Short-beaked Echidna

Short-beaked Echidna[1]
Conservation status

Least Concern (IUCN 3.1)[2]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Subclass: Prototheria
Order: Monotremata
Family: Tachyglossidae
Genus: Tachyglossus
(Illiger, 1811)
Species: T. aculeatus
Binomial name
Tachyglossus aculeatus
(Shaw, 1792)
Subspecies
  • T. a. acanthion
  • T. a. aculeatus
  • T. a. lawesii
  • T. a. multiaculeatus
  • T. a. setosus
Range of Short-beaked Echidna

The Short-beaked Echidna (Tachyglossus aculeatus), also known as the Spiny Anteater because of its diet of ants and termites, is one of four living species of echidna and the only member of the genus Tachyglossus. The Short-beaked Echidna is covered in fur and spines and has a distinctive snout and a specialized tongue, which it uses to catch its prey at a great speed. Like the other extant monotremes, the Short-beaked Echidna lays eggs; the monotremes are the only group of mammals to do so.

The species is found throughout Australia, where it is the most widespread native mammal, and in coastal and highland regions of southwestern New Guinea, where it is known as the Mungwe in the Daribi and Chimbu languages.[3] It is not threatened with extinction, but human activities, such as hunting, habitat destruction, and the introduction of foreign predatory species and parasites, have reduced the distribution of the Short-beaked Echidna in Australia.

Contents

Taxonomy and naming

The Short-beaked Echidna was first described by George Shaw in 1792. He named the species Myrmecophaga aculeata, thinking it might be related to the South American anteater. Since Shaw first described the species, its name has undergone four revisions: from M. aculeata to Ornithorhynchus hystrix, Echidna hystrix, Echidna aculeata and, finally, Tachyglossus aculeatus.[4][5] The name Tachyglossus means "quick tongue", in reference to the speed with which the Echidna uses its tongue to catch ants and termites,[4] and aculeatus means "spiny" or "equipped with spines".[4]

The Short-beaked Echidna is the only member of its genus,[6] sharing the family Tachyglossidae with the extant species of the genus Zaglossus that occur in New Guinea.[7] Zaglossus species, which include the Western Long-beaked, Sir David's Long-beaked and Eastern Long-beaked Echidna,[8] are all significantly larger than T. aculeatus, and their diet consists mostly of worms and grubs rather than ants and termites.[9] Species of the Tachyglossidae are egg-laying mammals; together with the related family Ornithorhynchidae, they are the only extant monotremes in the world.[10]

There are five subspecies of the Short-beaked Echidna, each found in a different geographical location. The subspecies also differ from one another in their hairiness, spine length and width, and the size of the grooming claws on their hind feet.[11]

T. a. setosus, Barrington, Tasmania

The Short-beaked Echidna was commonly called the Spiny Anteater in older books, though this term has fallen out of fashion since the Echidna bears no relation to the true anteaters. It has a variety of names in the indigenous languages of the regions where it is found. The Noongar people from southwestern Western Australia call it the Nyingarn. In Central Australia southwest of Alice Springs, the Pitjantjatjara term is tjilkamata or tjirili, from the word tjiri for spike of Porcupine Grass (Triodia irritans). The word can also mean slowpoke.[13] In central Cape York Peninsula, it is called (minha) kekoywa in Pakanh, where minha is a qualifier meaning 'meat' or 'animal', (inh-)ekorak in Uw Oykangand and (inh-)egorag in Uw Olkola, where inh- is a qualifier meaning 'meat' or 'animal'.[14] In the highland regions of southwestern New Guinea it is known as the Mungwe in the Daribi and Chimbu languages.[3]

Description

Spines and fur of an echidna

Short-beaked Echidnas are typically 30 to 45 centimetres (12 to 18 in) in length, have a 75-millimetre (3 in) snout, and weigh between 2 and 5 kilograms (4.4 and 11 lb).[15] However, the Tasmanian subspecies, T. a. setosus, is smaller than its Australian mainland counterparts.[16] Because the neck is not externally visible, the head and body appear to merge together.[17] The earholes are on either side of the head,[17] with no external pinnae.[17] The eyes are small and at the base of the wedge-shaped snout. The nostrils and the mouth are at the distal end of the snout;[17] the mouth of the Short-beaked Echidna cannot open wider than 5 mm (0.2 in).[18] The body of the Short-beaked Echidna is, with the exception of the underside, face and legs, covered with cream-coloured spines. The spines, which may be up to 50 mm (2 in) long, are modified hairs,[19] mostly made of keratin.[20] Insulation is provided by fur between the spines, which ranges in colour from honey to a dark reddish-brown and even black; the underside and short tail are also covered in fur.[19] Colouration of the fur and spines varies with geographic location. The Echidna's fur may be infested with what is said to be the world's largest flea, Bradiopsylla echidnae, which is about 4 mm (0.16 in) long.[19]

The limbs of the Short-beaked Echidna are adapted for rapid digging: they are short and have strong claws.[19] The claws on the hind feet are elongated and curve backwards to enable cleaning and grooming between the spines. Like the Platypus, the Echidna has a low body temperature – between 30 and 32 °C (86 and 90 °F) – but, unlike the Platypus, which shows no evidence of torpor or hibernation, the body temperature of the Echidna may fall as low as 5 °C (41 °F).[21] The Echidna does not pant or sweat[22] and normally seeks shelter in hot conditions.[23] In autumn and winter the Echidna shows periods of torpor or deep hibernation.[24] Because of the low body temperature of the Short-beaked Echidna, it becomes sluggish in very hot and very cold weather. Like all monotremes, it has one orifice, known as the cloaca, for the passage of faeces, urine and reproductive products.[24] The male has internal testes, no external scrotum and a highly unusual penis with four knobs on the tip.[25] The gestating female develops a pouch on its underside, where it raises its young.[26]

A Short-beaked Echidna curled into a ball; the snout is visible on the right.

The musculature of the Short-beaked Echidna has a number of unusual aspects. The panniculus carnosus is an enormous muscle that is just beneath the skin and covers the entire body. By contraction of various parts of the panniculus carnosus, the Short-beaked Echidna can change shape, the most characteristic shape change being achieved by rolling itself into a ball when threatened, so protecting its belly and presenting a defensive array of sharp spines. It has one of the shortest spinal cords of any mammal, extending only as far as the thorax.[27]

The musculature of the face, jaw and tongue is specialised to allow the Echidna to feed. The tongue of the Short-beaked Echidna is the animal's sole means of catching prey, and can protrude up to 180 mm (7 in)) outside the snout.[15] The tongue is sticky because of the presence of glycoprotein-rich mucous, which both lubricates movement in and out of the snout and helps to catch ants and termites, which adhere to it. Protrusion of the tongue is achieved by contracting circular muscles that change the shape of the tongue and force it forwards and contracting two genioglossal muscles attached to the caudal end of the tongue and to the mandible. The protruded tongue is stiffened by the rapid flow of blood, allowing it to penetrate wood and soil. Retraction requires the contraction of two internal longitudinal muscles, known as the sternoglossi. When the tongue is retracted, the prey is caught on backward-facing keratinous "teeth", located along the roof of the buccal cavity, allowing the animal both to capture and grind food.[18] The tongue moves with great speed, and has been measured to move in and out of the snout 100 times a minute.[15][28]

Numerous physiological adaptations aid the lifestyle of the Short-beaked Echidna. Because the animal burrows, it can tolerate very high levels of carbon dioxide in inspired air, and will voluntarily remain in situations where carbon dioxide concentrations are high. Its ear is sensitive to low-frequency sound, which may be ideal for detecting sounds emitted by termites and ants underground. The leathery snout is covered in mechano- and thermoreceptors, which provide information about the surrounding environment.[29] The Short-beaked Echidna has a well-developed olfactory system, which may be used to detect mates and prey. It has a highly sensitive optic nerve, and has been shown to have visual discrimination and spatial memory comparable to those of a rat.[30] The brain and central nervous system of the Short-beaked Echidna have been extensively studied for evolutionary comparison with placental mammals. The Short-beaked Echidna has the largest prefrontal cortex relative to body size of any mammal, it shows rapid eye movement during sleep, and its brain has been shown to contain a claustrum similar to that of placental mammals, so linking this structure to their common ancestor.[31][32]

Ecology and behaviour

No systematic study of the ecology of the Short-beaked Echidna has been published; however, there have been studies of several aspects of their ecological behaviour. Short-beaked Echidnas live alone and, apart from the burrow created for rearing young, they have no fixed shelter or nest site. They do not have a home territory, but range over a wide area. Short-beaked Echidnas are typically active in the daytime; however, they are ill-equipped to deal with heat, because they have no sweat glands and do not pant. Therefore, in warm weather they change their pattern of activity, becoming crepuscular or nocturnal. They can tolerate cold temperatures, and hibernate during the winter in very cold regions.[33]

Short-beaked Echidnas can live anywhere that has a good supply of food. They locate food by smell, using sensors in the tip of their snout, and regularly feast on ants and termites.[34] They are powerful diggers, using their clawed front paws to dig out prey and create burrows for shelter. They may rapidly dig themselves into the ground if they cannot find cover when in danger.[19]

In Australia they are most common in forested areas where there are abundant termite-filled fallen logs. In agricultural areas, they are most likely to be found in uncleared scrub; they may be found in grassland, arid areas, and in the outer suburbs of the capital cities. Little is known about their distribution in New Guinea; they have been found in southern New Guinea between Merauke in the west and the Kelp Welsh River, east of Port Moresby, in the east, where they may be found in open woodland.[3]

Reproduction

The underside of a female Short-beaked Echidna; the pouch which carries eggs is towards the rear of the abdomen.

The solitary Short-beaked Echidna looks for a mate between May and September;[19] the precise timing of the mating season varies with geographic location. Both males and females give off a strong odour during the mating season. During courtship – observed for the first time in 1989 – males locate and pursue females. Trains of up to ten males may follow a single female in a courtship ritual that may last for up to four weeks; the duration of the courtship period varies with location.[15][35] In cooler parts of their range, such as Tasmania, females may mate within a few hours of arousal from hibernation.[36]

Before mating, the male smells the female, paying particular attention to the cloaca. The male is often observed to roll the female onto her side and then assume a similar position himself so that the two animals are abdomen to abdomen. Each side of the bilaterally symmetrical, rosette-like, four-headed penis [similar to that of reptiles] is used alternately, with the other half being shut down between ejaculations. Sperm bundles of around 100 each appear to confer increased sperm motility, which may provide the potential for sperm competition between males.[37] Each mating results in the production of a single egg, and females are known to mate only once during the breeding season; each mating is successful.[38]

Fertilisation occurs in the oviduct. Gestation takes between 21 and 28 days, during which time the female constructs a nursery burrow. Following the gestation period, a single rubbery-skinned egg[15] between 13 and 17 millimetres (0.5 and 0.7 in) in diameter[39] is laid directly into a small, backward-facing pouch that has developed on her abdomen. Ten days after it is laid, the egg hatches within the pouch.[19] The embryo develops an "egg tooth" during incubation, which it uses to tear open the egg; the tooth disappears soon after hatching.[40]

Hatchlings are about 1.5 centimetres (0.6 in) long and weigh between 0.3 and 0.4 gram (0.011 and 0.014 oz).[41] After hatching, young Echidnas are known as puggles. Hatchlings attach themselves to their mothers' milk areolae, specialised patches on the skin that secrete milk (monotremes lack nipples).[15][19] The way in which puggles imbibe the milk is not yet known, but they have been observed ingesting large amounts during each feeding period, and mothers may leave them unattended in the burrow for between five and ten days.[42] The principal components of the milk are fucosyllactose and saialyllactose; it has a high iron content, which gives it a pink colour.[43] Juveniles are eventually ejected from the pouch at around two to three months of age, because of the continuing growth in the length of their spines.[19] Suckling gradually decreases until juveniles are weaned at about six months of age. The duration of lactation is about 200 days,[15] and the young leave the burrow after 180 to 205 days.[44]

The age of sexual maturity is uncertain, but may be four to five years. A twelve-year field study, published in 2003, found that the Short-beaked Echidna reaches sexual maturity between five and twelve years of age, and that the frequency of reproduction varies from once every two years to once every six years.[41] The Short-beaked Echidna can live as long as forty-five years in the wild.[45]

Like its fellow monotreme the Platypus, the Short-beaked Echidna has a system of multiple sex chromosomes, in which males have four Y chromosomes and five X chromosomes. Male individuals appear to be X1Y1X2Y2X3Y3X4Y4X5 (figure[46]), while females are X1X1X2X2X3X3X4X4X5X5. Weak identity between chromosomes results in meiotic pairing that yields only two possible genotypes of sperm, X1X2X3X4X5 or Y1Y2Y3Y4, thus preserving this complex system.[46]

Conservation status

A Short-beaked Echidna on the move

The Short-beaked Echidna is common throughout most of temperate Australia and lowland New Guinea, and is not listed as endangered.[15][19] In Australia, the number of Short-beaked Echidnas has been less affected by land clearance than have some other species, since Short-beaked Echidnas do not require a specialized habitat beyond a good supply of ants and termites.[19] Despite their spines, they are preyed on by birds, the Tasmanian Devil,[19] dingoes,[15] cats and foxes.[47] They were eaten by indigenous Australians and the early European settlers of Australia.[19] In contrast, hunting and eating of the Echidna in New Guinea has increased over time and caused a decline in the population and distribution areas; it is now believed to have disappeared from highland areas. The killing of Echidnas were a taboo in traditional culture, but since the tribespeople have become increasingly Christianised by western missionaries, hunting has increased, and the animals have been more easily tracked down due to the use of dogs.[48] In Australia, the Echidna remains widespread across a wide range of conditions, including urban outskirts, coastal forests and dry inland areas. They are especially widespread in Tasmania and on Kangaroo Island.[49]

The most common threats to the animal in Australia are motor vehicles and habitat destruction, which have led to localized extinctions.[49] Infection with the introduced parasite Spirometra erinaceieuropaei is considered fatal for the Echidna.[50] The Wildlife Preservation Society of Queensland runs an Australia-wide survey called Echidna Watch to monitor the species in Australia.

Captive breeding is difficult, partly due to the relatively infrequent breeding cycle. In 2009, Perth Zoo managed to breed some captive Short-beaked Echidnas.[51] Up until 2006, only five zoos have managed to breed a captive Short-beaked Echidna, but no captive-bred young have survived to maturity.[52] Of these five institutions, only one in Australia—Sydney's Taronga Zoo— managed to breed echidnas, in 1977. The other four cases occurred in the northern hemisphere, two in the United States and the others in western Europe. In these cases, breeding occurred six months out of phase compared to Australia after the animals had adapted to northern hemisphere seasons.[52] The failure of captive breeding programs has conservation implications for the endangered species of echidna from the genus Zaglossus, and to a lesser extent for the Short-beaked Echidna.[52]

Cultural references

Short-beaked Echidnas feature in the animistic culture of indigenous Australians, including their visual arts and stories. The species was a totem for some groups, including the Noongar people from Western Australia. Many groups have myths about the animal; one myth explains that it was created when a group of hungry young men went hunting at night and stumbled across a wombat. They threw spears at the wombat, but lost sight of it in the darkness. The wombat adapted the spears for its own defence and turned into an Echidna.[53] Another story tells of a greedy man who kept food from his tribe; warriors speared him and he crawled away into the bushes, where he turned into an Echidna, the spears becoming his spines.

The Short-beaked Echidna is an iconic animal in contemporary Australia, notably appearing on the Australian five-cent piece (the smallest denomination),[54] and on a $200 commemorative coin released in 1992.[55] The Short-beaked Echidna has been included in several postal issues: it was one of four native species to appear on Australian postage stamps in 1974, when it was on the 25-cent stamp; it appeared on a 37-cent stamp in 1987, and in 1992 it was on the 35-cent stamp.[56] The anthropomorphic echidna Millie was a mascot for the 2000 Summer Olympics.[57]

See also

References

Cited references

  1. Groves, Colin P. (16 November 2005). "Order Monotremata (pp. 1-2)". In Wilson, Don E., and Reeder, DeeAnn M., eds. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press, 2 vols. (2142 pp.). p. 1. ISBN 978-0-8018-8221-0. OCLC 62265494. http://www.bucknell.edu/msw3/browse.asp?id=10300004. 
  2. Aplin, K., Dickman, C., Salas, L. & Helgen, K. (2008). Tachyglossus aculeatus. In: IUCN 2008. IUCN Red List of Threatened Species. Downloaded on 10 October 2008. Database entry includes justification for why this species is of least concern
  3. 3.0 3.1 3.2 Flannery, T. 1990. Mammals of New Guinea Robert Brown & Associates ISBN 1-86273-029-6
  4. 4.0 4.1 4.2 Augee, Gooden and Musser, p. 5.
  5. Iredale, T. and Troughton, E. 1934. A checklist of mammals recorded from Australia. Memoirs of the Australian Museum 6: i–xii, 1–122
  6. http://animaldiversity.ummz.umich.edu/site/accounts/classification/Tachyglossus.html
  7. Augee, Gooden and Musser, pp. 5–9.
  8. http://animaldiversity.ummz.umich.edu/site/accounts/classification/Zaglossus.html
  9. Augee, Gooden and Musser, pp. 7–10.
  10. Augee, Gooden and Musser, p. 10.
  11. 11.0 11.1 11.2 Augee, Gooden and Musser, pp. 6–7.
  12. Griffiths, M.E. 1978. The Biology of Monotremes. Academic Press : New York ISBN 0-12-303850-2
  13. Cliff Goddard (1992). Pitjantjatjara/Yankunytjatjara To English Dictionary (2 ed.). Alice Springs, Northern Territory: Institute for Aboriginal Development. p. 152. ISBN 0-949659-64-9. 
  14. Philip Hamilton (1997). "short-beaked echidna, Tachyglossus aculeatus". Australian Institute of Aboriginal and Torres Strait Islander Studies. http://www.geocities.com/Athens/Delphi/2970/echidna.htm. Retrieved 2007-06-08. 
  15. 15.0 15.1 15.2 15.3 15.4 15.5 15.6 15.7 15.8 Egerton, p. 38.
  16. Augee, Gooden and Musser, p. 6.
  17. 17.0 17.1 17.2 17.3 Augee, Gooden and Musser, p. 2.
  18. 18.0 18.1 Murray, P.F. 1981. A unique jaw mechanism in the echidna, Tachglossus aculeatus (Monotremata). Australian Journal of Zoology 29: 1–5
  19. 19.00 19.01 19.02 19.03 19.04 19.05 19.06 19.07 19.08 19.09 19.10 19.11 19.12 "DPIW - Short-beaked Echidna". Dpiw.tas.gov.au. 2009-08-10. http://www.dpiw.tas.gov.au/internnsf/WebPages/BHAN-5357K5?open. Retrieved 2010-05-29. 
  20. Augee, Gooden and Musser, p. 35.
  21. Dawson, T.J., Grant, T.R. and Fanning, D. 1979. Standard metabolism of monotremes and the evolution of homeothermy. Australian Journal of Zoology 27:511–515
  22. Augee, Gooden and Musser, pp. 90–91.
  23. Augee, Gooden and Musser, p. 91.
  24. 24.0 24.1 "Science & Nature - Wildfacts - Short-beaked echidna, common echidna, spiny anteater". BBC. 2008-07-25. http://www.bbc.co.uk/nature/wildfacts/factfiles/680.shtml. Retrieved 2010-05-29. 
  25. Augee, Gooden and Musser, p. 79.
  26. Augee, Gooden and Musser, p. 4.
  27. Hassiotis, M., Paxinos, G., and Ashwell, K.W.S. 2004. Anatomy of the central nervous system of the Australian echidna. Proceedings of the Linnean Society of New South Wales 125:287–300
  28. Doran, G.A. and Baggett, H. 1970. The vascular stiffening mechanism in the tongue of the echidna (Tachyglossus aculeatus). Anatomical Record 167: 197–204
  29. Iggo, A., McIntyre, A.K. & Proske, U. (1985). Responses of mechanoreceptors and thermoreceptors in skin of the snout of the echidna Tachyglossus aculeatus. Proceedings of the Royal Society of London B 223: 261–277
  30. Burke, D. 2002. Win-shift and win-stay learning in the short-beaked echidna (Tachyglossus aculeatus). Animal Cognition 5:79–84
  31. Nicol, S.C., et al. 2000. The echidna manifests typical characteristics of rapid eye movement sleep. Neuroscience Letters 283:49–52
  32. Ashwell, K.W.S., Hardman, C.D., and Paxinos, G. 2004. The claustrum is not missing from all monotreme brains. Brain Behaviour and Evolution 64:223–241 PMID 15319553
  33. Grigg, G.C., Beard, L.A., and Augee, M.L. 1989. Hibernation in a monotreme, the echidna (Tachyglossus aculeatus). Comparative Biochemistry and Physiology. A, Comparative Physiology 92:609–612 PMID 2566425
  34. "Short-beaked Echidna - Zoo Aquarium Association". Arazpa.org.au. http://www.arazpa.org.au/Short-beaked-Echidna/default.aspx. Retrieved 2010-05-29. 
  35. Rismiller, P.D., Seymour, R.S., 1991. The echidna. Scientific American 264, 96–103.
  36. Nicol, S.C., Andersen, N.A., and Jones, S.M., 2005. Seasonal variations in reproductive hormones of free-ranging echidnas (Tachyglossus aculeatus): interaction between reproduction and hibernation. General and Comparative Endocrinology 144: 204–210
  37. Johnston S.D., Smith B., Pyne M., Stenzel D., and Holt W.V. One-Sided Ejaculation of Echidna Sperm Bundles (Tachyglossus aculeatus). Am. Nat. 2007. Vol. 170, p. E000. [AN:42629 DOI:10.1086/522847]
  38. Rismiller, P.D. and McKelvey, M.W. 2000. Frequency of breeding recruitment in the Short-beaked Echidna, Tachyglossus aculeatus. Journal of Mammalogy 81:1–17
  39. Augee, Gooden and Musser, p. 84.
  40. Augee, Gooden and Musser, p. 85.
  41. 41.0 41.1 Rismiller, P.D. and McKelvey, M.W. 2003. Body mass, age and sexual maturity in short-beaked echidnas, Tachyglossus aculeatus. Comparative Biochemistry and Physiology. A, Molecular & Integrative Physiology 136:851–865
  42. Augee, Gooden and Musser, p. 86.
  43. Augee, Gooden and Musser, p. 87.
  44. Augee, Gooden and Musser, p. 88.
  45. "Parks Victoria Fact File". Parkweb.vic.gov.au. http://www.parkweb.vic.gov.au/education/factfiles/15.htm. Retrieved 2010-05-29. 
  46. 46.0 46.1 Rens, W., et al (2007). "The multiple sex chromosomes of platypus and echidna are not completely identical and several share homology with the avian Z". Genome Biology 8 (11): R243. doi:10.1186/gb-2007-8-11-r243. PMID 18021405. PMC 2258203. http://genomebiology.com/2007/8/11/R243. 
  47. Augee, Gooden and Musser, pp. 120–121.
  48. Augee, Gooden and Musser, p. 119.
  49. 49.0 49.1 Augee, Gooden and Musser, p. 120.
  50. Augee, Gooden and Musser, p. 121.
  51. "Baby echidnas make first appearance at Perth Zoo". Perthnow.com.au. 2009-11-04. http://www.perthnow.com.au/news/western-australia/baby-echidnas-make-first-appearance-at-perth-zoo/story-e6frg15c-1225794333104. Retrieved 2010-05-29. 
  52. 52.0 52.1 52.2 Augee, Gooden and Musser, p. 126.
  53. Robinson, R. 1966. Aboriginal Myths and Legends. Sun Books, Melbourne
  54. "Five cents". Ramint.gov.au. 1966-02-14. http://www.ramint.gov.au/designs/ram-designs/5c.cfm. Retrieved 2010-05-29. 
  55. http://heavymetalholdings.com.au/index_files/RAMGoldCoins.htm
  56. Australia Post. "Australian Wildlife (1992)". http://www.stamps.com.au/shop/reprints/australian-wildlife-1992. Retrieved 2010-05-08. 
  57. Rohrer, Finlo (2008-06-16). "How not to have an Olympic mascot nightmare". BBC News. http://news.bbc.co.uk/2/hi/uk_news/magazine/7456330.stm. Retrieved 2010-05-29. 

General references

  • Augee, M and Gooden, B. 1993. Echidnas of Australia and New Guinea. Australian National History Press, Sydney ISBN 978-0-86840-046-4
  • Augee, M.L. 1983. R. Strahan Ed. The Australian Museum Complete Book of Australian Mammals. pp. 8–9. Angus & Robertson ISBN 0-207-14454-0
  • Egerton, L. ed. 2005. Encyclopedia of Australian wildlife. Reader's Digest ISBN 1-876689-34-X
  • Griffiths, M. 1989. Tachyglossidae. pp. 407–435 in Fauna of Australia (D. W. Walton and B. J. Richardson, eds.). Mammalia, Canberra, Australian Capital Territory 1B:1–1227.

External links