portugese is 'tamandua-bandeira'; the word tamandua is of Indian origin and means 'insect-eater' and 'bandeira' is a portugese word meaning 'flag'. This name refers to the large tail of the species, which is thought to look like a flag (3).
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Biology

Giant anteaters are predominantly solitary, except for mothers and their offspring (7). A single offspring is produced after a gestation period of 190 days (7). They are carried on their mother's back, aligned with the white stripe, so that they are very well camouflaged (2). Young are weaned after two months, although they may continue to be carried on their mother's back until they are nine months old (7). Recently, females have been discovered to occasionally produce a second infant, carrying the newborn on the back whilst accompanied by the older infant on foot (3). There is considerable regional variation in behaviour of giant anteaters. In some areas, they are largely nocturnal, but in other regions they are active mainly during the day (8). They are almost blind, but have a very keen sense of smell (3). The majority of the diet is made up of ants and termites, which are detected by smell (5). Once a termite mound or anthill has been located, the anteater rips it open with its powerful claws and 'drinks' in the prey by creating a vacuum in its throat, sucking the insects in, aided by their long sticky tongues (3). An individual may eat up to 30,000 ants in one day (7). Contrary to popular belief, giant anteaters rarely feed from tall termite mounds, but seem to prefer low mounds (3). Ants and termites are not especially nutritious and to compensate for this, the giant anteater conserves energy by moving around slowly. They allow their body temperatures to fluctuate within limits, and they have the lowest body temperature of any terrestrial mammal, reaching as low as 32 degrees Celsius (3). Anteaters sleep for as much as 16 hours a day out in the open with their tail wrapped around them like a blanket (3).
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Distribution

Giant anteaters are found in Central and South America, from southern Belize and Guatemala to northern Argentina.

Biogeographic Regions: neotropical

  • Redford, K., J. Eisenberg. 1992. Mammals of the Neotropics. Chicago, IL: University of Chicago Press.
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Range Description

Myrmecophaga tridactyla has been recorded from Honduras in Central America, south through South America to the Gran Chaco region of Bolivia, Paraguay and Argentina. Within Central America, the species has disappeared from much of its range, with recent sightings generally confined to highland regions. The giant anteater is considered the most threatened mammal of Central America; it seems to be extinct in Belize and Guatemala, and has disappeared from parts of Costa Rica. It is not listed in Owen and Girón's (2002) checklist of mammals of El Salvador. In South America, this species seems to be extinct in Uruguay (Fallabrino and Castiñeira 2006) and in the state of Santa Catarina, Brazil (Cherem et al. 2004). It is listed as Regionally Extinct in the Brazilian states of Rio de Janeiro (Bergallo et al. 2000) and Espírito Santo (Chiarello et al. 2007). It is classified as Critically Endangered in Paraná (Mikich and Bérnils 2004) and in Rio Grande do Sul, Brazil (Fontana et al. 2003) but will be categorized as Extinct in the next update of this state's Red List (C. Kasper pers. comm. 2009). Its presence in Ecuador west of the Andes needs to be confirmed.
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Range

The range extends from the southern tip of Mexico to Uruguay and northwestern Argentina, although the highest densities are found in South America (5). They are found in all regions of Brazil but are very rare or locally extinct in highly disturbed areas and are commonly found in protected areas (6).
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Physical Description

Morphology

Giant anteaters are quite distinctive morphologically, they are the largest of the anteater species. The snout is long (up to 45 cm in length) and the skull is streamlined with small eyes and ears. The tail is large and bushy and is nearly as long as the body. Head and body length measures 1,000 to 1,200 mm and tail length 650 to 900 mm. Weight ranges from 18 to 39 kg. Giant anteaters have fur that is thick and coarse and longer towards the tail. Their coat is brown with black and white stripes on the shoulders and a crest of hair along the middle of the back. The forelegs are white with black bands at the toes. Their hindfeet have 5 short claws, while their forefeet have 5 claws with the inner 3 being very long and sharp. They walk on the wrists of their forefeet, with these large claws curled out of the way. Giant anteaters have no teeth. The tongue can be extended 610 mm outside of the mouth and has spine-like protrusions.

Range mass: 18 to 39 kg.

Range length: 1 to 1.2 m.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: sexes alike

Average basal metabolic rate: 14.543 W.

  • Naples, V. 1999. Morphology, evolution, and function of feeding in the giant anteater. Journal of Zoology-London, 249: 19-41.
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Ecology

Habitat

Chocó-Darién Moist Forests Habitat

This taxon can be found in the Chocó-Darién moist forests ecoregion, one of the most species rich lowland areas on Earth, with exceptional abundance and endemism over a broad range of taxa including plants, birds, amphibians and arthropods. The biological distinctiveness is exceptional, with considerable biodiversity.

There are three principal geomorphologic types in the ecoregion: alluvial plains of recent origin, low mountains formed by the relatively recent dissection of sediments from the Tertiary and Pleistocene periods, and the complexes in mountain areas consisting of mesozoic rocks. The high precipitation and the topography mean that the ecoregion includes a complex of great hydrographic basins, the most important being those of the Atrato, Baudó, and San Juan Rivers and the Micay and Patía Rivers in the south. The force of the water in many of these rivers form deep gorges cutting through the mountains, creating spectacular rapids and waterfalls in the mountains. At lower elevations, large rivers become very wide and with many meanders. Given the high precipitation in the region, it is not surprising that the soils are severely leached and poor in nutrients. Most of the ecoregion has typical laterite soils with reddish clay, although the soils are younger and less leached in some areas, especially close to the base of the Andes and in the floodplains of the major rivers. Of particular botanical interest are the white clay soils in the region of Bajo Calima in Colombia, which are associated with the gigantic sclerophyllous leafed and unusually large fruited vegetation.

Depending on the altitudinal gradient, soil water content and the effect of the sea, there are various types of vegetation that make up the ecoregion. In broad terms, in the northern part of the ecoregion, the lowland rainforests correlate to the Brosimun utilis alliance, including communities dominated by the deciduous Cuipo tree (Cavanillesia platanifolia), the Espavé wild cashew (Anacardium excelsum), the Panamanian rubber tree (Castilla elastica), Brosimum guianense, Bombacopsis spp., Ceiba pentandra, Dipteryx panamensis, and others. In the undergrowth Mabea occidentalis, Clidemia spp., Conostegia spp. and Miconia spp. are abundant. In zones that are occasionally flooded, the Cativo (Prioria copaifera) flourishes as well. In the southern part of the ecoregion, these rainforests have multiple strata, with two layers of trees, lianas, and epiphytes with vigorous growth rates. The number of deciduous plants increases in the north and south, where there is a dry season, particularly near the coast. The forests at higher altitudes, starting at 600 meters, have communities with the following species: Guamos (Inga spp.), Billia columbiana, Brosimum sp., Sorocea spp., Jacaranda hesperia, Pourouma chocoana, Guatteria ferruginea, Cecropia spp., Elaegia utilis, and Brunellia spp.

There are at least 127 species of amphibians in the Choco-Darien, including the following endemic anuran species: Isla Bonita robber frog (Craugastor crassidigitus); Kokoe poison frog (Phyllobates aurotaenia NT), found on western slopes of the Cordillera Occidental , along the Ra­o San Juan drainage south to the Ra­o Raposo; Golden poison frog (Phyllobates terribilis EN); La Brea poison frog (Oophaga occultator); Andagoya robber frog (Pristimantis roseus); Antioquia beaked toad (Rhinella tenrec); Atrato glass frog (Hyalinobatrachium aureoguttatum); Blue-bellied poison arrow frog (Ranitomeya minuta); Colombian egg frog (Ctenophryne minor), known only to the in the upper Ra­o Saija drainage; Condoto stubfoot toad (Atelopus spurrelli VU); Flecked leaf frog (Phyllomedusa psilopygion); LeDanubio robber frog (Strabomantis zygodactylus). An endemic salamander present in the Choco-Darien is the Finca Chibigui salamander (Bolitoglossa medemi VU).

Some other non-endemic anurans found here are: Anatipes robber frog (Strabomantis anatipes); Banded horned treefrog (Hemiphractus fasciatus); Black-legged poison frog (Phyllobates bicolor NT); Horned marsupial frog (Gastrotheca cornuta EN), known for having the largest amphibian eggs in the world; El Tambo stubfoot toad (Atelopus longibrachius EN); Elegant stubfoot toad (Atelopus elegans CR). Endemic caecilians in the ecoregion include the Andagoya caecilian (Caecilia perdita).

There are a number of reptilian taxa within the ecoregion, including: Adorned graceful brown snake (Rhadinaea decorata); the endemic Black centipede snake (Tantilla nigra); Boulenger's least gecko (Sphaerodactylus scapularis VU); the endemic Iridescent ground snake (Atractus iridescens); the endemic Cauca coral snake (Micrurus multiscutatus); the endemic Colombian coral snake (Micrurus spurelli); the endemic Dark ground snake (Atractus melas); the endemic Colombian mud turtle (Atractus melas VU); and the endemic Echternacht's ameiva (Ameiva anomala).

There are 577 species of birds recorded; Tyrannidae is listed as the most diverse avian family, presenting 28 genera and 60 species within the ecoregion. The Choco-Daroemis is a center of avian endemism of the Neotropics; moreover, according to Stattersfield, this ecoregion spans two Endemic Bird Areas, one in Central America and one in South America.

Between these two Endemic Bird Areas there are over sixty restricted range species, including the Chocó tinamou (Crypturellus kerriae VU), Chestnut-mantled Oropendola (Psarocolius cassini EN), Viridian dacnis (Dacnis viguieri), Crested ant-tanager (Habia cristata), Lita woodpecker (Piculus litea), and Plumbeous forest-falcon (Micrastur plumbeus EN). Also to be noted is the presence of the Harpy eagle (Harpia harpyja), the Black and white crowned eagle (Spizastur melanoleucus), taxa increasingly rare in many areas of the Neotropics, and possibly the Speckled antshrike (Xenornis setifrons EN) although one has not been recorded in Colombia since the 1940s.

The region is rich in mammalian taxa, but the larger animals have received inadequate research. These include the Bush dog (Speothos venaticus NT); Chocó tamarin (Saguinus geoffroyi EN), the Baird's Tapir (Tapirus bairdii EN), the Giant anteater (Myrmecophaga tridactyla VU), the Brown-headed spider monkey (Ateles fuscipens CR), the Puma (Puma concolor VU), the Ocelot (Leopardus pardalis LC), and the jaguar (Panthera onca NT).

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Mesoamerican Gulf-Caribbean Mangroves Habitat

This taxon is found in the Mesoamerican Gulf-Caribbean mangroves ecoregion, but not necessarily exclusive to this region.The Mesoamerican Gulf-Caribbean mangroves occupy a long expanse of disjunctive coastal zone along the Caribbean Sea and Gulf of Mexico for portions of Central America and Mexico. The ecoregion has a very high biodiversity and species richness of mammals, amphibians and reptiles. As with most mangrove systmems, the Mesoamerican Gulf-Caribbean ecoregion plays an important role in shoreline erosion prevention from Atlantic hurricanes and storms; in addition these mangroves are significant in their function as a nursery for coastal fishes, turtles and other marine organisms.

This disjunctive Neotropical ecoregion is comprised of elements lying along the Gulf of Mexico coastline of Mexico south of the Tampico area, and along the Caribbean Sea exposures of Belize, Honduras, Guatemala, Nicaragua, Costa Rica and Panama.There are 507 distinct vertebrate species that have been recorded in the Mesoamerican Gulf-Caribbean mangroves ecoregion.

Chief mangrove tree species found in the central portion of the ecoregion (e.g. Belize) are White Mangrove (Laguncularia racemosa), Red Mangrove (Rhizophora mangle), and Black Mangrove (Avicennia germinans); Buttonwood (Conocarpus erectus) is a related tree associate. Red mangrove tends to occupy the more seaward niches, while Black mangrove tends to dominate the more upland niches. Other plant associates occurring in this central part of the ecoregion are Swamp Caway (Pterocarpus officinalis), Provision Tree (Pachira auatica) and Marsh Fern (Acrostichum aureum).

The Mesoamerican Gulf-Caribbean mangroves ecoregion has a number of mammalian species, including: Mexican Agouti (Dasyprocta mexicana, CR); Mexican Black Howler Monkey (Alouatta pigra, EN); Baird's Tapir (Tapirus bairdii, EN); Central American Spider Monkey (Ateles geoffroyi, EN); Giant Anteater (Myrmecophaga tridactyla); Deppe's Squirrel (Sciurus deppei), who ranges from Tamaulipas, Mexico to the Atlantic versant of Costa Rica; Jaguar (Panthera onca, NT), which requires a large home range and hence would typically move between the mangroves and more upland moist forests; Margay (Leopardus wiedii, NT); Mantled Howler Monkey (Alouatta palliata); Mexican Big-eared Bat (Plecotus mexicanus, NT), a species found in the mangroves, but who mostly roosts in higher elevation caves; Central American Cacomistle (Bassariscus sumichrasti).

A number of reptiles have been recorded within the ecoregion including the Green Turtle (Chelonia mydas, EN); Hawksbill Sea Turtle (Eretmochelys imbricata, CR); Central American River Turtle (Dermatemys mawii, CR), distributed along the Atlantic drainages of southern Mexico to Guatemala; Morelets Crocodile (Crocodylus moreletii, LR/CD), a crocodile found along the mangroves of Yucatan, Belize and the Atlantic versant of Guatemala.

Some of the other reptiles found in this ecoregion are the Adorned Graceful Brown Snake (Rhadinaea decorata); Allen's Coral Snake (Micrurus alleni); Eyelash Palm Pitviper (Bothriechis schlegelii); False Fer-de-lance (Xenodon rabdocephalus); Blood Snake (Stenorrhina freminvillei); Bridled Anole (Anolis frenatus); Chocolate Anole (Anolis chocorum), found in Panamanian and Colombian lowland and mangrove subcoastal forests; Furrowed Wood Turtle (Rhinoclemmys areolata. NT); Brown Wood Turtle (LR/NT); Belize Leaf-toed Gecko (Phyllodactylus insularis), which occurs only in this ecoregion along with the Peten-Veracruz moist forests.

Salamanders found in this ecoregion are: Cukra Climbing Salamander (Bolitoglossa striatula); Rufescent Salamander (Bolitoglossa rufescens); Alta Verapaz Salamander (Bolitoglossa dofleini, NT), the largest tropical lungless salamander, whose coastal range spans Honduras, Guatemala and the Cayo District of Belize; Colombian Worm Salamander (Oedipina parvipes), which occurs from central Panama to Colombia; La Loma Salamander (Bolitoglossa colonnea), a limited range taxon occurring only in portions of Costa Rica and Panama;.Central American Worm Salamander (Oedipina elongata), who inhabits very moist habitats; Cienega Colorado Worm Salamander (Oedipina uniformis, NT), a limited range taxon found only in parts of Costa Rica and Panama, including higher elevation forests than the mangroves; Limon Worm Salamander (Oedipina alfaroi, VU), a restricted range caecilian found only on the Atlantic versant of Costa Rica and extreme northwest Panama. Caecilians found in the ecoregion are represented by: La Loma Caecilian (Dermophis parviceps), an organism found in the Atlantic versant of Panama and Costa Rica up to elevation 1200 metres

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Panamanian Dry Forests Habitat

This taxon is found in the Panamanian dry forests, but not necessarily limited to this ecoregion. The Panamanian dry forests ecoregion occupies approximately 2000 square miles of coastal and near-coastal areas on the Pacific versant of Panama, around portions of the Gulf of Panama. Plant endemism is intermediate, and vertebrate species richness is quite high in the Panamanian dry forests.This key ecoregion is highly threatened from its extensive ongoing exploitation. Beyond the endemism and species richness, the ecoregion is further significant, since it offers a biological corridor from the moist forests to the coastal mangroves.

Plant endemism is intermediate in value within the Panamanian dry forests, likely elevated due to the (a) isolation of this ecoregion from the surrounding and intervening moist forest habitat; (b) arid conditions which likely enhanced speciation and hence species richness; and (c) absence of prehistoric glaciation, which has extinguished many species in more extreme latitudes.

Many of the plants are well adapted to herbivory defense through such morphologies as spiny exteriors and other features. Forest canopies are typically less than twenty meters, with a few of the highest species exceeding that benchmark. Caesalpinia coriaria is a dominant tree in the Azuero Peninsula portion of the dry forests, while Lozania pittieri is a dominant tree in the forests near Panama City. The vegetative palette is well adapted to the dry season, where water is a precious commodity.

Faunal species richness is high in the Panamanian dry forests, as in much of Mesoamerica, with a total of 519 recorded vertebrates alone within the Panamanian dry forests. Special status reptiles in the Panamanian dry forests include the American  Crocodile (Crocodylus acutus), the Lower Risk/Near Threatened Brown Wood Turtle (Rhinoclemmys annulata), the Lower Risk/Near Threatened Common Caiman (Caiman crocodilus), the Lower Risk/Near Threatened Common Slider (Trachemys scripta), and the Critically Endangered Leatherback Turtle (Dermochelys coriacea). There are two special status amphibian in the ecoregion: the Critically endangered plantation Glass Frog (Hyalinobatrachium colymbiphyllum) and the Vulnerable Camron mushroom-tongued salamander (Bolitoglossa lignicolor).

Threatened mammals found in the Panamanian dry forests include the: Endangered Central American Spider Monkey (Ateles geoffroyi), the Vulnerable Giant Anteater (Myrmecophaga tridactyla), the Near Threatened Handley’s Tailless Bat (Anoura cultrata), the Vulnerable Lemurine Night Monkey (Aotus lemurinus), the Near Threatened Margay (Leopardus wiedii), the Near Threatened Yellow Isthmus Rat (Isthmomys flavidus), the Near Threatened White-lipped Peccary (Tayassu pecari), and the Near Threatened Spectral Bat (Vampyrum spectrum). There are two special status bird species occurring in the ecoregion: the Endangered Great Green Macaw (Ara ambiguus) and the Near Threatened Olive-sided Flycatcher (Contopus cooperi).

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Giant anteaters use a variety of habitats, including swamp, forests, and grasslands. They choose secluded, covered areas for sleeping. Giant anteaters can be found in both rural and densely populated areas.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: savanna or grassland

  • Shaw, , Machado-Neto, Carter. 1987. Behavior of free-living giant anteaters. Biotropica, 19(3): 255-259.
  • Anderson, S. 1967. Recent Mammals of the World. New York: The Ronald Press Company.
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Habitat and Ecology

Habitat and Ecology
This terrestrial anteater is found in tropical moist forest, dry forest, savanna habitats and open grasslands; it has also been reported from the Gran Chaco (Meritt 2008, Noss et al. 2008) and timber plantations (Kreutz et al. 2013). Recent camera trap data suggest that the species may be more abundant in areas of upland forest in the Brazilian Amazon than previously thought (F. Miranda pers. comm. 2013). Giant anteaters require large areas for their survival, which should contain forested patches.

Animals are generally solitary. Once per year, the female gives birth to a single young. Gestation length is 171 to 184 days (Bartmann 1983, Patzl et al. 1998, Knott et al. 2013). The mother carries the offspring on its back for approximately six months. Data from captive animals in European zoos show several females reproducing for the first time at 18-22 months, and the oldest known reproductive captive dams were 20-24 years old (unpublished report provided by I. Schappert). For wild females, sexual maturity is usually indicated as being around 2 years however, as it is not possible to determine their age once they reach adult size and long-term population studies on giant anteaters are lacking, there are no data on the longevity, survival rates, or reproductive rates of wild giant anteaters.
The generation length of M. tridactyla in the wild is therefore unknown, but it is suspected to be around 7 years

Systems
  • Terrestrial
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Inhabits grassland savannas, deciduous woodlands and rainforests (5). In Brazil, it occurs in the cerrado, a large area of woodland and savannah that is one of the world's most important 'hot-spots' for biodiversity (3).
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Trophic Strategy

Giant anteaters eat ants, termites, and soft-bodied grubs. Using the long, sharp claws on their forelimbs, they open insect colonies and tree trunks. They then use the tongue to collect the eggs, larvae, and adult insects. The salivary glands secrete sticky saliva during feeding that coats the tongue. They only stay at one ant colony for a short period of time because soldier ants arrive but giant anteaters can consume a few thousand insects in minutes. The tongue is attached to the sternum and moves very quickly, flicking 150 times per minute. They may sometimes eat fruit.

Animal Foods: insects

Plant Foods: fruit

Primary Diet: carnivore (Insectivore )

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Associations

Giant anteaters, through their diet, have an enormous impact on local insect communities.

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Giant anteaters can use their immense front claws to defend themselves from predators, though their typical response to threat is to run away. Their size makes them invulnerable to all but the largest of predators, jaguars and pumas primarily. They are often killed by humans, either intentionally through hunting or unintentionally through collisions with cars.

Known Predators:

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Known predators

Myrmecophaga tridactyla is prey of:
Panthera onca
Puma concolor

This list may not be complete but is based on published studies.
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Known prey organisms

Myrmecophaga tridactyla preys on:
Insecta

This list may not be complete but is based on published studies.
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Life History and Behavior

Behavior

Most communication occurs between young and their mothers or during fighting. It consists of snorts, sniffs, and hisses, as well as roaring during fights. Sight and hearing are diminished. Smell is highly developed-40 times that of humans.

Communication Channels: tactile ; acoustic ; chemical

Perception Channels: visual ; tactile ; acoustic ; chemical

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Life Expectancy

Giant anteaters have been known to live up to 25 years and 10 months in captivity. Their longevity in the wild is unknown.

Range lifespan

Status: captivity:
25 (high) years.

Average lifespan

Status: captivity:
26.0 years.

Average lifespan

Status: captivity:
25.8 years.

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Lifespan, longevity, and ageing

Maximum longevity: 31 years (captivity) Observations: One specimen was estimated to be 31 years old when it died in captivity (Richard Weigl 2005).
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Reproduction

The mating system of M. tridactyla is not known. Reproductive behavior is primarily observed in captivity. The male stands over the female who lays on her side during copulation.

Gestation is approximately 190 days, after which females give birth to a single young which weighs about 1.3 kg. Females give birth standing up and immediately the young anteater climbs onto her back. Young are born with a full coat of hair and adult-like markings. Breeding occurs year-round in captivity and the wild, though seasonal breeding times have been reported in portions of their range. Inter-birth intervals can be as low as 9 months. Sexual maturity is reached between 2.5 and 4 years. The mammary glands are lateral to the 'armpits' on the chest.

Breeding interval: Giant anteaters can breed as often as every 9 months, though it is often longer.

Breeding season: Giant anteaters may breed year round, or seasonally depending on region.

Average number of offspring: 1.

Average gestation period: 190 days.

Average weaning age: 6 months.

Average time to independence: 24 months.

Range age at sexual or reproductive maturity (female): 2.5 to 4 years.

Range age at sexual or reproductive maturity (male): 2.5 to 4 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous

Average birth mass: 1600 g.

Average gestation period: 184 days.

Average number of offspring: 1.

Young are born with a full coat of hair and adult markings and are capable of clinging to their mother at birth. A mother will carry the baby on her back until it is almost half her size, about 6 to 9 months. Young suckle for 2 to 6 months. They become independent after about 2 years, or when the mother becomes pregnant again, which may be sooner.

Parental Investment: precocial ; female parental care

  • Shaw, , Machado-Neto, Carter. 1987. Behavior of free-living giant anteaters. Biotropica, 19(3): 255-259.
  • Redford, K., J. Eisenberg. 1992. Mammals of the Neotropics. Chicago, IL: University of Chicago Press.
  • Nowak, R. 1999. Walker's Mammals of the World, Volume I. Baltimore and London: The Johns Hopkins University Press.
  • Jones, M. 1982. Longevity of captive mammals. Zool. Garten, 52: 113-128.
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Evolution and Systematics

Functional Adaptations

Functional adaptation

Limbs dig efficiently: echigo mole
 

Echigo moles and other digging animals break the soil surface and move as much earth as possible per stroke with short, powerful limbs and sharp claws.

   
  "Quite different in character are the feet of the diggers, animals that habitually burrow into the earth. The friction drag of moving through the ground is potentially enormous, so the size of the limbs and the area through which they move must be kept to an absolute minimum; but at the same time, great strength is needed. The limbs of animals that lead an almost completely subterranean life, like the mole, are short and thick, and their feet are broad and powerful. Each short stroke of a foot must move as much earth as possible, and the mole's feet are spade-like with widely spaced digits. In addition, the claws of digging animals are usually large, sharp and strong, to do the work of a pickaxe in breaking the soil surface. The aardvark of South Africa (its Afrikaans name, 'earth-pig', refers to its rather pig-like head) is a curious animal that digs for food in termite's nests. Its feet are short and massive with large, almost hoof-like claws on each toe. It is said that one aardvark can dig a hole faster than six men with shovels. Not only does it dig into termite nests to eat the insects, the aardvark digs burrows 4m or more in length in which to hide during the day.

The armadillos of Central and South America are also powerful diggers, able to conceal themselves at amazing speed; they too have short, strong legs with daunting claws. The feet of the giant anteater, another excavator of ant and termite nests, are not massive as those of the aardvark. They are long and curved -- so much so that the anteater is forced to walk on the sides of its feet with an ungainly bow-legged gait. The anteater is a scratch-digger, not a maker of burrows, so its claws do not need to be as large." (Foy and Oxford Scientific Films 1982:179-180)
  Learn more about this functional adaptation.
  • Foy, Sally; Oxford Scientific Films. 1982. The Grand Design: Form and Colour in Animals. Lingfield, Surrey, U.K.: BLA Publishing Limited for J.M.Dent & Sons Ltd, Aldine House, London. 238 p.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage: Myrmecophaga tridactyla

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Specimens with Barcodes: 1
Species With Barcodes: 1
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Conservation

Conservation Status

Habitat destruction is the primary threat to giant anteaters. They are listed as Appendix II by the Convention on International Trade in Endangered Species (CITES). Appendix II is defined as a species not necessarily threatened to extinction but one that should be controlled in trade to avoid overuse. They are listed as vulnerable by the International Union for Conservation of Nature and Natural Resources (IUCN). 'Vulnerable' is defined as an estimated population reduction of 20% in the next 10 years.

CITES: appendix ii

IUCN Red List of Threatened Species: vulnerable

  • CITES Secretariat. 2003. "Convention on International Trade in Endangered Species of Wild Flora and Fauna" (On-line ). Accessed 02/02/03 at http://www.cites.org.
  • IUCN. 1997. "International Union for Conservation of Nature and Natural Resources" (On-line ). Accessed 02/02/03 at http://www.redlist.org.
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IUCN Red List Assessment


Red List Category
VU
Vulnerable

Red List Criteria
A2c

Version
3.1

Year Assessed
2014

Assessor/s
Miranda, F., Bertassoni, A. & Abba, A.M.

Reviewer/s
Superina, M.

Contributor/s
Rojano, C. & González-Maya, J.F.

Justification
Myrmecophaga tridactyla is widespread geographically, but there have been many records of population extirpation, especially in Central America (where it is considered the most threatened mammal) and the southern parts of its range. The dietary specificity, low reproductive rates, large body size, along with threats to habitat degradation in many parts of its range, have proved to be significant factors in its decline. The giant anteater is currently listed in a threat category in virtually all regional and national Red Lists. A population loss of at least 30% over the past 10 years has been estimated based on local extinctions, habitat loss, and deaths caused by fires and road kills. Based on this decline and past threats that are still ongoing today, it is likely that the population has suffered an overall reduction in population size of >30% over the last three generations (suspected to be around 21 years). Because of the real threats to this species and the noticeable declines, a precautionary assessment of Vulnerable is given. More data and population monitoring are required for this species, and a reassessment is recommended as soon as additional information becomes available.

History
  • 2010
    Vulnerable
  • 2006
    Near Threatened
  • 2006
    Near Threatened
    (IUCN 2006)
  • 1996
    Vulnerable
  • 1994
    Vulnerable
    (Groombridge 1994)
  • 1990
    Vulnerable
    (IUCN 1990)
  • 1988
    Vulnerable
    (IUCN Conservation Monitoring Centre 1988)
  • 1986
    Vulnerable
    (IUCN Conservation Monitoring Centre 1986)
  • 1982
    Vulnerable
    (Thornback and Jenkins 1982)
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Status

Classified as Near Threatened (NT) on the IUCN Red List 2007 (1), and listed on Appendix II of CITES (4).
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Population

Population
Myrmecophaga tridactyla is locally uncommon to rare. Habitat loss, roadkills, hunting, and wildfires have been substantially affecting the populations over the past 10 years, and there have been many records of population extirpation, especially in Central America and in the southern parts of its range.

Population Trend
Decreasing
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Threats

Major Threats
Myrmecophaga tridactyla is at risk from habitat loss in parts of its range, and this is a significant threat to Central American populations in particular. Where this species inhabits grassland habitats it is particularly susceptible to fires. In Brazil, burning of sugar cane plantations prior to their harvest leads to the death of significant numbers of giant anteaters due to severe burn injuries (F. Miranda pers. comm. 2013). Animals are sometimes killed on roads or by dogs. Giant anteaters are hunted for food throughout their distribution, and are additionally hunted as a pest, for pets or for illegal trade in some parts of their range.
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The giant anteater and its habitat are threatened by agricultural encroachment and fires, both natural and started by people (3).
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Management

Conservation Actions

Conservation Actions
Myrmecophaga tridactyla is listed on Appendix II of CITES. It has been recorded from many protected areas. It is listed on several national Red Data lists, and is protected as a national heritage species in some provinces in Argentina. There is a need to improve fire management practices, especially in sugarcane plantations and within the regions of grassland habitat occupied by this species. Population and genetic data, as well as habitat use information, are needed, especially for areas that are being subjected to land use change.
A reintroduction program is being carried out in Corrientes province, Argentina.
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Conservation

Law protects this species in the majority of countries where it is found (9), and it is listed on Appendix II of the Convention on International Trade in Endangered Species (CITES), which stipulates that any exports require a permit (4). The giant anteater also occurs within some protected areas such as Conservation International's Guayana Shield, one of the largest blocks of undisturbed rainforest in the world (10).
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Relevance to Humans and Ecosystems

Benefits

Giant anteaters are increasingly killed in car accidents.

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Giant anteaters are hunted for food, fur, and sport. They are also valuable for the criical ecosystem roles they play.

Positive Impacts: body parts are source of valuable material; controls pest population

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Wikipedia

Giant anteater

The giant anteater (Myrmecophaga tridactyla), also known as the ant bear, is a large insectivorous mammal native to Central and South America. It is one of four living species of anteaters and is classified with sloths in the order Pilosa. This species is mostly terrestrial, in contrast to other living anteaters and sloths, which are arboreal or semi-arboreal. The giant anteater is the largest of its family, 182–217 cm (5.97–7.12 ft) in length, with weights of 33–41 kg (73–90 lb) for males and 27–39 kg (60–86 lb) for females. It is recognizable by its elongated snout, bushy tail, long fore claws, and distinctively colored pelage.

The anteater can be found in multiple habitats, including grassland and rainforest. It forages in open areas and rests in more forested habitats. It feeds primarily on ants and termites, using its fore claws to dig them up and its long, sticky tongue to collect them. Though giant anteaters live in overlapping home ranges, they are mostly solitary except during mother-offspring relationships, aggressive interactions between males, and when mating. Mother anteaters carry their offspring on their backs until weaning them.

The giant anteater is listed as Vulnerable by the International Union for Conservation of Nature. It has been extirpated from many parts of its former range, including nearly all of Central America. Threats to its survival include habitat destruction, fire, and poaching for fur and bushmeat, although some anteaters inhabit protected areas. With its distinctive appearance and habits, the anteater has been featured in pre-Columbian myths and folktales, as well as modern popular culture.

Taxonomy and phylogeny[edit]

From The Cambridge Natural History, Volume X—Mammalia (1902)

The giant anteater got its binomial name from Carl Linnaeus in 1758. Its generic name, Myrmecophaga, and specific name, tridactyla, are both Greek, meaning "anteater" and "three fingers", respectively.[4] Myrmecophaga jubata was used as a synonym. Three subspecies have been tentatively proposed: M. t. tridactyla (ranging from Venezuela and the Guianas to northern Argentina), M. t. centralis (native to Central America, northwestern Colombia, and northern Ecuador), and M. t. artata (native to northeastern Colombia and northwestern Venezuela). The giant anteater is grouped with the semi-arboreal northern and southern tamanduas in the family Myrmecophagidae. Together with the family Cyclopedidae, whose only extant member is the arboreal silky anteater, the two families comprise the suborder Vermilingua.[5]

Anteaters and sloths belong to order Pilosa and share superorder Xenarthra with Cingulata (whose only extant members are armadillos). The two orders of Xenarthra split 66 million years ago (Mya) during the Late Cretaceous epoch. Anteaters and sloths diverged around 55 Mya, between the Paleocene and Eocene epochs. The Cyclopes lineage emerged around 30 Mya in the Oligocene epoch, while the Myrmecophaga and Tamandua lineages split 10 Mya in the Late Miocene subepoch.[6] During most of the Cenozoic era, anteaters were confined to South America, which was formerly an island continent. Following the formation of the Isthmus of Panama about 3 Mya, anteaters of all three extant genera invaded Central America as part of the Great American Interchange.[7]

The fossil record for anteaters is generally sparse.[8] Some known fossils include the Pliocene genus Palaeomyrmidon, a close relative to the silky anteater, Protamandua, the sister taxon to the clade that includes the giant anteater and the tamanduas from the Miocene, and Neotamandua, a sister taxon to Myrmecophaga.[9] Protamandua was larger than the silky anteater but smaller than a tamandua, while Neotamandua was larger, falling somewhere between a tamandua and a giant anteater. Protamandua did not appear to have feet specialized for terrestrial or arboreal locomotion, but it may have had a prehensile tail. Neotamandua, though, is unlikely to have had a prehensile tail and its feet were intermediate in form between those of the tamanduas and the giant anteater.[8] The species Neotamandua borealis was suggested to be an ancestor of the latter.[10]

The giant anteater is the most terrestrial of the living anteater species. Its ancestors may originally have been adapted to arboreal life;[9] the transition to life on the ground could have been aided by the expansion of open habitats such as savanna in South America and the availability there of colonial insects, such as termites, that provided a larger potential food source. Both the giant anteater and the southern tamandua are well represented in the fossil record of the late Pleistocene and early Holocene.[8]

Description[edit]

Lateral view

The giant anteater can be identified by its large size, elongated muzzle, and long tail.[11] It has a total body length of 182–217 cm (5.97–7.12 ft). Males weigh 33–41 kg (73–90 lb) and females weigh 27–39 kg (60–86 lb),[4] making the giant anteater the largest extant species in its suborder. The head of the giant anteater, at 30 cm (12 in) long,[12] is particularly elongated, even when compared to other anteaters.[13] Its tubular snout, which ends in its tiny mouth opening and nostrils, takes up most of its head. Its eyes and ears are relatively small.[4] It has poor eyesight,[11] but its sense of smell is 40 times more sensitive than that of humans.[14] Giant anteaters can live around 16 years in captivity.[11]

Mounted skeleton

Unlike those of other mammals, the necks of anteaters, especially the giant, are thicker than the backs of their heads.[13] A small hump can be found at the back of its neck. The giant anteater's coat is mostly grey and salted with white. Its fore limbs are white, with black bands around the wrists, while its hind limbs are dark. Thick black bands with white outlines stretch from throat to shoulder, where they end in triangular points. The body ends in a brown tail. The coat hairs are long, especially on the tail, which makes the tail look larger than it actually is. A stiff mane stretches along its back.[4] The bold pattern was thought to be disruptive camouflage, but a 2009 study suggests it is warning coloration.[15] While adult males are slightly larger and more muscular than females, with wider heads and necks, visual sex determination can be difficult. The (retracted) penis and testes are located internally between the rectum and urinary bladder in males, and females have a single pair of mammary glands near the armpits.[16]

The giant anteater has broad ribs. Despite its specific name, it has five toes on each foot. Four toes on the front feet have claws, which are particularly elongated on the second and third digits.[16] The giant anteater walks on its front knuckles, similar to the African apes, specifically gorillas and chimpanzees. Doing this allows the anteater to keep its claws out of the way while walking. The middle digits, which support most of its weight, are extended at the metacarpophalangeal joints and bent at the interphalangeal joints.[17] Unlike the front feet, the hind feet have short claws on all five toes and walk plantigrade.[4] As a "hook-and-pull" digger, the anteater's enlarged supraspinous fossa gives the teres major more leverage—increasing the front limbs' pulling power—and the triceps muscle helps power the flexion of the thickened third digit of the front feet.[18]

The giant anteater has a low body temperature for a mammal, about 33°C (91°F), a few degrees lower than typical a mammalian temperature of 36–38°C (97-100°F).[14] Xenarthrans in general tend to have lower metabolic rates than most other mammals, a trend thought to correlate with their dietary specializations and low mobility.[19]

Feeding anatomy[edit]

Tongue extended

The anteater has no teeth and is capable of very limited jaw movement. It relies on the rotation of the two halves of its lower jaw, held together by a ligament at the tip,[13] to open and close its mouth. This is accomplished by its masticatory muscles,[20] which are relatively underdeveloped.[13] Jaw depression creates an oral opening large enough for the slender tongue to flick out. It is typically 60 cm (24 in)[4] long and is triangular posteriorly, rounded anteriorly, and ends in a small, rounded tip.[13] The tongue is covered in backward-curving papillae and coated in thick, sticky saliva secreted from its enlarged salivary glands,[12] which allows the anteater to collect insects with it.

The tube-like rostrum and small mouth opening restrict the tongue to protrusion-retraction movements. During feeding, the tongue moves in and out around 160 times per minute (nearly three times per second). These movements are powered by the unique musculature of the anteater's long, large and flexible hyoid apparatus. The animal relies on the orientation of its head for aim. When fully extended, the tongue can reach 45 cm (18 in); longer than the length of the skull. The buccinators allow it to slide back in without losing attached food and tighten the mouth to prevent food from escaping as it protracts. When retracted, the tongue is held in the oropharynx by the hyoid muscles and the secondary palate, preventing it from blocking respiration. This retraction is aided by the long sternoglossus muscle, which is formed by the fusion of the sternohyoid and the hyoglossus, and does not attach to the hyoid.[13] Thus, the tongue is directly anchored to the sternum.[21]

Anteaters swallow at a much higher rate than most other mammals; when feeding they swallow almost continuously.[13] Before being swallowed, insects are crushed against the palate.[12] The anteater's stomach, similar to a bird's gizzard, has hardened folds and uses strong contractions to grind up the insects. The digestive process is assisted by small amounts of ingested sand and soil.[16] The anteater cannot produce stomach acid of its own, but uses the formic acid of its prey for digestion.[12]

Distribution and status[edit]

In the Pantanal, Brazil

The giant anteater is native to Central and South America. Its known range stretches from Honduras to northern Argentina, and fossil remains have been found as far north as northwestern Sonora, Mexico.[7] It is largely absent from the Andes[4] and has been extirpated in Uruguay.[2] It may also be extirpated in Belize, Costa Rica and Guatemala.[2] The species can be found in a number of habitats including both tropical rainforests and xeric shrublands,[11] provided enough prey is present to sustain it.[22]

The species is listed as Vulnerable by the IUCN, due to the number of regional extirpations,[2] and under Appendix II by CITES, tightly restricting trade in specimens of the animal.[23] Between 2000 and 2010, the total population declined by 30%.[2] In 1994, some 340 anteaters died due to wildfires at Emas National Park in Brazil.[24] The animal is particularly vulnerable to fires due to its slow movement and flammable coat.[4]

Human-induced threats include collision with vehicles, attacks by dogs, and destruction of habitat. The anteater is commonly hunted in Bolivia, both for sport and sustenance. The animal's thick, leathery hide is used to make equestrian equipment in the Chaco. In Venezuela, it is hunted for its claws.[22] Anteaters are killed for safety reasons, due to their reputation as dangerous animals. The giant anteater remains widespread. Some populations are stable and the animal can be found in various protected areas in the Amazon and the Cerrado.[22] It is officially protected in some Argentine provinces as a national heritage species.[2]

Behavior and ecology[edit]

Anteater sleeping under its tail

The giant anteater may use multiple habitats. A 2007 study of anteaters in the Brazilian Pantanal found the animals generally forage in open areas and rest in forested areas, possibly because forests are warmer than grasslands on cold days and cooler on hot days.[25] Anteaters can be either diurnal or nocturnal. A 2006 study in the Pantanal found those anteaters to be mostly nocturnal: they are most active during nighttime and early morning, and retire as the temperature rises. On colder days, anteaters start and end periods of activity earlier, shifting them into daylight hours, and may become diurnal.[26] Diurnal anteaters have been observed at Serra da Canastra.[27] Nocturnality in anteaters may be a response to human disturbances.[28]

Giant anteaters typically rest in dense brush, but may use tall grass on cooler days. They carve a shallow cavity in the ground for resting. The animal sleeps curled up with its bushy tail folded over its body. The tail serves both to conserve body heat and as camouflage. One anteater was recorded sleeping with its tail stretched out on a sunny morning with an ambient temperature of 17°C (63°F); possibly it was positioned this way to allow its body to absorb the sun's rays for warmth.[29]

Giant anteaters are good swimmers and are capable of moving through wide rivers.[4] They have been observed to bathe.[30] They are also able to climb and have been recorded ascending both termite mounds and trees while foraging. One individual was observed holding onto a branch with its feet just touching the ground.[31]

Spacing[edit]

Moving through vegetation

Giant anteaters' home ranges vary in size depending on the location. At Serra da Canastra National Park in Brazil, female anteaters have home ranges of around 3.67 km2 (910 acres), while males live in home ranges of 2.74 km2 (680 acres). In other parts of Brazil, anteaters may have 9-km2 ranges. Anteaters from Venezuela have been recorded living in home ranges as large as 25 km2 (6,200 acres).[27] Individuals mostly live alone, except for females that nurse their young and males courting females. Anteaters communicate their presence, status, and sexual condition with secretions from their anal glands. They also advertise their presence to other individuals though tree markings and urine.[32] They appear to be able to recognize each other's saliva by scent.[14]

Females appear to be more tolerant of females than males are of males, leading to greater overlap in female home ranges. Males are more likely to engage in agonistic behaviors,[27] such as slowly circling each other, chasing, or actual fighting.[14] Circling anteaters may cry a drawn-out "harrr". In combat, they slash at each other with their claws and wrestle each other to the ground.[4] Fighting anteaters may emit roars or bellows.[27] Males are possibly territorial.[14]

Foraging and predation[edit]

Anteater foraging

This animal is an insectivore, feeding mostly on ants or termites. In areas that experience seasonal flooding, like the grassy plains of the Venezuelan llanos, anteaters mainly feed on ants, because termites are less available.[14] Conversely, anteaters at Emas National Park eat mainly termites, which occur in high density on the grasslands.[33] At Serra da Canastra, anteaters switch between eating mainly ants during the wet season (October to March) and termites during the dry season (May to September).[14]

Anteaters track prey by their scent.[11] After finding a nest, the animal tears it open with its long fore claws and inserts its long, sticky tongue to collect its prey. An anteater spends one minute on average feeding at a nest, visiting up to 200 nests in one day and consuming as many as 30,000 insects.[16] The anteater may be driven away from a nest by the chemical or biting attacks of soldiers.[11] Some termite species rely on their fortified mounds for protection and many individuals escape while the predator digs.[33] These modes of defense prevent the entire colony from being eaten in one anteater attack.[14]

Other prey include the larvae of beetles and western honey bees that have located their hives in termite mounds. Anteaters may target termite mounds with bee hives.[14] In captivity, anteaters are fed mixtures made of milk, eggs, mealworms, and ground beef.[28] To drink, an anteater may dig for water when no surface water is available, creating waterholes for other animals.[30] Giant anteaters are prey for jaguars and pumas. They typically flee from danger by galloping, but if cornered,[28] will rear up on their hind legs and slash at the attacker.[34] The giant anteater's front claws are formidable weapons, capable of killing a jaguar.[35]

Although shy and will typically attempt to avoid humans, this species has been known to kill or injure humans who corner the animals, then wound and agitate them. Giant anteaters can deliver fatal wounds with their front claws. Two hunters were recently killed by giant anteaters in Brazil. In both cases, the hunters were agitating, wounding, and cornering the animals and the attacks appeared to be defensive behaviours.[36] In April 2007, an anteater at the Florencio Varela Zoo mauled a zookeeper with its front claws.[37][a]

Reproduction and parenting[edit]

Adult with offspring clinging to her back

Giant anteaters can mate throughout the year.[14] During courtship, a male consorts with an estrous female, following and sniffing her. Male and female pairs are known to feed at the same insect nest.[27] While mating, the female lies on her side as the male crouches over her. A couple may stay together for up to three days and mate several times during that period.[14] Gestation lasts around 190 days[27] and ends with the birth of a single pup,[16] which typically weighs around 1.4 kg (3.1 lb).[35] Females give birth standing upright.[14]

Pups are born with eyes closed and begin to open them after six days. The mother carries its dependent pup on its back.[28] The pup's black and white band aligns with its mother's, camouflaging it.[16] The young communicate with their mothers with sharp whistles and use their tongues during nursing. After three months, the pup begins to eat solid food and is fully weaned by ten months. The mother grooms her offspring during rest periods lasting up to an hour. Grooming peaks during the first three months and declines as the young reaches nine months of age, ending by ten months. The decline mirrors that of the weakening bond between mother and offspring; young anteaters usually become independent by nine or ten months.[4] Anteaters are sexually mature in 2.5–4 yr.[28]

In culture[edit]

Anteater mask and scratcher used by Kayapo boys in their ceremonies

In the mythology and folklore of the indigenous peoples of the Amazon Basin, the giant anteater is depicted as a trickster foil to the jaguar, as well as a humorous figure due to its long snout. In one Shipib tale, an anteater challenged a jaguar to a breath-holding contest under water, which the jaguar accepted. After the two removed their pelts and submerged, the anteater jumped out of the water and stole the jaguar's pelt, leaving the jaguar with the anteater's pelt. In a Yarabara myth, the evil ogre Ucara is transformed into one by the sun. This myth emphasizes the nearly immobile nature of the anteater's mouth, which was considered a burden.[39] The Kayapo people wear masks of various animals and spirits, including the anteater, during naming and initiation ceremonies. They believe women who touched anteater masks or men who stumbled while wearing them would die or receive some sort of physical disorder.[40]

During the Spanish colonization of the Americas, the giant anteater was one of many native fauna taken to Europe for display. At first, Europeans believed all anteaters were female and mated with their noses, a misconception corrected by naturalist Félix de Azara.[41] In the 20th century, Salvador Dalí wrote imaginatively that the giant anteater "reaches sizes bigger than the horse, possesses enormous ferocity, has exceptional muscle power, is a terrifying animal." Dalí depicted an anteater in the style of The Great Masturbator. It was used as a bookplate for André Breton, who compared the temptations a man experiences in life to what "the tongue of the anteater must offer to the ant."[42]

The 1940 Max Fleischer cartoon Ants in the Plants features a colony of ants fighting off a villainous anteater. Released during the Phoney War, the film may have alluded to France's Maginot Line.[43] An anteater is also a recurring character in the comic strip B.C.. This character was the inspiration for Peter the Anteater, the University of California, Irvine team mascot.[44] The Flash cartoon Happy Tree Friends features an anteater named Sniffles. In the Stephen King miniseries Kingdom Hospital, the character Antubis appears in the form of an anteater-like creature with razor-sharp teeth.[45]

Notes[edit]

  1. ^ The animal in question, a male named Ramon, had a reputation for aggressiveness.[38]

References[edit]

  1. ^ Gardner, A. L. (2005). "Order Pilosa". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. p. 102. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  2. ^ a b c d e f Miranda, F., Bertassoni, A. & Abba, A.M. (2014). "Myrmecophaga tridactyla". IUCN Red List of Threatened Species. Version 2014.1. International Union for Conservation of Nature. Retrieved 2014-07-07. 
  3. ^ Linnæus, Carl (1758). Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I (in Latin) (10 ed.). Holmiæ: Laurentius Salvius. p. 35. Retrieved 23 November 2012. 
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