Temporal range: Miocene–Holocene
|Western long-beaked echidna|
Echidnas //, sometimes known as spiny anteaters, belong to the family Tachyglossidae in the monotreme order of egg-laying mammals. The four extant species, together with the platypus, are the only surviving members of that order and are the only extant mammals that lay eggs. Their diet consists of ants and termites, but they are not closely related to the true anteaters of the Americas. They live in Australia and New Guinea.
The echidnas are named after Echidna, a creature from Greek mythology who was half-woman, half-snake, as the animal was perceived to have qualities of both mammals and reptiles. Aboriginal Australians, who regard the echidna as a food delicacy, rarely use the term echidna, instead almost invariably calling the monotreme and its meat, erroneously "porcupine". 
Echidnas are medium-sized, solitary mammals covered with coarse hair and spines. Superficially, they resemble the anteaters of South America and other spiny mammals such as hedgehogs and porcupines. They are usually black or brown in colour. There have been several reports of albino echidnas, their eyes pink and their spines white. They have elongated and slender snouts that function as both mouth and nose. Like the platypus, they are equipped with electrosensors, but while the platypus has 40,000 electroreceptors on its bill, the long-billed echidna has only 2,000, and the short-billed echidna, which lives in a drier environment, has no more than 400 located at the tip of its snout. They have very short, strong limbs with large claws, and are powerful diggers. Echidnas have tiny mouths and toothless jaws. The echidna feeds by tearing open soft logs, anthills and the like, and using its long, sticky tongue, which protrudes from its snout, to collect prey.
The short-beaked echidna's diet consists largely of ants and termites, while the Zaglossus species typically eats worms and insect larvae. They have no teeth, and break down their food by grinding it between the bottoms of their mouths and their tongues. The echidnas' ears are slits on the sides of their heads that are usually unseen due to the fact that they are blanketed by their spines. The external ear is called the "pinna", which is created by a large cartilaginous funnel, deep in the muscle.
Echidnas do not tolerate extreme temperatures; they use caves and rock crevasses to shelter from harsh weather conditions. Echidnas are found in forests and woodlands, hiding under vegetation, roots or piles of debris. They sometimes use the burrows of animals such as rabbits and wombats. Individual echidnas have large, mutually overlapping territories. 
Echidnas are capable swimmers. When swimming, they expose their snout and some of their spines. They are known to journey to water in order to groom and bathe themselves.
The tongues of long-beaked echidnas have sharp, tiny spines that help them capture their prey. Echidnas feces are 7 centimeters long and are cylindrical in shape; they are usually broken and unrounded, and composed largely of dirt and ant-hill material.
Echidnas and the platypus are the only egg-laying mammals, known as monotremes. The neocortex makes up half of the echidna's brain, compared to 80% of a human brain. Due to their low metabolism and accompanying stress resistance, echidnas are long-lived for their size; the longest recorded lifespan for a captive echidna is 50 years, with anecdotal accounts of wild individuals reaching 45 years. Contrary to previous research, the echidna does enter REM sleep, but only when the ambient temperature is around 25 °C (77 °F). At temperatures of 15 °C (59 °F) and 28 °C (82 °F), REM sleep is suppressed. The average lifespan of an echidna in the wild is estimated around 16 years. When fully grown a female can weigh up to 4.5 kilograms and a male can weigh up to 6 kilograms. You can determine the echidnas' sex by their size; males are 25% larger than females. As well, the reproductive organs differ even though both sexes have a single opening they use to urinate, release their faeces and use to mate.
|This section requires expansion. (November 2012)|
The female lays a single soft-shelled, leathery egg 22 days after mating, and deposits it directly into her pouch. While hatching, the baby echidna opens the leather shell with a reptile-like egg tooth  Hatching takes place after 10 days; the young echidna then sucks milk from the pores of the two milk patches (monotremes have no nipples) and remains in the pouch for 45 to 55 days, at which time it starts to develop spines. The mother digs a nursery burrow and deposits the young, returning every five days to suckle it until it is weaned at seven months.
Male echidnas have a four-headed penis. During mating, the heads on one side "shut down" and do not grow in size; the other two are used to release semen into the female's two-branched reproductive tract. Each time it has sex, it alternates heads in sets of two. When not in use, the penis is retracted inside a preputial sac in the cloaca. The male echidna's penis is 7 centimeters long when erect, and its shaft is covered with penile spines. These may be used to induce ovulation in the female. 
It is a challenge to study the echidna in their natural habitat and they show no interest in mating while in captivity. Therefore, no one has ever seen an echidna ejaculate. There have been previous attempts, trying to force the echidna to ejaculate through the use of electrically stimulated ejaculation in order to obtain semen samples but has only resulted in the penis swelling.
Breeding season begins in late June and extends through September. Males will form lines up to ten individuals long, the youngest echidna trailing last, that follow the female and attempt to mate. During a mating season an echidna may switch between lines. This is known as the "train" system. Two weeks after mating, a single fertilized egg, weighing 380 milligrams and being about 1.4 centimeters long, is implanted in a rear-facing pouch that has developed on the female, where it is held for ten days before hatching. The young echidna, called a puggle, is then held in the pouch for two to three months before being expelled. Puggles will stay within their mother's den for up to a year before leaving.
Echidnas are very timid animals. When they feel endangered they attempt to bury themselves or if exposed they will curl into a ball, both methods using their spines to shield them. Strong front arms allow echidnas to continue to dig themselves in whilst holding fast against a predator attempting to remove them from the hole. Although they have a way to protect themselves, the echidnas still face many dangers. Some predators include wild cats, foxes, domestic dogs and goannas. Snakes pose as a large threat to the echidna species because they slither into their burrows and prey on the young spineless puggles. Some precautions that can be taken include keeping the environment clean by picking up litter and causing less pollution, planting vegetation for echidnas to use as shelter, supervising pets, reporting hurt echidnas or just leaving them undisturbed. Merely grabbing them may cause stress, and not picking them up correctly may even result in injury.
The first divergence between oviparous (egg-laying) and viviparous (offspring develop internally) mammals is believed to have occurred during the Triassic period.  However, there is still some disagreement on this estimated time of divergence. Though most findings from genetics studies (especially those concerning nuclear genes) are in agreement with the paleontological findings, some results from other techniques and sources, like mitochondrial DNA, are in slight disagreement with findings from fossils.
Molecular clock data suggest echidnas split from platypuses between 19 and 48 million years ago, and that platypus-like fossils dating back to over 112.5 million years ago therefore represent basal forms, rather than close relatives of the modern platypus. This would imply that echidnas evolved from water-foraging ancestors that returned to living completely on the land, even though this put them in competition with marsupials. Further evidence of possible water-foraging ancestors can be found in some of the echidna's phenotypic traits as well. Traits such as: as aqua dynamic streamlining, dorsally projecting hind limbs acting as rudders, and locomotion founded on hypertrophied humeral long-axis rotation, which provides a very efficient swimming stroke. Consequently, oviparous reproduction in monotremes may have given them an advantage over marsupials, a view consistent with present ecological partitioning between the two groups. This advantage could as well be in part responsible for the observed associated adaptive radiation of echidnas and expansion of the niche space, which together contradict the fairly common assumption of halted morphological and molecular evolution that continues to be associated with monotremes. Furthermore, studies of mitochondrial DNA in platypuses have also found that monotremes and marsupials are most likely sister taxa. It also implies that any shared derived morphological traits between marsupials and placental mammals either occurred independently from one another or were lost in the lineage to monotremes.
Echidnas are classified into three genera. The genus Zaglossus includes three extant species and two species known only from fossils, while only one extant species from the genus Tachyglossus is known. The third genus, Megalibgwilia, is known only from fossils.
The three living Zaglossus species are endemic to New Guinea. They are rare and are hunted for food. They forage in leaf litter on the forest floor, eating earthworms and insects. The species are:
- Western long-beaked echidna (Z. bruijni), of the highland forests;
- Sir David's long-beaked echidna (Z. attenboroughi), described in 1961 and preferring a still higher habitat;
- Eastern long-beaked echidna (Z. bartoni), of which four distinct subspecies have been identified.
The two fossil species are:
The short-beaked echidna (Tachyglossus aculeatus) is found in southern, southeast and northeast New Guinea, and also occurs in almost all Australian environments, from the snow-clad Australian Alps to the deep deserts of the Outback, essentially anywhere ants and termites are available. It is smaller than the Zaglossus species, and it has longer hair. Despite the similar dietary habits and methods of consumption to those of an anteater, there is no evidence supporting the idea that echidna-like monotremes have been myrmecophagic (ant/termite-eating) since the Cretaceous. The fossil evidence of invertebrate-feeding bandicoots and rat-kangaroos, from around the time of the platypus–echidna divergence and pre-dating Tachyglossus, show evidence that echidnas expanded into new ecospace despite competition from marsupials.
The genus Megalibgwilia is known only from fossils:
Echidnas in popular culture
- The echidna appears on the reverse of the Australian 5-cent coin.
- Knuckles the Echidna is a red echidna featured in the video game series Sonic the Hedgehog.
- Millie, an echidna, was a mascot for the 2000 Summer Olympics.
- Frank Zappa wrote and performed the instrumental "Echidna's Arf (Of You)" on his 1974 LP Roxy & Elsewhere.
- The Echidna, an oracular guide, appears in the film Legend of the Guardians: The Owls of Ga'Hoole.
- Echidna Capital Management is a fictional front company for HYDRA in Marvel's Agents of S.H.I.E.L.D. (Season 2, Episode 18).
•The Echidna appears as a character in the app Crossy Road as part of the Australian update.
- "Short-Beaked Echidna, Tachyglossus aculeatus". Park & Wildlife Service Tasmania. Retrieved October 2012.
- Stewart, Doug (April–May 2003). "The Enigma of the Echidna". National Wildlife.
- Phillips, MJ; Bennett, TH; Lee, MS (October 2009). "Molecules, morphology, and ecology indicate a recent, amphibious ancestry for echidnas". Proc. Natl. Acad. Sci. U.S.A. 106 (40): 17089–94. doi:10.1073/pnas.0904649106. PMC 2761324. PMID 19805098.
- "echidna". Online Etymology Dictionary. Retrieved November 23, 2014.
- "echidna". Macquarie Dictionary. Retrieved July 7, 2015.
- Augee, Michael; Gooden, Brett; Musser, Anne (2006). Echidna : extraordinary egg-laying mammal (PDF) (2nd ed.). CSIRO. p. 3. ISBN 978-0-643-09204-4.
- "Electroreception in fish, amphibians and monotremes". Map of Life. 7 July 2010.
- "Zaglossus bruijni". AnimalInfo.org.
- Carritt, Rachel. "Echidnas: Helping them in the wild" (PDF). NSW National Parks and Wildlife Service. Retrieved 13 April 2013.
- "Short-beaked Echidna". Department of Primary Industries, Parks, Water, and Environment. Retrieved 13 April 2013.
- Gill, Victoria (19 November 2012). "Are these animals too 'ugly' to be saved?". BBC News.
- Dunbar, R.I.M. (1993). "Coevolution of neocortical size, group size and language in humans". Behavioural and Brain Sciences 16 (4): 681–735. doi:10.1017/S0140525X00032325.
- Dunbar, R.I.M. "The Social Brain Hypothesis" (PDF). University of Colorado at Boulder, Department of Psychology and Neuroscience. Retrieved 4 January 2014.
- Cason, M. (2009). "Tachyglossus aculeatus". Animal Diversity. Retrieved October 2012.
- Nicol, SC; Andersen, NA; Phillips, NH; Berger, BJ (March 2000). "The echidna manifests typical characteristics of rapid eye movement sleep". Neurosci. Lett. 283 (1): 49–52. doi:10.1016/S0304-3940(00)00922-8. PMID 10729631.
- Griffiths, Mervyn (1978). The biology of the monotremes. New York: Academic Press. ISBN 0123038502.
- O'Neil, Dennis. "Echidna Reproduction" 12 February 2011. Retrieved on 17 June 2015.
- "Short-beaked echidna (Tachyglossus aculeatus)". Arkive.org.
- Shultz, N. (26 October 2007). "Exhibitionist spiny anteater reveals bizarre penis". New Scientist. Retrieved 27 October 2006.
- Larry Vogelnest; Rupert Woods (18 August 2008). Medicine of Australian Mammals. Csiro Publishing. ISBN 978-0-643-09928-9. Retrieved 15 March 2013.
- Hayssen, V.D.; Van Tienhoven, A. (1993). "Order Monotremata, Family Tachyglossidae". Asdell's Patterns of Mammalian Reproduction: A Compendium of Species-specific Data. Cornell University Press. pp. 8–9. ISBN 0-8014-1753-8.
- Rowe T, Rich TH, Vickers-Rich P, Springer M, Woodburne MO (2008). "The oldest platypus and its bearing on divergence timing of the platypus and echidna clades". Proc. Natl. Acad. Sci. U.S.A. 105 (4): 1238–42. doi:10.1073/pnas.0706385105. PMC 2234122. PMID 18216270.
- Musser AM (2003). "Review of the monotreme fossil record and comparison of palaeontological and molecular data". Comp. Biochem. Physiol., Part A Mol. Integr. Physiol. 136 (4): 927–42. doi:10.1016/s1095-6433(03)00275-7. PMID 14667856.
- Janke A, Xu X, Arnason U (1997). "The complete mitochondrial genome of the wallaroo (Macropus robustus) and the phylogenetic relationship among Monotremata, Marsupialia, and Eutheria". Proc. Natl. Acad. Sci. U.S.A. 94 (4): 1276–81. doi:10.1073/pnas.94.4.1276. PMC 19781. PMID 9037043.
- Flannery, T.F.; Groves, C.P. (1998). "A revision of the genus Zaglossus (Monotremata, Tachyglossidae), with description of new species and subspecies". Mammalia 62 (3): 367–396. doi:10.1515/mamm.19188.8.131.527.
- Phillips, Matthew; Bennett, T.; Lee, Michael (2010). "Reply to Camens: How recently did modern monotremes diversify?". Proc. Natl. Acad. Sci. U.S.A. 107 (4): E13. doi:10.1073/pnas.0913152107.
|Wikimedia Commons has media related to Echidna.|
|Wikispecies has information related to: Tachyglossidae|
- Stewart, Doug (April–May 2003). "The Enigma of the Echidna". National Wildlife. Retrieved October 2005.
- Parker, J. (1 June 2000). "Echidna Love Trains". ABC Science. Australian Broadcasting Corporation.
- Rismiller, Peggy (2005). "Echidna research, Kangaroo island". Pelican Lagoon Research & Wildlife Centre.
- "Tachyglossidae". NCBI Taxonomy Browser. 9259.