Platypus: Difference between revisions

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The platypus (Ornithorhynchus anatinus), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative of its family Ornithorhynchidae and genus Ornithorhynchus, though a number of related species appear in the fossil record. Together with the four species of echidna, it is one of the five extant species of monotremes, mammals that lay eggs instead of giving birth to live young. Like other monotremes, the platypus has a sense of electrolocation, which it uses to detect prey in water while its eyes, ears and nostrils are closed. It is one of the few species of venomous mammals, as the male platypus has a spur on each hind foot that delivers an extremely painful venom.

The unusual appearance of this egg-laying, duck-billed, beaver-tailed mammal at first baffled European naturalists. In 1799, the first scientists to examine a preserved platypus body judged it a fake made of several animals sewn together. The unique features of the platypus make it important in the study of evolutionary biology, and a recognisable and iconic symbol of Australia. It is culturally significant to several Aboriginal peoples, who also used to hunt it for food, and has appeared on stamps and currency.

The platypus was hunted for its fur, but it has been a legally protected species in all states where it occurs since 1912. Captive breeding programs have had slight success, and it is vulnerable to pollution, bycatching and climate change. It is classified as a near-threatened species by the IUCN, but a November 2020 report has recommended that it be upgraded to threatened species under the federal EPBC Act, due to habitat destruction and declining numbers in all states.

Taxonomy and naming

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Australian Aboriginal people have referred to the platypus in various ways depending on Australian indigenous languages and dialects. Among the names found: boondaburra, mallingong, tambreet, watjarang^[1] (names in Yass, Murrumbidgee, and Tumut),^[2] tohunbuck (region of Goomburra, Darling Downs),^[2] dulaiwarrung^[3] or dulai warrung (Woiwurrung language, Wurundjeri, Victoria),^[4] djanbang (Bundjalung, Queensland),^[4] djumulung (Yuin language, Yuin, New South Wales),^[4] maluŋgaŋ (ngunnawal language, Ngunnawal, Australian Capital Territory),^[4] biladurang, wamul, dyiimalung, oornie, dungidany (Wiradjuri language, Wiradjuri, Vic, NSW),^[4] oonah,^[5] etc. The name chosen and approved in Palawa kani (reconstructed Tasmanian language) is larila.^[6]

When the platypus was first encountered by Europeans in 1798, a pelt and sketch were sent back to Great Britain by Captain John Hunter, the second Governor of New South Wales.^[7] British scientists' initial hunch was that the attributes were a hoax.^[8] George Shaw, who produced the first description of the animal in the Naturalist's Miscellany in 1799, stated it was impossible not to entertain doubts as to its genuine nature,^[9] and Robert Knox believed—because it arrived in England via the Indian Ocean—that it might have been created by Chinese sailors.^[8] It was thought somebody had sewn a duck's beak onto the body of a beaver-like animal. Shaw used a pair of scissors to check for stitches.^[10]^[9]

The common name "platypus" means 'flat-foot', deriving from the Greek word Template:Transliteration (Template:Wikt-lang),^[11] from Template:Transliteration (Template:Wikt-lang 'broad, wide, flat')^[12]Template:Efn and Template:Transliteration (Template:Wikt-lang 'foot').^[13]^[14] Shaw initially assigned the species the Linnaean name Platypus anatinus when he described it,^[9]^[7] but the genus term was quickly discovered to already be in use as the name of a beetle genus Platypus.^[7] It was independently described as Ornithorhynchus paradoxus by Johann Blumenbach in 1800 (from a specimen given to him by Sir Joseph Banks)^[15] and following the rules of priority of nomenclature, it was later officially recognised as Ornithorhynchus anatinus.^[16]

Various dictionaries list "platypuses" or simply "platypus" as the plural. Alternatively, the term "platypi" is also used for the plural, although this is a form of pseudo-Latin; going by the word's Greek roots the plural would be "platypodes".^[10] Early European settlers called it by many names, such as "watermole", "duckbill", and "duckmole".^[7] Occasionally it is specifically called the "duck-billed platypus".^[8] There is no official term for platypus young, but the term "platypup" sees unofficial use, as does "puggle".^[17]^[18]

The scientific name Ornithorhynchus anatinus literally means 'duck-like bird-snout',^[9] deriving its genus name from the Greek root Template:Transliteration (Template:Wikt-lang ornith or Script error: No such module "Lang". órnīs 'bird')^[19] and the word Template:Transliteration (Template:Wikt-lang 'snout', 'beak').^[20] Its species name is derived from Latin Template:Wikt-lang ('duck-like') from Template:Wikt-lang 'duck'.^[9]^[21] The platypus is the sole living representative or monotypic taxon of its family (Ornithorhynchidae).^[22]

A book for children published in Germany in 1798

A book for children published in Germany in 1798
Illustration from the first scientific description in 1799

Illustration from the first scientific description in 1799
Colour print from 1863

Colour print from 1863

Description

File:Platypus skeleton Pengo.jpg

Platypus skeleton

Most of the platypus' small streamlined body is covered with short, dense, brown, fur that traps a layer of insulating air to keep the animal warm, both in and out of water.^[10]^[16]^[23]Template:Rp The fur coat is waterproof and consists of flattened guard hairs and curvy underfur hairs.^[16]^[23]Template:Rp It is one of the most densely furred mammals, behind only otters.^[24] It is also biofluorescent and glows cyan and green when under ultraviolet light; this may serve to camouflage it in low lighting from UV-sensitive predators.^[25] The duck-like bill consists of a long snout and lower jaw which is covered in soft skin. The nostrils are located near the tip of the snout's dorsal surface, while the eyes and ears are just behind the snout in a groove which closes underwater.^[16] It has cheek pouches for storing food. The platypus's wide, flat tail is compared to a beaver's but is furry rather than scaly;^[26] it stores fat reserves and can act as a rudder during swimming.^[16]^[23]Template:Rp The legs are short and have a sprawling stance. Webbing is more significant on the front feet. While walking on land, the feet are folded up in knuckle-walking to protect the webbing.^[23]Template:Rp^[27]

The platypus has an interclavicle in the shoulder girdle, a trait which they share in common with reptiles.^[23]Template:Rp As in many other aquatic and semiaquatic vertebrates, the bones show osteosclerosis, increasing their density to reduce buoyancy.^[28] Adult platypuses lack teeth and instead have heavily keratinised food-grinding pads.^[16] Young platypuses have one premolar tooth and two molars on each maxillae, and three molars on the dentaries. The first upper and third lower cheek teeth have only one major cusp, while the rest have two.^[29] They lose their teeth around the time they leave their natal burrow.^[16]

Male platypuses have an average length of Template:Convert and weight of Template:Convert, while females are smaller with an average length of Template:Convert and weight of Template:Convert.^[16] The species follows Bergmann's rule, with individuals being larger the farther south they are, due to colder climates; there are local variations, however.^[30] The platypus has an average body temperature of about Template:Convert, lower than the Template:Convert typical of placental mammals.^[31] Research suggests this has been a gradual adaptation to harsh environmental conditions among the few marginal surviving monotreme species, rather than a general characteristic of past monotremes.^[32]^[33]

The platypus has a single opening, called a cloaca, for both the reproductive and waste systems.^[23]Template:Rp The male platypus has penile spines and an asymmetrical glans penis that is larger on the left side.^[34] The female has two ovaries, with the right one being non-functional,^[34]^[35] and teats are absent.^[16]

Diving

Diving
Surfacing

Surfacing
File:Duck-billed platypus (Ornithorhynchus anatinus) Scottsdale 4.jpg

Senses

File:Platypus electrolocation.svg

The platypus has secondarily acquired electroreception. Its receptors are arranged in stripes on its bill, giving it high sensitivity to the sides and below; it makes quick turns of its head as it swims to detect prey.^[36]

Monotremes are the only mammals (apart from the Guiana dolphin)^[37] known to have a sense of electroreception.^[38]^[36] The playtpus relies on electrolocation when feeding, as the eyes, ears, and nose are closed while underwater.^[39]^[40] Digging in the bottom of streams with its bill, its electroreceptors detect tiny electric currents generated by the muscular contractions of its prey.^[36] Experiments have shown the platypus will even react to an "artificial shrimp" if a small electric current is passed through it.^[41]

The 40,000 electroreceptors are arranged in rows in the skin of the bill from front to back, while mechanoreceptors for touch are uniformly distributed across the bill. The electrosensory area of the cerebral cortex is in the tactile somatosensory area, and some cortical cells receive input from both electroreceptors and mechanoreceptors, suggesting the platypus feels electric fields as touches. These receptors in the bill dominate the somatotopic map of the platypus brain, in the same way human hands dominate the Penfield homunculus map.^[24]^[42] The platypus can feel the direction of an electric source, perhaps by comparing differences in signal strength across the array of electroreceptors, enhanced by the characteristic side-to-side motion of the animal's head while hunting. It may also be able to determine the distance of moving prey via the timing difference between electrical and mechanical pressure sensations.^[36] Monotreme electrolocation for hunting in murky waters may be tied to their tooth loss. The extinct Obdurodon was electroreceptive, but unlike the modern platypus it foraged in open water.^[43]

The eyes of the platypus have basal traits also found in lungfish and amphibians, such as scleral cartilage, double cones, and droplets.^[44] The platypus's eyes are small and shut under water, though several features indicate its ancestors relied on vision. As with other aquatic mammals, the eye has a flattened cornea and surrounding lens, while the posterior surface of the lens is sharply inclined. A temporal (ear side) concentration of retinal ganglion cells, important for binocular vision, indicates a vestigial role in predation, though the actual visual acuity is insufficient for such activities. Limited acuity is matched by low cortical magnification, a small lateral geniculate nucleus, and a large optic tectum, suggesting that the visual midbrain plays a more important role than the visual cortex, as in some rodents. These features suggest that the platypus has adapted to an aquatic and nocturnal lifestyle, developing its electrosensory system at the cost of its visual system. This contrasts with the small number of electroreceptors in the short-beaked echidna, which dwells in dry environments, while the long-beaked echidna, which lives in wetter habitats is intermediate between the other two monotremes.^[24]

The ears of the platypus are adapted for hearing while out of water.^[24] As in all true mammals, it has three middle ear bones, though the cochlea lacks spirals,^[16] but is described as "well organised". Within the cochlea, there are rows of inner and outer hair cells. As in placental mammals, the outer hair cells of the platypus are adapted for hearing high frequencies, suggesting it is an ancestral mammalian trait. However it also possesses more rows of inner hair cells.^[24] The olfactory (smelling) systems of the platypus and the echidna independently evolved from an ancestor with less advanced smelling. The main olfactory bulb of the platypus lacks the complex layers of the echidna, while both the piriform cortex and flaps (lamella) are simpler. Monotremes differ from placental mammals in that their mitral cells are distributed throughout the outer plexiform layer of the olfactory bulb rather than packed as a monolayer.^[45]

Venom

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File:Platypus spur.JPG

The calcaneus spur on the male's hind limb is used to inject venom.

While both male and female platypuses are born with back ankle spurs, only the males retain them into adulthood.^[26] Similar spurs are found on many archaic mammal groups, indicating that this was an ancient general characteristic among mammals.^[46] The spurs of the male inject venom, which is powerful enough to inflict pain in humans.^[26] Starting from the wounded area, the affect limb develops edema (swelling via fluid buildup) which can lead to an excruciating hyperalgesia (heightened sensitivity to pain) that can last as long as months.^[47]

The venom is composed largely of defensin-like proteins (DLPs) produced by the immune system, some of which are unique to the species.^[48] It is produced in kidney-shaped alveolar glands located in each of the thighs of the hind limbs and connected to the spur.^[16] The venomous spurs of male platypuses serve as weapons in battles with other males for breeding.^[26]^[47]

Distribution and habitat

The platypus is native to the freshwaters of eastern Australia, from Queenland to Tasmania (including King Island but not the Furneaux Group).^[26]^[49] It was believed to be extinct on the South Australian mainland, with the last sighting recorded at Renmark in 1975.^[50] Platypuses were captively bred at Warrawong Sanctuary in 1990-91.^[51] In October 2020 a nesting platypus was filmed in the wild after the previously abandoned Sanctuary reopened.^[52] There is a population on Kangaroo Island^[53] introduced in the 1920s, said to stand at 150 individuals in the Rocky River region of Flinders Chase National Park. In the 2019–20 Australian bushfire season, large portions of the island burned, decimating wildlife. However, SA Department for Environment and Water recovery teams worked to restore their habitat, with a number of sightings reported by April 2020.^[54] The platypus has almost disappeared from the Murray–Darling Basin, possibly due to poor water management.^[55] Platypuses can be found in a variety of freshwater habitats including rivers, streams, lakes and lagoon-like pools. The surrounding terrestrial environment includes tropical rainforests and colder alpine areas.^[26]

Ecology and behaviour

File:Ornithorhynchus anatinus -Sydney Aquarium, Sydney, Australia -swimming-6a.ogv

Swimming underwater at Sydney Aquarium, Australia

The platypus is semiaquatic and requires permanent freshwater habitat.^[26] Its swimming style is unique among mammals, propelling itself by alternating strokes of each front foot, while the webbed hind feet and tail are used for steering.^[56] It can maintain its relatively low body temperature when feeding in colder depths of below Template:Convert.^[16] In one study, dives lasted on average thirty-five seconds, with surfacing intervals averaging thirteen seconds.^[57] The species is mainly nocturnal but is also active at dusk during the summer and daytime during winter. A platypus may spend half the day in water and then retreat into its burrow, which is constructed by digging into the bank. These vary between simple resting banks and complex nesting/breeding burrows.^[26]^[16]^[58] It may have a range of up to Template:Convert, with a male's home range overlapping those of three or four females.^[59] Platypuses are not very vocal; they have been recorded to growl when disturbed and squeak when feeling pain.^[24]

The platypus is a carnivore and forages by probing along the bottom. It feeds on insect larvae, annelid worms, shrimp, crayfish, bivalves, tadpoles and fish eggs. It stores food in its cheek pouches for later consumption.^[26]^[58] In captivity, platypuses have survived up to thirty years, and wild specimens have been recaptured at twenty-four years old.^[60] They are preyed upon by eels, snakes, goannas, birds of prey, crocodiles, feral cats and foxes.^[23]Template:Rp Parasites and viruses also affect their mortality, though platypuses appear to have a high tolerance for them. Externally, platypuses may carry fleas, mites, and ticks, the latter being more prominent in young.^[26] The platypus is a common host for the tick species Ixodes ornithorhynchi.^[61] Internally, the platypus may host protozoans, trematodes, and nematodes.^[26] The fungus Mucor amphibiorum has been reported in Tasmanian platypuses, which causes the disease mucormycosis, symptoms of which include skin lesions and ulcers along the body.^[62]

Reproduction

File:Ornithorhynchus anatinus - nest with eggs - MUSE.JPG

Platypus's nest with eggs (replica)

Platypuses are seasonal breeders, with the more southerly populations breeding later in the year than those further north; those in New South Wales mate during fall and winter.^[58] Research has found both resident and transient platypuses, and suggest a polygynous mating system.^[63] Females are believed to become sexually mature at two years of age and continue to breed beyond nine years.^[63] During copulation, the male swims behind the female and grabs the tail with his bill, positioning his own tail under her from the side, then grips her neck or shoulder.^[64]Template:Rp The female retreats into a nesting/breeding burrow made of grass, which can be Template:Convert long, to lay eggs and raise young.^[26]^[16]^[65] Early European naturalists did not believe that the female platypus lays eggs, but this was confirmed by William Hay Caldwell in 1884.^[16]^[48]

Most mammal zygotes go through holoblastic cleavage, splitting into several divisible daughter cells. However, monotremes such as the platypus, along with reptiles and birds, undergo meroblastic cleavage, in which the ovum does not split completely. The cells at the edge of the yolk remain continuous with the egg's cytoplasm, allowing the yolk and embryo to exchange waste and nutrients with the egg through the cytoplasm.^[66]^[67] The female normally lays two small, leathery eggs about Template:Convert long. The eggs develop in utero for about twenty-eight days, followed by a ten day external incubation.^[35] The female curls around the incubating eggs,^[23]Template:Rp as the embryo continues to develop.^[67] Newly hatched platypuses are vulnerable; blind and hairless, they are fed by the mother's milk, which is thicker than in placental mammals and provides all the requirements for growth and development.^[16]^[68]^[69] With no teats, the milk is released through pores in the skin from which the young lap it up in her fur.^[70] The offspring are milk-fed for around four months; a minority of young are weaned after they exit the burrow but usually within five days.^[65] Hatched young also have a remnant yolk sac which disappears within four days.^[69]

During incubation and weaning, the mother initially leaves the burrow only for short periods to forage. She leaves behind her a number of thin soil plugs along the length of the burrow; pushing past these on her return squeezes water from her fur and allows the burrow to remain dry.^[70]^[65] The female spends less time with her offspring after five weeks, who emerge from the burrow around four months.^[71] By then, they are fully covered in fur and may be around 67% the weight of an adult and 80% the length.^[16] Juvenile males disperse further than females.^[72] They are fully grown at around two years.^[64]Template:Rp

Evolution

Template:Cladogram

In separate publications in 1934, 1947, and 1951, William King Gregory theorised that placental mammals and marsupials may have diverged earlier, and a subsequent branching divided the monotremes and marsupials. Later research and fossil discoveries have suggested this is incorrect.^[73]^[74] Modern monotremes are the survivors of an early branching of the mammal tree, and a later branching is thought to have led to the marsupial and placental groups.^[73]^[75] Both molecular clock and fossil dating suggest that the platypus split from echidnas around 19–48Script error: No such module "String".million years ago.^[76]

File:Steropodon BW.jpg

Reconstruction of ancient platypus relative Steropodon

The oldest discovered fossil of the modern platypus dates back to about 100,000 years ago during the Quaternary period, though a limb bone of Ornithorhynchus is known from Pliocene-epoch strata.^[77] The extinct monotremes Teinolophos, Steropodon and Kollikodon from the Cretaceous period are considered to be basal to the platypus and echidnas.^[78] The remains of Steropodon were discovered in New South Wales, composed of an opalised lower jawbone with three molar teeth (whereas the adult contemporary platypus is toothless). The molar teeth were initially thought to be tribosphenic, which would have supported a variation of Gregory's theory, but later research has suggested that, while they have three cusps, they evolved under a separate process.^[79] The fossil jaw of Teinolophos is elongated but unlike the modern platypus (and echidnas), lacks a beak.^[80]

In 2024, Late Cretaceous (Cenomanian)-aged fossil specimens of early platypus relatives were recovered from the same rocks as Steropodon, including the basal Opalios and the more derived Dharragarra, the latter of which may be the oldest member of the platypus family Ornithorhynchidae, as it retains the same dental formula found in Cenozoic platypus relatives.^[81] Monotrematum and Patagorhynchus, also fossil relatives of the platypus, are known from the latest Cretaceous (Maastrichtian) and the mid-Paleocene of Argentina, indicating that some monotremes managed to colonize South America from Australia when the two continents were connected via Antarctica. These are also considered potential members of the platypus Ornithorhynchidae.^[79]^[82]^[81] The closest fossil relative of the platypus is Obdurodon, known from the late Oligocene to the Miocene of Australia. It closely resembled the modern platypus, aside from the presence of molar teeth.^[81] A fossilised tooth of the giant platypus Obdurodon tharalkooschild was dated 5–15Script error: No such module "String".million years ago. Judging by the tooth, the animal measured 1.3 metres long, making it the largest platypus known.^[83]^[84]

The loss of teeth in the modern platypus has long been enigmatic, as a distinctive lower molar tooth row was present in its lineage for over 95 million years. Even its closest relative, Obdurodon, which otherwise closely resembles the platypus, retained this tooth row. More recent studies indicate that this tooth loss was geologically a very recent event, occurring only around the Plio-Pleistocene about 2.5 million years ago, when the rakali, a large semiaquatic rodent, colonized Australia from New Guinea. The platypus, which previously fed on a wide array of hard and soft-bodied prey, was outcompeted by the rakali for hard-bodied prey such as crayfish and mussels. This competition may have selected for the loss of teeth in the platypus and their replacement by horny pads, as a way of specializing for softer-bodied prey, over which the rakali did not compete.^[81]

Genome

Because of the early divergence from the therian mammals and the low numbers of extant monotreme species, the platypus is a frequent subject of research in evolutionary biology. In 2004, researchers at the Australian National University discovered that the platypus has ten sex chromosomes, compared with two (XY) in most other mammals. These ten chromosomes form five unique pairs of XY in males and XX in females, i.e. males are X₁Y₁X₂Y₂X₃Y₃X₄Y₄X₅Y₅. One of the X chromosomes of the platypus has close homology to the bird Z chromosome.^[85] The platypus genome also has both reptilian and mammalian genes associated with egg fertilisation.^[40]^[86] Though the platypus lacks the mammalian sex-determining gene SRY, a study found that the mechanism of sex determination is the AMH gene on the oldest Y chromosome.^[87]^[88] A draft version of the platypus genome sequence was published in Nature in May 2008, revealing both reptilian and mammalian elements, as well as two genes previously found only in non-mammalian vertebrates. More than 80% of the platypus's genes exist in other amniotes whose genomes have been compared.^[40] An updated genome, the most complete on record, was published in 2021, together with the genome of the short-beaked echidna.^[89]

Conservation

The International Union for Conservation of Nature categorised the platypus as "near threatened" in 2016,^[90] based on estimates that numbers had fallen by about thirty percent on average since European settlement. Other biologists have shown concern that the estimates of the 2016 baseline numbers could be wrong, and numbers may have been reduced by as much as fifty percent.^[91] The species was hunted for its fur until the early years of the 20th century. Although the species gained legal protections beginning in Victoria in 1890^[92] and throughout Australia by 1912,^[93] it continues to drown in the nets of inland fisheries.^[94] The use of "opera house traps" by recreational fishers for catching yabbies is banned in the ACT, South Australia, Tasmania and Victoria, and restricted in NSW and Queensland, due to the traps drowning non-targeted species including platypuses.^[95] The platypus was listed as endangered is South Australia, under the National Parks and Wildlife Act 1972. In November 2020 a recommendation was made to list the platypus as a vulnerable species across all states.^[96] In January 2021, Victoria officially adopted the vulnerable species designation, under the state's Flora and Fauna Guarantee Act 1988.^[97] The platypus is not covered under the federal EPBC Act.^[91]^[98]

The platypus continues to be adversely affected by habitat disruption caused by dams, pollution, urban expansion, and urban runoff.^[99] Droughts and the demands for water for human use are also considered threats.^[90] In January 2020, researchers from the University of New South Wales presented evidence that the platypus is at risk of extinction, due to factors such as water extraction, land clearing, climate change, and invasive species. The study predicted that, considering current threats, the animals' abundance would drop by 47–66% and metapopulation occupancy by 22–32% over fifty years, causing "extinction of local populations across about 40% of the range". Using climate change projections to 2070, reduced habitat due to drought would lead to 51–73% lower abundance and 36–56% lower metapopulation occupancy after at least fifty years. These predictions suggested that the species would fall under the "Vulnerable" classification. The authors stressed the need for national conservation efforts to ensure healthy platypus habitat, which may include conducting more surveys and tracking trends, as well as better river management while reducing threats.^[100]

A November 2020 report by scientists from the University of New South Wales, funded by a research grant from the Australian Conservation Foundation in collaboration with the World Wildlife Fund Australia and the Humane Society International Australia revealed that over the past thirty years platypus habitat in Australia has dropped by 22%, and supported listing the platypus as a threatened species under the EPBC Act, as the declines have been mostly in the Murray–Darling basin and NSW in general.^[96]

Sanctuaries and captivity

Few platypuses have been successfully raised and bred by humans. One notable example is at Healesville Sanctuary in Victoria under David Fleay, where breeding was successful in 1943.^[101]^[102]^[103] More platypuses were successfully bred and raised in 1998 and again in 2000 using a stream tank.^[103] Between 2008 and 2012, platypuses were bred regularly at Healesville,^[104] including twins.^[105] In 1990-91, there was successful breeding of platypuses at Warrawong Sanctuary.^[51] Taronga Zoo in Sydney bred twins in 2003,^[103] and the facility has since bred more platypuses to be released into the wild in NSW.^[106] Template:As of, the only platypuses in captivity outside of Australia are in the San Diego Zoo Safari Park in the U.S. state of California.^[107]^[108] Three platypuses were given to the Bronx Zoo in 1947, two females and a male. One of the females; Penelope had a false proto-pregnancy.^[109]

Human interactions

Usage

Aboriginal Australians hunted and ate platypuses, particularly for their fatty nutritious tails, while, after colonisation, Europeans killed them for fur from the late 19th century until 1912, when it was prohibited by law. In addition, European researchers captured and killed platypus or removed their eggs, partly in order to increase scientific knowledge, but also to gain prestige and outcompete rivals from different countries.^[96] During the Second World War, in spite of an export ban,^[110] Australia gave live platypuses as diplomatic gifts to Allied nations as part of an initiative to increase military assistance.^[111] One of which was intended as a gift to Winston Churchill died from neglect while en route.^[110]

Cultural references

The platypus has been a subject in the Dreamtime stories of Aboriginal Australians, some of whom believed the animal was a hybrid of a duck and a water rat.^[112]Template:Rp Aboriginals from the upper Darling River region have a story of a large water-rat called Biggoon who kidnaps a duck what wandered too far from its tribe. After managing to escape, she returned and laid two eggs which hatched the first platypuses. They were all exiled and went to live in the mountains. In another story from the upper Darling, the major animal groups, the land animals, water animals and birds, all competed for the platypus to join their respective groups, but the platypus ultimately decided to not join any of them, feeling that he did not need to be part of a group to be special,^[112]Template:Rp and wished to remain friends with all of those groups.^[96]

The platypus is also featured as a totem for some Aboriginal peoples, which is to them "a natural object, plant or animal that is inherited by members of a clan or family as their spiritual emblem", and the animal holds special meaning for the Wadi Wadi people at the Murray River. Because of their cultural significance and importance in connection to country, the platypus is protected and conserved by these Indigenous peoples.^[96] The platypus has often represented Australia's cultural identity and its image has also been used for stamps and currency and as a mascot in the 2000 Summer Olympics in Sydney.^[96]

Platypus fur cape made in 1890, now in the National Gallery of Victoria

Platypus fur cape made in 1890, now in the National Gallery of Victoria
Early 20th-century matchbox label

Early 20th-century matchbox label
9d postage stamp from 1937

9d postage stamp from 1937

Footnotes

Template:Notelist

Citations

Template:Reflist

External links

Biodiversity Heritage Library bibliography for Ornithorhynchus anatinus
Platypus facts (archived 10 September 2019)
View the platypus genome in Ensembl
PBS Nature "The Platypus Guardian"

Template:Mammaliaformes Template:Monotremata Template:Taxonbar

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Platypus: Difference between revisions

Latest revision as of 21:57, 12 November 2025

Contents

Taxonomy and naming

Description

Senses

Venom

Distribution and habitat

Ecology and behaviour

Reproduction

Evolution

Genome

Conservation

Sanctuaries and captivity

Human interactions

Usage

Cultural references

See also

Footnotes

Citations

External links

Navigation menu

@@ Line 12: / Line 12: @@
 | status = NT
 | status_system = IUCN3.1
-| status_ref = <ref name="iucn">{{cite iucn |author=Woinarski, J. |author2=Burbidge, A.A. |date=2016 |title=''Ornithorhynchus anatinus'' |volume=2016 |page=e.T40488A21964009 |doi=10.2305/IUCN.UK.2016-1.RLTS.T40488A21964009.en |access-date=19 November 2021}}</ref>
+| status_ref = <ref name="iucn">{{cite iucn |author=Woinarski, J. |author2=Burbidge, A.A. |date=2016 |title=''Ornithorhynchus anatinus'' |volume=2016 |article-number=e.T40488A21964009 |doi=10.2305/IUCN.UK.2016-1.RLTS.T40488A21964009.en |access-date=19 November 2021}}</ref>
 | taxon = Ornithorhynchus anatinus
 | parent_authority = [[Johann Friedrich Blumenbach|Blumenbach]], 1800
@@ Line 30: / Line 30: @@
 |''O. triton'' | O. Thomas, 1923
 }}
-| synonyms_ref = <ref>{{GBIF |id=2433376 |taxon=''Ornithorhynchus anatinus'' |access-date=13 July 2021}}</ref><ref>{{cite journal |last1=Pasitschniak-Arts |first1=Maria |last2=Marinelli |first2=Lui |title=Ornithorhynchus anatinus |journal=Mammalian Species |date=1 June 1998 |issue=585 |pages=1–9 |doi=10.2307/3504433 |jstor=3504433 |url=https://academic.oup.com/mspecies/article/doi/10.2307/3504433/2600754 |access-date=15 October 2024}}</ref>
+| synonyms_ref = <ref>{{GBIF |id=2433376 |taxon=''Ornithorhynchus anatinus'' |access-date=13 July 2021}}</ref><ref name=mammalianspecies/>
 | range_map = Distribution of the Platypus (Ornithorhynchus anatinus).png
 | range_map_caption = Platypus range{{break}}(red – native, yellow – introduced)
 }}
-The '''platypus''' ('''''Ornithorhynchus anatinus'''''), sometimes referred to as the '''duck-billed platypus''', is a [[semiaquatic]], egg-laying [[mammal]] endemic to [[Eastern states of Australia|eastern Australia]], including [[Tasmania]]. The platypus is the sole living representative or [[monotypic taxon]] of its [[Family (biology)|family]] [[Ornithorhynchidae]] and [[genus]] '''''Ornithorhynchus''''', though a number of [[Fossil Monotremes|related species]] appear in the fossil record.
+The '''platypus''' ('''''Ornithorhynchus anatinus'''''), sometimes referred to as the '''duck-billed platypus''', is a [[semiaquatic]], egg-laying [[mammal]] endemic to [[Eastern states of Australia|eastern Australia]], including [[Tasmania]]. The platypus is the [[monotypic taxon|sole living representative]] of its [[Family (biology)|family]] [[Ornithorhynchidae]] and [[genus]] '''''Ornithorhynchus''''', though a number of [[Fossil Monotremes|related species]] appear in the fossil record. Together with the four species of [[echidna]], it is one of the five [[wikt:extant|extant]] species of [[monotreme]]s, mammals that [[Oviparity|lay eggs]] instead of giving birth to live young. Like other monotremes, the platypus has a sense of [[electroreception|electrolocation]], which it uses to detect prey in water while its eyes, ears and nostrils are closed. It is one of the few species of [[venomous mammals]], as the male platypus has a [[spur (zoology)|spur]] on each hind foot that delivers an extremely painful [[Platypus venom|venom]].
-Together with the four species of [[echidna]], it is one of the five [[wikt:extant|extant]] species of [[monotreme]]s, mammals that lay [[Egg (biology)|eggs]] instead of giving birth to live young. Like other monotremes, the platypus has a sense of [[electroreception|electrolocation]], which it uses to detect prey in cloudy water. It is one of the few species of [[venomous mammals]], as the male platypus has a [[spur (zoology)|spur]] on the hind foot that delivers an extremely painful [[Platypus venom|venom]].
+The unusual appearance of this egg-laying, [[duck]]-billed, [[beaver]]-tailed mammal at first baffled European naturalists. In 1799, the first scientists to examine a preserved platypus body judged it a fake made of several animals sewn together. The unique features of the platypus make it important in the study of [[evolutionary biology]], and a recognisable and iconic symbol of [[Australia]]. It is culturally significant to several [[Aboriginal Australians|Aboriginal peoples]], who also used to hunt it for food, and has appeared on stamps and currency.
-The unusual appearance of this egg-laying, [[duck]]-billed, [[beaver]]-tailed, [[otter]]-footed mammal at first baffled European naturalists. In 1799, the first scientists to examine a preserved platypus body judged it a fake made of several animals sewn together.
+The platypus was hunted for its fur, but it has been a legally [[protected species]] in all states where it occurs since 1912. [[Captive breeding]] programs have had slight success, and it is vulnerable to pollution, [[Bycatch|bycatching]] and [[climate change]]. It is classified as a [[near-threatened species]] by the [[IUCN]], but a November 2020 report has recommended that it be upgraded to [[threatened species]] under the federal ''[[EPBC Act]]'', due to [[habitat destruction]] and declining numbers in all states.
-The unique features of the platypus make it important in the study of [[evolutionary biology]], and a recognisable and iconic symbol of [[Australia]]. It is culturally significant to several [[Aboriginal Australians|Aboriginal peoples]], who also used to hunt it for food. It has appeared as a national mascot, features on the [[Obverse and reverse|reverse]] of the [[Australian twenty-cent coin]], and is an emblem of the state of [[New South Wales]].
-The platypus was hunted for its fur, but it has been a legally [[protected species]] in all states where it occurs since 1912. Its population is not under severe threat, although [[captive breeding|captive-breeding]] programs have had slight success, and it is vulnerable to pollution. It is classified as a [[near-threatened species]] by the [[IUCN]], but a November 2020 report has recommended that it be upgraded to [[threatened species]] under the federal ''[[EPBC Act]]'', due to [[habitat destruction]] and declining numbers in all states.
 == Taxonomy and naming ==
 {{See also|Plural form of words ending in -us#Platypus}}
-[[Australian Aboriginal people]] name or have named the platypus in various ways depending on [[Australian Aboriginal languages|Australian indigenous languages and dialects]]. Among the names found: ''boondaburra'', ''mallingong'', ''tambreet'', ''watjarang''<ref>{{cite web|access-date=2024-10-03 |date=2024-03-06 |language=en-US |title=Platypus names (including "What's the plural of platypus?") - Australian Platypus Conservancy |url=https://platypus.asn.au/platypus-names-including-whats-the-plural-of-platypus/ |website=platypus.asn.au}}<!-- auto-translated from French by Module:CS1 translator --></ref> (names in [[Yass, New South Wales|Yass]], [[Murrumbidgee Shire|Murrumbidgee]], and [[Tumut]]),<ref name=":2" /> ''tohunbuck'' (region of [[Goomburra]], [[Darling Downs]]),<ref name=":2">{{cite web|access-date=2024-10-03 |language=en-US |title=Platypus {{!}} Wildlife Preservation Society of Queensland |url=https://wildlife.org.au/news-resources/educational-resources/species-profiles/mammals/platypus/}}<!-- auto-translated from French by Module:CS1 translator --></ref> ''dulaiwarrung''<ref>{{cite book|author1=M. Serena & G.A. Williams |date=2010 |publisher=Australian Platypus Conservancy |title=Conserving platypus and rat waters |url=https://www.tweed.nsw.gov.au/files/assets/public/v/1/documents/environment/waterways-amp-coastline/conserving-platypus-information-and-guidelines.pdf}}<!-- auto-translated from French by Module:CS1 translator --></ref> or ''dulai warrung'' ([[Woiwurrung–Taungurung language|Woiwurrung]] language, [[Wurundjeri]], [[Victoria (State)|Victoria]]),<ref name=":1">{{cite web|access-date=2024-10-03 |language=en |title=Platypus- WWF-Australia {{!}} Platypus {{!}} WWF Australia |url=https://wwf.org.au/what-we-do/species/platypus/ |website=wwf.org.au}}<!-- auto-translated from French by Module:CS1 translator --></ref> ''djanbang'' ([[Yugambeh–Bundjalung languages|Bundjalung]], [[Queensland]]),<ref>{{cite web|access-date=2024-10-03 |language=en |title=Platypus- WWF-Australia {{!}} Platypus {{!}} WWF Australia |url=https://wwf.org.au/what-we-do/species/platypus/ |website=wwf.org.au}}<!-- auto-translated from French by Module:CS1 translator --></ref> ''djumulung'' ([[Yuin–Kuric languages|Yuin]] language, [[Yuin]], [[New South Wales]]),<ref>{{cite web|access-date=2024-10-03 |language=en |title=Platypus- WWF-Australia {{!}} Platypus {{!}} WWF Australia |url=https://wwf.org.au/what-we-do/species/platypus/ |website=wwf.org.au}}<!-- auto-translated from French by Module:CS1 translator --></ref> ''maluŋgaŋ'' (ngunnawal language, [[Ngunnawal]], [[Australian Capital Territory]]<ref>{{cite web|access-date=2024-10-03 |language=en |title=Platypus- WWF-Australia {{!}} Platypus {{!}} WWF Australia |url=https://wwf.org.au/what-we-do/species/platypus/ |website=wwf.org.au}}<!-- auto-translated from French by Module:CS1 translator --></ref>), ''biladurang'', ''wamul'', ''dyiimalung'', ''oornie'', ''dungidany'' ([[Wiradjuri language|Wiradjuri]] language, [[Wiradjuri]], [[Victoria (State)|Vic]], [[New South Wales|NSW]]),<ref name=":1" /> ''oonah'',<ref>{{cite web | url=https://steemit.com/animals/@mostly.nature/platypus-out-of-the-water | title=Platypus Out of the Water | date=3 July 2017 }}</ref> etc. The name chosen and approved in [[Palawa kani]] (reconstructed [[Tasmanian languages|Tasmanian]] language) is ''larila''.<ref>{{cite web|access-date=2024-10-03 |language=en-AU |title=Larila's Legacy |url=https://hobartrivuletplatypus.org/larilas-legacy/ |website=Hobart Rivulet Platypus}}<!-- auto-translated from French by Module:CS1 translator --></ref>
+[[Australian Aboriginal people]] have referred to the platypus in various ways depending on [[Australian Aboriginal languages|Australian indigenous languages and dialects]]. Among the names found: ''boondaburra'', ''mallingong'', ''tambreet'', ''watjarang''<ref>{{cite web|access-date=2024-10-03 |date=2024-03-06 |language=en-US |title=Platypus names (including "What's the plural of platypus?") - Australian Platypus Conservancy |url=https://platypus.asn.au/platypus-names-including-whats-the-plural-of-platypus/ |website=platypus.asn.au}}<!-- auto-translated from French by Module:CS1 translator --></ref> (names in [[Yass, New South Wales|Yass]], [[Murrumbidgee Shire|Murrumbidgee]], and [[Tumut]]),<ref name=":2" /> ''tohunbuck'' (region of [[Goomburra]], [[Darling Downs]]),<ref name=":2">{{cite web|access-date=2024-10-03 |language=en-US |title=Platypus {{!}} Wildlife Preservation Society of Queensland |url=https://wildlife.org.au/news-resources/educational-resources/species-profiles/mammals/platypus/}}<!-- auto-translated from French by Module:CS1 translator --></ref> ''dulaiwarrung''<ref>{{cite book|author1=M. Serena & G.A. Williams |date=2010 |publisher=Australian Platypus Conservancy |title=Conserving platypus and rat waters |url=https://www.tweed.nsw.gov.au/files/assets/public/v/1/documents/environment/waterways-amp-coastline/conserving-platypus-information-and-guidelines.pdf}}<!-- auto-translated from French by Module:CS1 translator --></ref> or ''dulai warrung'' ([[Woiwurrung–Taungurung language|Woiwurrung]] language, [[Wurundjeri]], [[Victoria (State)|Victoria]]),<ref name=":1">{{cite web|access-date=2024-10-03 |language=en |title=Platypus- WWF-Australia {{!}} Platypus {{!}} WWF Australia |url=https://wwf.org.au/what-we-do/species/platypus/ |website=wwf.org.au}}<!-- auto-translated from French by Module:CS1 translator --></ref> ''djanbang'' ([[Yugambeh–Bundjalung languages|Bundjalung]], [[Queensland]]),<ref name=":1"/> ''djumulung'' ([[Yuin–Kuric languages|Yuin]] language, [[Yuin]], [[New South Wales]]),<ref name=":1"/> ''maluŋgaŋ'' (ngunnawal language, [[Ngunnawal]], [[Australian Capital Territory]]),<ref name=":1"/> ''biladurang'', ''wamul'', ''dyiimalung'', ''oornie'', ''dungidany'' ([[Wiradjuri language|Wiradjuri]] language, [[Wiradjuri]], [[Victoria (State)|Vic]], [[New South Wales|NSW]]),<ref name=":1" /> ''oonah'',<ref>{{cite web | url=https://steemit.com/animals/@mostly.nature/platypus-out-of-the-water | title=Platypus Out of the Water | date=3 July 2017 }}</ref> etc. The name chosen and approved in [[Palawa kani]] (reconstructed [[Tasmanian languages|Tasmanian]] language) is ''larila''.<ref>{{cite web|access-date=2024-10-03 |language=en-AU |title=Larila's Legacy |url=https://hobartrivuletplatypus.org/larilas-legacy/ |website=Hobart Rivulet Platypus}}<!-- auto-translated from French by Module:CS1 translator --></ref>
-When the platypus was first encountered by Europeans in 1798, a [[Pelage|pelt]] and sketch were sent back to [[Kingdom of Great Britain|Great Britain]] by [[John Hunter (Royal Navy officer)|Captain John Hunter]], the second Governor of [[New South Wales]].<ref name="Paradox">{{cite journal|journal=BioScience|jstor=1313511|title=The Paradoxical Platypus|first=Brian K. |last=Hall|volume=49|issue=3|pages=211–8|date=March 1999|doi=10.2307/1313511|doi-access=free}}</ref> British scientists' initial hunch was that the attributes were a hoax.<ref name="hoax">{{cite web|url=http://www.museumofhoaxes.com/hoax/archive/permalink/the_duckbilled_platypus|title=Duck-billed Platypus|publisher=Museum of hoaxes|access-date=21 July 2010|archive-date=29 July 2014|archive-url=https://web.archive.org/web/20140729074144/http://www.museumofhoaxes.com/hoax/archive/permalink/the_duckbilled_platypus|url-status=live}}</ref> [[George Shaw (biologist)|George Shaw]], who produced the first description of the animal in the ''Naturalist's Miscellany'' in 1799, stated it was impossible not to entertain doubts as to its genuine nature,<ref name="Shaw-1799">{{cite journal |last1=Shaw |first1=George |last2=Nodder |first2=Frederick Polydore |title=The Duck-Billed Platypus, ''Platypus anatinus'' |journal=The Naturalist's Miscellany |year=1799 |volume=10 |issue=CXVIII |pages=385–386 |doi=10.5962/p.304567 |url=https://www.biodiversitylibrary.org/part/304567 }}</ref> and [[Robert Knox (surgeon)|Robert Knox]] believed it might have been produced by some Asian [[Taxidermy|taxidermist]].<ref name="hoax"/> It was thought somebody had sewn a duck's beak onto the body of a beaver-like animal. Shaw even took a pair of scissors to the dried skin to check for stitches.<ref name="APC">{{cite web|url=http://www.platypus.asn.au/|title=Platypus facts file|publisher=Australian Platypus Conservancy|access-date=13 September 2006|archive-date=10 November 2015|archive-url=https://web.archive.org/web/20151110113935/http://www.platypus.asn.au/|url-status=live}}</ref><ref name="Shaw-1799" />
+When the platypus was first encountered by Europeans in 1798, a [[Pelage|pelt]] and sketch were sent back to [[Kingdom of Great Britain|Great Britain]] by [[John Hunter (Royal Navy officer)|Captain John Hunter]], the second Governor of [[New South Wales]].<ref name="Paradox">{{cite journal|journal=BioScience|jstor=1313511|title=The Paradoxical Platypus|first=Brian K. |last=Hall|volume=49|issue=3|pages=211–8|date=March 1999|doi=10.2307/1313511|doi-access=free}}</ref> British scientists' initial hunch was that the attributes were a hoax.<ref name="hoax">{{cite web|url=http://www.museumofhoaxes.com/hoax/archive/permalink/the_duckbilled_platypus|title=Duck-billed Platypus|publisher=Museum of hoaxes|access-date=21 July 2010|archive-date=29 July 2014|archive-url=https://web.archive.org/web/20140729074144/http://www.museumofhoaxes.com/hoax/archive/permalink/the_duckbilled_platypus|url-status=live}}</ref> [[George Shaw (biologist)|George Shaw]], who produced the first description of the animal in the ''Naturalist's Miscellany'' in 1799, stated it was impossible not to entertain doubts as to its genuine nature,<ref name="Shaw-1799">{{cite journal |last1=Shaw |first1=George |last2=Nodder |first2=Frederick Polydore |year=1799 |title=The Duck-Billed Platypus, ''Platypus anatinus'' |url=https://www.biodiversitylibrary.org/part/304567 |journal=The Naturalist's Miscellany |volume=10 |issue=CXVIII |pages=385–386 |doi=10.5962/p.304567 |via=Biodiversity Heritage Library}}</ref> and [[Robert Knox (surgeon)|Robert Knox]] believed—because it arrived in England via the Indian Ocean—that it might have been created by Chinese sailors.<ref name="hoax"/> It was thought somebody had sewn a duck's beak onto the body of a beaver-like animal. Shaw used a pair of scissors to check for stitches.<ref name="APC">{{cite web|url=http://www.platypus.asn.au/|title=Platypus facts file|publisher=Australian Platypus Conservancy|access-date=13 September 2006|archive-date=10 November 2015|archive-url=https://web.archive.org/web/20151110113935/http://www.platypus.asn.au/|url-status=live}}</ref><ref name="Shaw-1799" />
-The common name "platypus" literally means 'flat-foot', deriving from the [[Ancient Greek|Greek]] word {{transliteration|grc|platúpous}} ({{wikt-lang|grc|πλατύπους}}),<ref>[https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dplatu%2Fpous πλατύπους] {{Webarchive|url=https://web.archive.org/web/20210225004134/http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dplatu%2Fpous |date=25 February 2021 }}, Henry George Liddell, Robert Scott, ''A Greek-English Lexicon'', on Perseus</ref> from {{transliteration|grc|platús}} ({{wikt-lang|grc|πλατύς}} 'broad, wide, flat')<ref>[https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dplatu%2Fs πλατύς] {{Webarchive|url=https://web.archive.org/web/20210225083230/http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dplatu%2Fs |date=25 February 2021 }}, ''A Greek-English Lexicon'', on Perseus</ref>{{efn|The same root gives rise to [[platysma]], a broad, wide and flat muscle of the neck.}} and {{transliteration|grc|poús}} ({{wikt-lang|grc|πούς}} 'foot').<ref>[https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dpou%2Fs πούς] {{Webarchive|url=https://web.archive.org/web/20210227054504/http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dpou%2Fs |date=27 February 2021 }}, ''A Greek-English Lexicon'', on Perseus</ref><ref>{{cite book|author1=Liddell, Henry George|author2=Scott, Robert|name-list-style=amp|year=1980|title=Greek-English Lexicon, Abridged Edition|publisher=Oxford University Press, Oxford, UK|isbn=978-0-19-910207-5|url-access=registration|url=https://archive.org/details/lexicon00lidd}}</ref> Shaw initially assigned the species the [[Linnaean taxonomy|Linnaean]] name ''Platypus anatinus'' when he described it,<ref name="Shaw-1799"/> but the genus term was quickly discovered to already be in use as the name of the wood-boring [[ambrosia beetle]] genus ''[[Platypus (beetle)|Platypus]]''.<ref name="ABRS">{{cite book|chapter-url=http://www.deh.gov.au/biodiversity/abrs/publications/fauna-of-australia/pubs/volume1b/16-ind.pdf |title=Fauna of Australia |chapter=16 |volume=1b |first=J.R. |last=Grant |publisher=Australian Biological Resources Study (ABRS) |access-date=13 September 2006 |url-status=dead |archive-url=https://web.archive.org/web/20050519143852/http://www.deh.gov.au/biodiversity/abrs/publications/fauna-of-australia/pubs/volume1b/16-ind.pdf |archive-date=19 May 2005 }}</ref> It was independently described as ''Ornithorhynchus paradoxus'' by [[Johann Blumenbach]] in 1800 (from a specimen given to him by [[Sir Joseph Banks]])<ref name="NLA">{{cite web|url=http://www.nla.gov.au/pub/gateways/archive/52/p16a01.html|title=Platypus Paradoxes|publisher=National Library of Australia|date=August 2001|access-date=14 September 2006|archive-date=5 March 2012|archive-url=https://web.archive.org/web/20120305074657/http://www.nla.gov.au/pub/gateways/archive/52/p16a01.html|url-status=live}}</ref> and following the [[principle of priority|rules of priority]] of nomenclature, it was later officially recognised as ''Ornithorhynchus anatinus''.<ref name="ABRS"/>
+The common name "platypus" means 'flat-foot', deriving from the [[Ancient Greek|Greek]] word {{transliteration|grc|platúpous}} ({{wikt-lang|grc|πλατύπους}}),<ref>[https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dplatu%2Fpous πλατύπους] {{Webarchive|url=https://web.archive.org/web/20210225004134/http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dplatu%2Fpous |date=25 February 2021 }}, Henry George Liddell, Robert Scott, ''A Greek-English Lexicon'', on Perseus</ref> from {{transliteration|grc|platús}} ({{wikt-lang|grc|πλατύς}} 'broad, wide, flat')<ref>[https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dplatu%2Fs πλατύς] {{Webarchive|url=https://web.archive.org/web/20210225083230/http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dplatu%2Fs |date=25 February 2021 }}, ''A Greek-English Lexicon'', on Perseus</ref>{{efn|The same root gives rise to [[platysma]], a broad, wide and flat muscle of the neck.}} and {{transliteration|grc|poús}} ({{wikt-lang|grc|πούς}} 'foot').<ref>[https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dpou%2Fs πούς] {{Webarchive|url=https://web.archive.org/web/20210227054504/http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dpou%2Fs |date=27 February 2021 }}, ''A Greek-English Lexicon'', on Perseus</ref><ref>{{cite book|author1=Liddell, Henry George|author2=Scott, Robert|name-list-style=amp|year=1980|title=Greek-English Lexicon, Abridged Edition|publisher=Oxford University Press, Oxford, UK|isbn=978-0-19-910207-5|url-access=registration|url=https://archive.org/details/lexicon00lidd}}</ref> Shaw initially assigned the species the [[Linnaean taxonomy|Linnaean]] name ''Platypus anatinus'' when he described it,<ref name="Shaw-1799"/><ref name="Paradox"/> but the genus term was quickly discovered to already be in use as the name of a beetle genus ''[[Platypus (beetle)|Platypus]]''.<ref name="Paradox"/> It was independently described as ''Ornithorhynchus paradoxus'' by [[Johann Blumenbach]] in 1800 (from a specimen given to him by [[Sir Joseph Banks]])<ref name="NLA">{{cite web|url=http://www.nla.gov.au/pub/gateways/archive/52/p16a01.html|title=Platypus Paradoxes|publisher=National Library of Australia|date=August 2001|access-date=14 September 2006|archive-date=5 March 2012|archive-url=https://web.archive.org/web/20120305074657/http://www.nla.gov.au/pub/gateways/archive/52/p16a01.html|url-status=live}}</ref> and following the [[principle of priority|rules of priority]] of nomenclature, it was later officially recognised as ''Ornithorhynchus anatinus''.<ref name="ABRS">{{cite book|chapter-url=http://www.deh.gov.au/biodiversity/abrs/publications/fauna-of-australia/pubs/volume1b/16-ind.pdf |title=Fauna of Australia |chapter=16 |volume=1b |first=J.R. |last=Grant |publisher=Australian Biological Resources Study (ABRS) |access-date=13 September 2006 |archive-url=https://web.archive.org/web/20050519143852/http://www.deh.gov.au/biodiversity/abrs/publications/fauna-of-australia/pubs/volume1b/16-ind.pdf |archive-date=19 May 2005 }}</ref>
-There is no universally agreed plural form of "platypus" in the English language. Scientists generally use "platypuses" or simply "platypus". Alternatively, the term "platypi" is also used for the plural, although this is a form of [[Dog Latin|pseudo-Latin]];<ref name="APC" /> going by the word's Greek roots the plural would be "platypodes". Early [[United Kingdom|British]] [[settler]]s called it by many names, such as "watermole", "duckbill", and "duckmole".<ref name="APC" /> Occasionally it is specifically called the "duck-billed platypus".
+Various dictionaries list "platypuses" or simply "platypus" as the plural. Alternatively, the term "platypi" is also used for the plural, although this is a form of [[Dog Latin|pseudo-Latin]]; going by the word's Greek roots the plural would be "platypodes".<ref name="APC" /> Early European [[settler]]s called it by many names, such as "watermole", "duckbill", and "duckmole".<ref name="Paradox"/> Occasionally it is specifically called the "duck-billed platypus".<ref name="hoax"/> There is no official term for platypus young, but the term "platypup" sees unofficial use, as does "puggle".<ref>{{cite book|last=Carmody|first=Judy|year=2011|title=Wet Tropics of Queensland World Heritage Area: Tour Guide Handbook|publisher=James Cook University, Marine and Tropical Science Research Facility|url=https://researchonline.jcu.edu.au/18787/1/Carmody-J-2011-Wet-Tropics-Tour-Guide-Handbook.pdf#page=160|access-date=8 February 2021|archive-date=7 June 2020|archive-url=https://web.archive.org/web/20200607085049/https://researchonline.jcu.edu.au/18787/1/Carmody-J-2011-Wet-Tropics-Tour-Guide-Handbook.pdf#page=160|url-status=live}}</ref><ref>{{cite web |author=Australian National Dictionary Centre |title=Oxford Word of the Month - November: platypup |url=https://www.oup.com.au/__data/assets/pdf_file/0027/108477/WotM_November_2017.pdf |publisher=Oxford University Press |date=November 2017|access-date=20 April 2022}}</ref>
-The scientific name ''Ornithorhynchus anatinus'' literally means 'duck-like bird-snout',<ref name="Shaw-1799"/> deriving its [[generic epithet|genus name]] from the Greek root {{transliteration|grc|ornith-}} ({{wikt-lang|grc|ὄρνις|όρνιθ}} ''ornith'' or {{lang|el|ὄρνις}} ''órnīs'' 'bird')<ref>{{cite web |url=http://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=o)/rnis |title=ὄρνις |last1=Liddell |first1=Henry George |last2=Scott |first2=Robert |work=A Greek-English Lexicon |publisher=Perseus Digital Library |date=1940 |access-date=29 September 2022 |archive-date=4 April 2023 |archive-url=https://web.archive.org/web/20230404195515/https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=o)/rnis |url-status=live }}</ref> and the word {{transliteration|grc|rhúnkhos}} ({{wikt-lang|grc|ῥύγχος}} 'snout', 'beak').<ref>{{cite web |url=http://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=r(u/gxos |title=ῥύγχος |last1=Liddell |first1=Henry George |last2=Scott |first2=Robert |work=A Greek-English Lexicon |publisher=Perseus Digital Library |date=1940 |access-date=29 September 2022 |archive-date=23 April 2023 |archive-url=https://web.archive.org/web/20230423120445/https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=r(u/gxos |url-status=live }}</ref> Its [[specific epithet|species name]] is derived from Latin {{wikt-lang|la|anatinus}} ('duck-like') from {{wikt-lang|la|anas}} 'duck'.<ref name="Shaw-1799"/><ref>{{cite encyclopedia |url= https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0059%3Aentry%3Danas1 |title= ănăs |last1= Lewis |first1= Charlton T. |last2= Short |first2= Charles |dictionary= A Latin Dictionary |edition=  |publisher= Perseus Digital Library |date= 1879 |access-date= 29 September 2022 |archive-date= 27 September 2023 |archive-url= https://web.archive.org/web/20230927152321/http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0059%3Aentry%3Danas1 |url-status= live }}</ref> The platypus is the sole living representative or monotypic taxon of its [[Family (biology)|family]] (Ornithorhynchidae).<ref>{{cite web |last=Bess |first=Anna |url=https://animaldiversity.org/accounts/Ornithorhynchidae/ |title=ADW: Ornithorhynchidae: INFORMATION |publisher=Animaldiversity.org |date= |access-date=11 February 2022 |archive-date=17 January 2022 |archive-url=https://web.archive.org/web/20220117035152/https://animaldiversity.org/accounts/Ornithorhynchidae/ |url-status=live }}</ref>
+The scientific name ''Ornithorhynchus anatinus'' literally means 'duck-like bird-snout',<ref name="Shaw-1799"/> deriving its [[generic epithet|genus name]] from the Greek root {{transliteration|grc|ornith-}} ({{wikt-lang|grc|ὄρνις|όρνιθ}} ''ornith'' or {{lang|el|ὄρνις}} ''órnīs'' 'bird')<ref>{{cite web |url=http://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=o)/rnis |title=ὄρνις |last1=Liddell |first1=Henry George |last2=Scott |first2=Robert |work=A Greek-English Lexicon |publisher=Perseus Digital Library |date=1940 |access-date=29 September 2022 |archive-date=4 April 2023 |archive-url=https://web.archive.org/web/20230404195515/https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=o)/rnis |url-status=live }}</ref> and the word {{transliteration|grc|rhúnkhos}} ({{wikt-lang|grc|ῥύγχος}} 'snout', 'beak').<ref>{{cite web |url=http://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=r(u/gxos |title=ῥύγχος |last1=Liddell |first1=Henry George |last2=Scott |first2=Robert |work=A Greek-English Lexicon |publisher=Perseus Digital Library |date=1940 |access-date=29 September 2022 |archive-date=23 April 2023 |archive-url=https://web.archive.org/web/20230423120445/https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=r(u/gxos |url-status=live }}</ref> Its [[specific epithet|species name]] is derived from Latin {{wikt-lang|la|anatinus}} ('duck-like') from {{wikt-lang|la|anas}} 'duck'.<ref name="Shaw-1799"/><ref>{{cite encyclopedia |url= https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0059%3Aentry%3Danas1 |title= ănăs |last1= Lewis |first1= Charlton T. |last2= Short |first2= Charles |dictionary= A Latin Dictionary |edition= |publisher= Perseus Digital Library |date= 1879 |access-date= 29 September 2022 |archive-date= 27 September 2023 |archive-url= https://web.archive.org/web/20230927152321/http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0059%3Aentry%3Danas1 |url-status= live }}</ref> The platypus is the sole living representative or monotypic taxon of its [[Family (biology)|family]] (Ornithorhynchidae).<ref>{{cite web |last=Bess |first=Anna |url=https://animaldiversity.org/accounts/Ornithorhynchidae/ |title=ADW: Ornithorhynchidae: INFORMATION |publisher=Animaldiversity.org |date= |access-date=11 February 2022 |archive-date=17 January 2022 |archive-url=https://web.archive.org/web/20220117035152/https://animaldiversity.org/accounts/Ornithorhynchidae/ |url-status=live }}</ref>
 <gallery mode="packed" heights="180px">
 platypus-plate.jpg|A book for children published in Germany in 1798
@@ Line 64: / Line 60: @@
 == Description ==
-In [[David Collins (lieutenant governor)|David Collins]]'s account of the new colony 1788–1801, he describes "an amphibious animal, of the mole species", with a drawing.<ref>{{cite book|url=https://archive.org/details/anaccountoftheen12668gut|title=An Account of the English Colony in New South Wales, Volume 2|first=David|last=Collins|access-date=5 July 2017|via=Internet Archive}}</ref>
+[[File:Platypus skeleton Pengo.jpg|thumb|Platypus skeleton]]
+Most of the platypus' small streamlined body is covered with short, dense, brown, [[fur]] that traps a layer of insulating air to keep the animal warm, both in and out of water.<ref name="APC" /><ref name="ABRS" /><ref name=Grant89>{{cite book|last=Grant|first=Tom|year=1989|title=The Platypus: A Unique Mammal|publisher=New South Wales Press|isbn=0-86840-243-5}}</ref>{{rp|1}} The fur coat is waterproof and consists of flattened guard hairs and curvy underfur hairs.<ref name="ABRS" /><ref name=Grant89/>{{rp|2}} It is one of the most densely furred mammals, behind only [[otter]]s.<ref name="sensory_platypus" /> It is also [[biofluorescent]] and glows [[cyan]] and green when under [[ultraviolet]] light; this may serve to camouflage it in low lighting from UV-sensitive predators.<ref>{{cite journal |last1=Anich |first1=Paula Spaeth |title=Biofluorescence in the platypus (Ornithorhynchus anatinus) |journal=[[Mammalia (journal)|Mammalia]] |date=15 October 2020 |volume=85 |issue=2 |pages=179–181 |doi=10.1515/mammalia-2020-0027 |doi-access=free }}</ref> The duck-like bill consists of a long snout and lower jaw which is covered in soft skin. The nostrils are located near the tip of the snout's dorsal surface, while the eyes and ears are just behind the snout in a groove which closes underwater.<ref name="ABRS" /> It has [[cheek pouches]] for storing food. The platypus's wide, flat tail is compared to a [[beaver]]'s but is furry rather than scaly;<ref name=mammalianspecies/> it stores fat reserves and can act as a rudder during swimming.<ref name="ABRS" /><ref name=Grant89/>{{rp|4}} The legs are short and have a sprawling stance. [[webbed foot|Webbing]] is more significant on the front feet. While walking on land, the feet are folded up in [[knuckle-walking]] to protect the webbing.<ref name=Grant89/>{{rp|2, 4}}<ref name="Fish">{{cite journal |last1=Fish |first1=F. E. |last2=Frappell |first2=P. B. |last3=Baudinette |first3=R. V. |last4=Macfarlane |first4=P. M. |title=Energetics of Terrestrial Locomotion of the Platypus Ornithorhynchus Anatinus |journal=Journal of Experimental Biology |date=15 February 2001 |volume=204 |issue=4 |pages=797–803 |doi=10.1242/jeb.204.4.797 |pmid=11171362 |bibcode=2001JExpB.204..797F |hdl-access=free |hdl=2440/12192 }}</ref>
-The body and the broad, flat tail of the platypus are covered with dense, brown, biofluorescent [[fur]] that traps a layer of insulating air to keep the animal warm.<ref name="APC" /><ref name="ABRS" /><ref>{{cite journal |last1=Anich |first1=Paula Spaeth |title=Biofluorescence in the platypus (Ornithorhynchus anatinus) |journal=[[Mammalia (journal)|Mammalia]] |date=15 October 2020 |volume=85 |issue=2 |pages=179–181 |doi=10.1515/mammalia-2020-0027 |doi-access=free }}</ref> The fur is waterproof, and textured like that of a [[Mole (animal)|mole]].<ref>{{cite web |url=http://animal.discovery.com/mammals/platypus/ |title=Platypus: Facts, Pictures: Animal Planet |publisher=Animal.discovery.com |date=16 November 2011 |access-date=8 September 2012 |archive-date=27 July 2011 |archive-url=https://web.archive.org/web/20110727171511/http://animal.discovery.com/mammals/platypus/ |url-status=live }}</ref> The platypus's tail stores fat reserves, an adaptation also found in the [[Tasmanian devil]].<ref name="Guiler">{{cite book|author=Guiler, E.R.|year=1983|chapter=Tasmanian Devil|editor=R. Strahan|title=The Australian Museum Complete Book of Australian Mammals|pages=27–28|publisher=Angus & Robertson|isbn=978-0-207-14454-7}}</ref> [[webbed foot|Webbing]] is more significant on the front feet, which in land walking are folded up in [[knuckle-walking]] to protect the webbing.<ref name="Fish">{{cite journal |last1=Fish |first1=F. E. |last2=Frappell |first2=P. B. |last3=Baudinette |first3=R. V. |last4=Macfarlane |first4=P. M. |title=Energetics of Terrestrial Locomotion of the Platypus Ornithorhynchus Anatinus |journal=Journal of Experimental Biology |date=15 February 2001 |volume=204 |issue=4 |pages=797–803 |doi=10.1242/jeb.204.4.797 |pmid=11171362 |bibcode=2001JExpB.204..797F |hdl-access=free |hdl=2440/12192 }}</ref> The elongated snout and lower jaw are covered in soft skin, forming the bill. The nostrils are located on the snout's dorsal surface, while the eyes and ears are just behind the snout in a groove which closes underwater.<ref name="ABRS" /> Platypuses can give a low growl when disturbed, and a range of vocalisations have been reported in captivity.<ref name="APC" />
-Size varies considerably in different regions, with average weight from {{convert|0.7|to|2.4|kg|lboz|abbr=on}}; males have average length {{convert|50|cm|in|0|abbr=on}}, while females are the smaller at {{convert|43|cm|in|frac=2|abbr=on}}.<ref name="ABRS" /> This variation does not seem to follow any particular climatic rule and may be due to other factors such as predation and human encroachment.<ref name="Workshop">{{cite web|url=http://www.medicine.utas.edu.au/research/mono/Taspaper.html|title=Current research on the platypus, ''Ornithorhynchus anatinus'' in Tasmania: Abstracts from the 1999 'Tasmanian Platypus Workshop'|author1=Munks, Sarah |author2=Nicol, Stewart |name-list-style=amp |publisher=University of Tasmania|date=May 1999| access-date = 23 October 2006|archive-url = https://web.archive.org/web/20060830075935/http://www.medicine.utas.edu.au/research/mono/Taspaper.html |archive-date = 30 August 2006|url-status=dead}}</ref>
-The platypus has an average [[Core temperature|body temperature]] of about {{convert|32|C|F}}, lower than the {{convert|37|C|F}} typical of [[placentalia|placental mammals]].<ref name="DC">{{cite web|url=http://www.bio.davidson.edu/courses/anphys/1999/White/thermal.htm |title=Thermal Biology of the Platypus |publisher=Davidson College |year=1999 |access-date=14 September 2006 |url-status=dead |archive-url=https://web.archive.org/web/20120306024923/http://www.bio.davidson.edu/courses/anphys/1999/White/thermal.htm |archive-date=6 March 2012 }}</ref> Research suggests this has been a gradual adaptation to harsh environmental conditions among the few marginal surviving monotreme species, rather than a general characteristic of past monotremes.<ref name="temp">{{cite journal |last1=Watson |first1=Jm |last2=Graves |first2=Jam |title=Monotreme Cell-Cycles and the Evolution of Homeothermy |journal=Australian Journal of Zoology |date=1988 |volume=36 |issue=5 |pages=573 |doi=10.1071/ZO9880573 }}</ref><ref name="temp2">{{cite journal |last1=Dawson |first1=Tj |last2=Grant |first2=Tr |last3=Fanning |first3=D |title=Standard Metabolism of Monotremes and the Evolution of Homeothermy. |journal=Australian Journal of Zoology |date=1979 |volume=27 |issue=4 |pages=511 |doi=10.1071/ZO9790511 }}</ref>
-In addition to laying eggs, the anatomy, [[ontogeny]], and genetics of monotremes shows traces of similarity to reptiles and birds. The platypus has a [[Reptile|reptilian]] gait with legs on the sides of the body, rather than underneath.<ref name="ABRS" /> The platypus's genes are a possible evolutionary link between the mammalian [[XY sex-determination system|XY]] and bird/reptile [[ZW sex-determination system|ZW]] sex-determination systems, as one of the platypus's five X chromosomes contains the [[DMRT1]] gene, which birds possess on their Z chromosome.<ref name="Graves">{{cite journal |last=Graves |first=Jennifer |date=10 March 2006 |title=Sex Chromosome Specialization and Degeneration in Mammals |journal=Cell |volume=124 |issue=5 |pages=901–914 |doi=10.1016/j.cell.2006.02.024 |pmid=16530039 |s2cid=8379688 |doi-access=free}}</ref>
+The platypus has an [[interclavicle]] in the shoulder girdle, a trait which they share in common with reptiles.<ref name=Grant89/>{{rp|7}} As in many other aquatic and semiaquatic [[vertebrates]], the bones show [[osteosclerosis]], increasing their density to reduce buoyancy.<ref>{{Cite journal | doi = 10.1371/journal.pone.0059146|pmid=23565143|pmc=3615000| title = Bone Inner Structure Suggests Increasing Aquatic Adaptations in Desmostylia (Mammalia, Afrotheria)| journal = PLOS ONE| volume = 8| issue = 4| article-number = e59146| year = 2013| last1 = Hayashi | first1 = S.| last2 = Houssaye | first2 = A.| last3 = Nakajima | first3 = Y.| last4 = Chiba | first4 = K.| last5 = Ando | first5 = T.| last6 = Sawamura | first6 = H.| last7 = Inuzuka | first7 = N.| last8 = Kaneko | first8 = N.| last9 = Osaki | first9 = T.|bibcode=2013PLoSO...859146H|doi-access=free}}</ref> Adult platypuses lack teeth and instead have heavily [[keratinisation|keratinised]] food-grinding pads.<ref name="ABRS" /> Young platypuses have one premolar tooth and two [[Molar (tooth)|molars]] on each [[maxilla]]e, and three molars on the [[Mandible|dentaries]]. The first upper and third lower cheek teeth have only one major cusp, while the rest have two.<ref>{{cite book |last=Ungar |first=Peter S. |title=Mammal Teeth: Origin, Evolution, and Diversity |publisher=The Johns Hopkins University Press |year=2010 |isbn=978-0-801-89668-2 |page=130 |chapter=Monotremata and Marsupialia}}</ref> They lose their teeth around the time they leave their natal burrow.<ref name="ABRS" />
-As in all true mammals, the tiny bones that conduct sound in the [[middle ear]] are fully incorporated into the skull, rather than lying in the jaw as in pre-mammalian [[synapsid]]s. However, the external opening of the ear still lies at the base of the jaw.<ref name="ABRS" /> The platypus has extra bones in the shoulder girdle, including an [[interclavicle]] not found in other mammals.<ref name="ABRS" /> As in many other aquatic and semiaquatic [[vertebrates]], the bones show [[osteosclerosis]], increasing their density to provide ballast.<ref>{{Cite journal | doi = 10.1371/journal.pone.0059146|pmid=23565143|pmc=3615000| title = Bone Inner Structure Suggests Increasing Aquatic Adaptations in Desmostylia (Mammalia, Afrotheria)| journal = PLOS ONE| volume = 8| issue = 4| pages = e59146| year = 2013| last1 = Hayashi | first1 = S.| last2 = Houssaye | first2 = A.| last3 = Nakajima | first3 = Y.| last4 = Chiba | first4 = K.| last5 = Ando | first5 = T.| last6 = Sawamura | first6 = H.| last7 = Inuzuka | first7 = N.| last8 = Kaneko | first8 = N.| last9 = Osaki | first9 = T.|bibcode=2013PLoSO...859146H|doi-access=free}}</ref>
+Male platypuses have an average length of {{convert|50|cm|in|0|abbr=on}} and weight of {{convert|1700|g|lb|abbr=on}}, while females are smaller with an average length of {{convert|43|cm|in|frac=2|abbr=on}} and weight of {{convert|900|g|lb|abbr=on}}.<ref name="ABRS" /> The species follows [[Bergmann's rule]], with individuals being larger the farther south they are, due to colder climates; there are local variations, however.<ref>{{cite journal|last1=Furlan|first1=E|last2=Griffiths|first2=J|last3=Gust|first3=N|last4=Armistead|first4=R|last5=Mitrovski|first5=P|last6=Handasyde|first6=K. A.|last7=Serena|first7=M|last8=Hoffmann|first8=A. A.|last9=Weeks|first9=A. R.|year=2012|title=Is body size variation in the platypus (Ornithorhynchus anatinus) associated with environmental variables?|journal=Australian Journal of Zoology|volume=59|issue=4|pages=201–215|doi=10.1071/ZO11056}}</ref> The platypus has an average [[Core temperature|body temperature]] of about {{convert|32|C|F}}, lower than the {{convert|37|C|F}} typical of [[placentalia|placental mammals]].<ref name="DC">{{cite web|url=http://www.bio.davidson.edu/courses/anphys/1999/White/thermal.htm |title=Thermal Biology of the Platypus |publisher=Davidson College |year=1999 |access-date=14 September 2006 |archive-url=https://web.archive.org/web/20120306024923/http://www.bio.davidson.edu/courses/anphys/1999/White/thermal.htm |archive-date=6 March 2012 }}</ref> Research suggests this has been a gradual adaptation to harsh environmental conditions among the few marginal surviving monotreme species, rather than a general characteristic of past monotremes.<ref name="temp">{{cite journal |last1=Watson |first1=Jm |last2=Graves |first2=Jam |title=Monotreme Cell-Cycles and the Evolution of Homeothermy |journal=Australian Journal of Zoology |date=1988 |volume=36 |issue=5 |page=573 |doi=10.1071/ZO9880573 }}</ref><ref name="temp2">{{cite journal |last1=Dawson |first1=Tj |last2=Grant |first2=Tr |last3=Fanning |first3=D |title=Standard Metabolism of Monotremes and the Evolution of Homeothermy. |journal=Australian Journal of Zoology |date=1979 |volume=27 |issue=4 |page=511 |doi=10.1071/ZO9790511 }}</ref>
-The platypus [[jaw]] is constructed differently from that of other mammals, and the jaw-opening muscle is different.<ref name="ABRS" /> Modern platypus young have three teeth in each of the [[maxilla]]e (one premolar and two [[Molar (tooth)|molars]]) and [[Mandible|dentaries]] (three molars), which they lose before or just after leaving the breeding burrow;<ref name="ABRS" /> adults instead develop heavily [[keratinisation|keratinised]] food-grinding pads called [[ceratodontes]].<ref name="ABRS" /><ref name="Piper-2007" /><ref name="Haeckel1895">{{cite book |last=Haeckel |url=https://books.google.com/books?id=AOi2AAAAIAAJ&pg=PA142 |title=Systematische Phylogenie der Wirbelthiere (Vertebrata) |publisher=Georg Reimer |year=1895 |edition=1 |series=Entwurf einer systematischen Stammesgeschichte |volume=3 |location=Berlin |pages=142–143 |language=de |access-date=16 July 2021 |archive-url=https://web.archive.org/web/20210716071006/https://books.google.com/books?id=AOi2AAAAIAAJ&pg=PA142 |archive-date=16 July 2021 |url-status=live}}</ref> The first upper and third lower cheek teeth of platypus nestlings are small, each having one principal cusp, while the other teeth have two main cusps.<ref>{{cite book |last=Ungar |first=Peter S. |title=Mammal Teeth: Origin, Evolution, and Diversity |publisher=The Johns Hopkins University Press |year=2010 |isbn=978-0-801-89668-2 |page=130 |chapter=Monotremata and Marsupialia}}</ref>
+The platypus has a single opening, called a [[cloaca]], for both the reproductive and waste systems.<ref name=Grant89/>{{rp|32–33}} The male platypus has [[penile spines]] and an asymmetrical [[glans penis]] that is larger on the left side.<ref name=Vogelnest>{{Cite book |last1=Vogelnest |first1=Larry |url=https://books.google.com/books?id=4d6VDwAAQBAJ&dq=platypus&pg=PA64 |title=Current Therapy in Medicine of Australian Mammals |last2=Portas |first2=Timothy |date=2019-05-01 |publisher=Csiro Publishing |isbn=978-1-4863-0752-4 |language=en|pages=64, 107}}</ref> The female has two [[Ovary|ovaries]], with the right one being non-functional,<ref name=Vogelnest/><ref name="EC">{{cite web|url=http://www.biology.iastate.edu/InternationalTrips/1Australia/04papers/CromerMonotrRepro.htm |title=Monotreme Reproductive Biology and Behavior |publisher=Iowa State University |author=Cromer, Erica |date=14 April 2004 |access-date=18 June 2009 |archive-url=https://web.archive.org/web/20090313030236/http://www.biology.iastate.edu/InternationalTrips/1Australia/04papers/CromerMonotrRepro.htm |archive-date=13 March 2009 }}</ref> and [[teat]]s are absent.<ref name="ABRS" />
 <gallery mode="packed" heights="180px">
 Duck-billed platypus (Ornithorhynchus anatinus) diving Scottsdale.jpg|Diving
-Duck-billed platypus (Ornithorhynchus anatinus) surfacing Scottsdale.jpg|Surfacing
+duck-billed platypus (Ornithorhynchus anatinus) surfacing Scottsdale.jpg|Surfacing
-Duck-billed platypus (Ornithorhynchus anatinus) Scottsdale 4.jpg
+duck-billed platypus (Ornithorhynchus anatinus) Scottsdale 4.jpg
 </gallery>
-=== Venom ===
+=== Senses ===
-{{Main|Platypus venom}}
+[[File:Platypus electrolocation.svg|thumb|upright=2<!--for readability of svg text-->|The platypus has secondarily acquired [[Electroreception and electrogenesis|electroreception]]. Its receptors are arranged in stripes on its bill, giving it high sensitivity to the sides and below; it makes quick turns of its head as it swims to detect prey.<ref name="Electro1"/>]]
-[[File:Platypus spur.JPG|right|thumb|The calcaneus spur on the male's hind limb is used to inject venom.]]
-While both male and female platypuses are born with back ankle spurs, only the males' deliver venom.<ref>{{cite web |url=http://www.australianfauna.com/platypus.php |title=Australian Fauna |publisher=Australian Fauna |access-date=14 May 2010 |url-status=dead |archive-url=https://web.archive.org/web/20120529040439/http://www.australianfauna.com/platypus.php |archive-date=29 May 2012 }}</ref><ref>{{cite web |url=http://www.usyd.edu.au/news/84.html?newsstoryid=2267 |title=Platypus venom linked to pain relief |publisher=University of Sydney |date=8 May 2008 |access-date=14 May 2010 |archive-date=21 August 2011 |archive-url=https://web.archive.org/web/20110821151157/http://www.usyd.edu.au/news/84.html?newsstoryid=2267 |url-status=live }}</ref><ref>{{cite web |url=http://rainforest-australia.com/platypus_poison.htm |title=Platypus poison |publisher=Rainforest Australia |access-date=14 May 2010 |url-status=dead |archive-url=https://web.archive.org/web/20100529165453/http://rainforest-australia.com/platypus_poison.htm |archive-date=29 May 2010 }}</ref>
+[[Monotremes]] are the only mammals (apart from the [[Guiana dolphin]])<ref>{{cite journal |last1=Czech-Damal |first1=Nicole U. |last2=Liebschner |first2=Alexander |last3=Miersch |first3=Lars |last4=Klauer |first4=Gertrud |last5=Hanke |first5=Frederike D. |last6=Marshall |first6=Christopher |last7=Dehnhardt |first7=Guido |last8=Hanke |first8=Wolf |date=22 February 2012 |title=Electroreception in the Guiana dolphin (Sotalia guianensis) |journal=Proceedings of the Royal Society B: Biological Sciences |volume=279 |issue=1729 |pages=663–668 |doi=10.1098/rspb.2011.1127 |pmc=3248726 |pmid=21795271}}</ref> known to have a sense of [[electroreception]].<ref>{{cite journal |last1=Proske |first1=Uwe |last2=Gregory |first2=J. E. |last3=Iggo |first3=A. |year=1998 |title=Sensory receptors in monotremes |journal=Philosophical Transactions of the Royal Society of London |volume=353 |issue=1372 |pages=1187–1198 |doi=10.1098/rstb.1998.0275 |pmc=1692308 |pmid=9720114}}</ref><ref name="Electro1">{{cite journal |last=Pettigrew |first=John D. |year=1999 |title=Electroreception in Monotremes |url=http://jeb.biologists.org/cgi/reprint/202/10/1447.pdf |url-status=live |journal=The Journal of Experimental Biology |volume=202 |issue=Part 10 |pages=1447–54 |doi=10.1242/jeb.202.10.1447 |pmid=10210685 |archive-url=https://web.archive.org/web/20060928054253/http://jeb.biologists.org/cgi/reprint/202/10/1447.pdf |archive-date=28 September 2006 |access-date=19 September 2006 |doi-access=free|bibcode=1999JExpB.202.1447P }}</ref> The playtpus relies on electrolocation when feeding, as the eyes, ears, and nose are closed while underwater.<ref>{{cite journal |author=Gregory, J.E. |author2=Iggo, A. |author3=McIntyre, A.K. |author4=Proske, U. |date=June 1988 |title=Receptors in the Bill of the Platypus |journal=Journal of Physiology |volume=400 |issue=1 |pages=349–366 |doi=10.1113/jphysiol.1988.sp017124 |pmc=1191811 |pmid=3418529}}</ref><ref name="draft_genome" /> Digging in the bottom of streams with its bill, its electroreceptors detect tiny electric currents generated by the muscular contractions of its prey.<ref name="Electro1" /> Experiments have shown the platypus will even react to an "artificial shrimp" if a small electric current is passed through it.<ref name="Manning">{{cite book |author1=Manning, A. |url=https://archive.org/details/introductiontoan0000mann_m2z8 |title=An Introduction to Animal Behaviour |author2=Dawkins, M.S. |publisher=Cambridge University Press |year=1998 |edition=5th|page=135|url-access=registration}}</ref>
-It is powerful enough to kill smaller animals such as dogs, and though it is not lethal to humans, it can inflict weeks of agony.<ref name="PS" /><ref>Weimann, Anya (4 July 2007) [https://web.archive.org/web/20121105043922/http://www.cosmosmagazine.com/news/1423/evolution-platypus-venom-revealed Evolution of platypus venom revealed]. ''[[Cosmos (magazine)|Cosmos]]''.</ref> [[Edema]] rapidly develops around the wound and gradually spreads through the affected limb, and it may develop into an excruciating [[hyperalgesia]] (heightened sensitivity to pain) persisting for days or even months.<ref name="JN">{{cite journal |author=de Plater, G.M. |author2=Milburn, P.J. |author3=Martin, R.L. |year=2001 |title=Venom From the Platypus, ''Ornithorhynchus anatinus'', Induces a Calcium-Dependent Current in Cultured Dorsal Root Ganglion Cells |journal=Journal of Neurophysiology |volume=85 |issue=3 |pages=1340–5 |doi=10.1152/jn.2001.85.3.1340 |pmid=11248005 |s2cid=2452708 }}</ref><ref name="venom">{{cite web |title=The venom of the platypus (''Ornithorhynchus anatinus'') |url=http://www.kingsnake.com/toxinology/old/mammals/platypus.html |url-status=dead |archive-url=https://web.archive.org/web/20120201185504/http://www.kingsnake.com/toxinology/old/mammals/platypus.html |archive-date=1 February 2012 |access-date=13 September 2006}}</ref>
-The venom is composed largely of [[defensin]]-like [[protein]]s (DLPs) produced by the immune system, three of which are unique to the platypus.<ref name="PS">{{cite journal| last = Gerritsen| first = Vivienne Baillie| title = Platypus poison| journal = Protein Spotlight| issue = 29| date = December 2002| url = http://www.expasy.org/spotlight/back_issues/sptlt029.shtml| access-date = 14 September 2006| archive-date = 20 October 2008| archive-url = https://web.archive.org/web/20081020054110/http://www.expasy.org/spotlight/back_issues/sptlt029.shtml| url-status = live}}</ref> In other animals, defensins [[lysis|kill]] pathogenic bacteria and viruses, but in platypuses they are also collected into a venom against predators. Venom is produced in the [[wikt:crural|crural]] glands of the male, which are kidney-shaped [[alveolar gland]]s connected by a thin-walled duct to a [[calcaneus]] spur on each hind limb. The female platypus, in common with echidnas, has rudimentary spur buds that do not develop (dropping off before the end of their first year) and lack functional crural glands.<ref name="ABRS" /> Venom production rises among males during the breeding season, and it may be used to assert dominance.<ref name="PS" />
+The 40,000 [[electroreceptor]]s are arranged in rows in the skin of the bill from front to back, while [[mechanoreceptor]]s for touch are uniformly distributed across the bill. The electrosensory area of the [[cerebral cortex]] is in the tactile [[somatosensory]] area, and some cortical cells receive input from both electroreceptors and mechanoreceptors, suggesting the platypus feels electric fields as touches. These receptors in the bill dominate the [[somatotopic map]] of the platypus brain, in the same way human hands dominate the [[Cortical homunculus|Penfield homunculus map]].<ref name="sensory_platypus">{{cite journal |last1=Pettigrew |first1=John D. |first2=P. R. |last2=Manger |first3=S. L. |last3=Fine |title=The sensory world of the platypus |journal=Philosophical Transactions of the Royal Society of London |pages=1199–1210 |issue=1372 |year=1998 |pmid=9720115|pmc=1692312 |doi=10.1098/rstb.1998.0276 |volume=353 }}</ref><ref name="Ancestors_Tale">{{cite book |first=Richard |last=Dawkins |author-link=Richard Dawkins |title=The Ancestor's Tale, A Pilgrimage to the Dawn of Life |chapter=The Duckbill's Tale |publisher=Houghton Mifflin |location=Boston, Massachusetts |year=2004 |isbn=978-0-618-00583-3 |title-link=The Ancestor's Tale }}</ref> The platypus can feel the direction of an electric source, perhaps by comparing differences in [[signal strength]] across the array of electroreceptors, enhanced by the characteristic side-to-side motion of the animal's head while hunting. It may also be able to determine the distance of moving prey via the timing difference between electrical and mechanical pressure sensations.<ref name="Electro1" /> Monotreme electrolocation for hunting in murky waters may be tied to their tooth loss. The extinct ''[[Obdurodon]]'' was electroreceptive, but unlike the modern platypus it foraged in open water.<ref name="Masakazu Asahara 2016">{{cite journal|last1=Asahara|first1=M|last2=Koizumi|first2=M|last3=Macrini|first3=T. E.|last4=Hand|first4=S, J.|last5=Archer|first5=M|year=2016|title=Comparative cranial morphology in living and extinct platypuses: Feeding behavior, electroreception, and loss of teeth|journal=Science Advances|volume=2|issue=10|article-number=e1601329|doi=10.1126/sciadv.1601329|pmid=27757425|pmc=5061491|bibcode=2016SciA....2E1329A}}</ref>
-Similar spurs are found on many archaic mammal groups, indicating that this was an ancient general characteristic among mammals.<ref>Jørn H. Hurum, Zhe-Xi Luo, and Zofia Kielan-Jaworowska,
+The eyes of the platypus have [[Basal (phylogenetics)|basal]] traits also found in [[lungfish]] and [[amphibian]]s, such as [[Scleral ring|scleral cartilage]], [[double cone (biology)|double cones]], and [[Oil droplet|droplets]].<ref>{{cite journal |last1=Zeiss|first1=Caroline|title=Comparative retinal morphology of the platypus|doi=10.1002/jmor.10959 |year=2011 |first2=Ivan R. |last3=Murphy|first3=Christopher J. |last4=Dubielzig|first4=Richard W. |journal=Journal of Morphology |volume=272 |issue=8 |pages=949–57 |pmid=21567446|last2=Schwab|bibcode=2011JMorp.272..949Z |s2cid=28546474}}</ref> The platypus's eyes are small and shut under water, though several features indicate its ancestors relied on vision. As with other aquatic mammals, the eye has a flattened cornea and surrounding lens, while the posterior surface of the lens is sharply inclined. A [[Temporal bone|temporal]] (ear side) concentration of [[retinal ganglion cell]]s, important for binocular vision, indicates a vestigial role in [[predation]], though the actual visual acuity is insufficient for such activities. Limited acuity is matched by low [[cortical magnification]], a small [[lateral geniculate nucleus]], and a large [[optic tectum]], suggesting that the [[Midbrain tectum|visual midbrain]] plays a more important role than the [[visual cortex]], as in some rodents. These features suggest that the platypus has adapted to an aquatic and nocturnal lifestyle, developing its electrosensory system at the cost of its visual system. This contrasts with the small number of electroreceptors in the [[short-beaked echidna]], which dwells in dry environments, while the [[long-beaked echidna]], which lives in wetter habitats is intermediate between the other two monotremes.<ref name="sensory_platypus" />
-Were mammals originally venomous?, Acta Palaeontologica Polonica 51 (1), 2006: 1–11</ref>
-=== Electrolocation ===
+The ears of the platypus are adapted for hearing while out of water.<ref name="sensory_platypus" /> As in all true mammals, it has three [[middle ear]] bones, though the [[cochlea]] lacks spirals,<ref name="ABRS" /> but is described as "well organised". Within the cochlea, there are rows of inner and outer [[hair cell]]s. As in placental mammals, the outer hair cells of the platypus are adapted for hearing high frequencies, suggesting it is an ancestral mammalian trait. However it also possesses more rows of inner hair cells.<ref name="sensory_platypus" /> The [[olfactory]] (smelling) systems of the platypus and the echidna independently evolved from an ancestor with less advanced smelling. The main olfactory bulb of the platypus lacks the complex layers of the echidna, while both the [[piriform cortex]] and flaps ([[Lamella (surface anatomy)|lamella]]) are simpler. Monotremes differ from placental mammals in that their [[mitral cell]]s are distributed throughout the [[outer plexiform layer]] of the [[olfactory bulb]] rather than packed as a [[monolayer]].<ref>{{cite journal|last1=Ashwell|first1=K. W. S.|year=2011|title=Development of the olfactory pathways in platypus and echidna|journal=Brain, Behavior and Evolution|volume=79|issue=1|pages=45–56|doi=10.1159/000332804 |pmid=22156550 }}</ref>
-[[File:Platypus electrolocation.svg|thumb|upright=2<!--for readability of svg text-->|The platypus has secondarily acquired [[Electroreception and electrogenesis|electroreception]]. Its receptors are arranged in stripes on its bill, giving it high sensitivity to the sides and below; it makes quick turns of its head as it swims to detect prey.<ref name="Electro1"/>]]
-[[Monotremes]] are the only mammals (apart from the [[Guiana dolphin]])<ref>{{cite journal |last1=Czech-Damal |first1=Nicole U. |last2=Liebschner |first2=Alexander |last3=Miersch |first3=Lars |last4=Klauer |first4=Gertrud |last5=Hanke |first5=Frederike D. |last6=Marshall |first6=Christopher |last7=Dehnhardt |first7=Guido |last8=Hanke |first8=Wolf |date=22 February 2012 |title=Electroreception in the Guiana dolphin (Sotalia guianensis) |journal=Proceedings of the Royal Society B: Biological Sciences |volume=279 |issue=1729 |pages=663–668 |doi=10.1098/rspb.2011.1127 |pmc=3248726 |pmid=21795271}}</ref> known to have a sense of [[electroreception]], and the platypus's electroreception is the most sensitive of any monotreme.<ref>{{cite journal |last1=Proske |first1=Uwe |last2=Gregory |first2=J. E. |last3=Iggo |first3=A. |year=1998 |title=Sensory receptors in monotremes |journal=Philosophical Transactions of the Royal Society of London |volume=353 |issue=1372 |pages=1187–1198 |doi=10.1098/rstb.1998.0275 |pmc=1692308 |pmid=9720114}}</ref><ref name="Electro1">{{cite journal |last=Pettigrew |first=John D. |year=1999 |title=Electroreception in Monotremes |url=http://jeb.biologists.org/cgi/reprint/202/10/1447.pdf |url-status=live |journal=The Journal of Experimental Biology |volume=202 |issue=Part 10 |pages=1447–54 |doi=10.1242/jeb.202.10.1447 |pmid=10210685 |archive-url=https://web.archive.org/web/20060928054253/http://jeb.biologists.org/cgi/reprint/202/10/1447.pdf |archive-date=28 September 2006 |access-date=19 September 2006 |doi-access=free|bibcode=1999JExpB.202.1447P }}</ref> Feeding by neither sight nor smell,<ref name="draft_genome" /> the platypus closes its eyes, ears, and nose when it dives.<ref>{{cite journal |author=Gregory, J.E. |author2=Iggo, A. |author3=McIntyre, A.K. |author4=Proske, U. |date=June 1988 |title=Receptors in the Bill of the Platypus |journal=Journal of Physiology |volume=400 |issue=1 |pages=349–366 |doi=10.1113/jphysiol.1988.sp017124 |pmc=1191811 |pmid=3418529}}</ref> Digging in the bottom of streams with its bill, its electroreceptors detect tiny electric currents generated by the muscular contractions of its prey, enabling it to distinguish between animate and inanimate objects.<ref name="Electro1" /> Experiments have shown the platypus will even react to an "artificial shrimp" if a small electric current is passed through it.<ref name="Manning">{{cite book |author1=Manning, A. |url=https://archive.org/details/introductiontoan0000mann_m2z8 |title=An Introduction to Animal Behaviour |author2=Dawkins, M.S. |publisher=Cambridge University Press |year=1998 |edition=5th |url-access=registration}}</ref>
+=== Venom ===
+{{Main|Platypus venom}}
-The [[electroreceptor]]s are located in rostrocaudal rows in the skin of the bill, while [[mechanoreceptor]]s for touch are uniformly distributed across the bill. The electrosensory area of the [[cerebral cortex]] is in the tactile [[somatosensory]] area, and some cortical cells receive input from both electroreceptors and mechanoreceptors, suggesting the platypus feels electric fields like touches. These receptors in the bill dominate the [[somatotopic map]] of the platypus brain, in the same way human hands dominate the [[Cortical homunculus|Penfield homunculus map]].<ref name="sensory_platypus">{{cite journal |last1=Pettigrew |first1=John D. |first2=P. R. |last2=Manger |first3=S. L. |last3=Fine |title=The sensory world of the platypus |journal=Philosophical Transactions of the Royal Society of London |pages=1199–1210 |issue=1372 |year=1998 |pmid=9720115|pmc=1692312 |doi=10.1098/rstb.1998.0276 |volume=353 }}</ref><ref name="Ancestors_Tale">{{cite book |first=Richard |last=Dawkins |author-link=Richard Dawkins |title=The Ancestor's Tale, A Pilgrimage to the Dawn of Life |chapter=The Duckbill's Tale |publisher=Houghton Mifflin |location=Boston, Massachusetts |year=2004 |isbn=978-0-618-00583-3 |title-link=The Ancestor's Tale }}</ref>
+[[File:Platypus spur.JPG|right|thumb|The calcaneus spur on the male's hind limb is used to inject venom.]]
-The platypus can feel the direction of an electric source, perhaps by comparing differences in [[signal strength]] across the sheet of electroreceptors, enhanced by the characteristic side-to-side motion of the animal's head while hunting. It may also be able to determine the distance of moving prey from the time lag between their electrical and mechanical pressure pulses.<ref name="Electro1" />
-Monotreme electrolocation for hunting in murky waters may be tied to their tooth loss.<ref name="Masakazu Asahara 2016">Masakazu Asahara; Masahiro Koizumi; Thomas E. Macrini; Suzanne J. Hand; Michael Archer (2016). "Comparative cranial morphology in living and extinct platypuses: Feeding behavior, electroreception, and loss of teeth". Science Advances. 2 (10): e1601329. {{doi|10.1126/sciadv.1601329}}.</ref> The extinct ''[[Obdurodon]]'' was electroreceptive, but unlike the modern platypus it foraged [[pelagic zone|pelagically]].<ref name="Masakazu Asahara 2016" />
+While both male and female platypuses are born with back ankle spurs, only the males retain them into adulthood.<ref name=mammalianspecies/> Similar spurs are found on many archaic mammal groups, indicating that this was an ancient general characteristic among mammals.<ref>{{cite journal|last1=Hurum|first1=J. H|last2=Zhe-Xi|first2=L|last3=Kielan-Jaworowska|first3=Z|year=2006|title=Were mammals originally venomous?|journal=Acta Palaeontologica Polonica|volume=51|issue=1|pages=1–11|url=https://www.app.pan.pl/article/item/app51-001.html}}</ref> The spurs of the male inject venom, which is powerful enough to inflict pain in humans.<ref name=mammalianspecies>{{cite journal|last1=Pasitschniak-Arts|first1=M.|last2=Marinelli|first2=L|year=1998|title=Ornithorhynchus anatinus|journal=Mammalian Species|issue=585 |pages=1–9|doi=10.2307/3504433 |jstor=3504433 }}</ref> Starting from the wounded area, the affect limb develops [[edema]] (swelling via fluid buildup) which can lead to an excruciating [[hyperalgesia]] (heightened sensitivity to pain) that can last as long as months.<ref name="JN">{{cite journal |author=de Plater, G.M. |author2=Milburn, P.J. |author3=Martin, R.L. |year=2001 |title=Venom From the Platypus, ''Ornithorhynchus anatinus'', Induces a Calcium-Dependent Current in Cultured Dorsal Root Ganglion Cells |journal=Journal of Neurophysiology |volume=85 |issue=3 |pages=1340–5 |doi=10.1152/jn.2001.85.3.1340 |pmid=11248005 |s2cid=2452708 }}</ref>
-=== Eyes ===
+The venom is composed largely of [[defensin]]-like [[protein]]s (DLPs) produced by the immune system, some of which are unique to the species.<ref name="PS">{{cite journal| last = Gerritsen| first = Vivienne Baillie| title = Platypus poison| journal = Protein Spotlight| issue = 29| date = December 2002| url = http://www.expasy.org/spotlight/back_issues/sptlt029.shtml| access-date = 14 September 2006| archive-date = 20 October 2008| archive-url = https://web.archive.org/web/20081020054110/http://www.expasy.org/spotlight/back_issues/sptlt029.shtml| url-status = live}}</ref> It is produced in kidney-shaped [[alveolar gland]]s located in each of the thighs of the hind limbs and connected to the spur.<ref name="ABRS" /> The venomous spurs of male platypuses serve as weapons in battles with other males for breeding.<ref name=mammalianspecies/><ref name="JN"/>
-In recent studies it has been suggested that the eyes of the platypus are more similar to those of [[Pacific hagfish]] or Northern Hemisphere [[lamprey]]s than to those of most tetrapods. The eyes also contain [[Double cone (biology)|double cones]], unlike most mammals.<ref>{{cite journal |last1=Zeiss|first1=Caroline|title=Comparative retinal morphology of the platypus|doi=10.1002/jmor.10959 |year=2011 |first2=Ivan R. |last3=Murphy|first3=Christopher J. |last4=Dubielzig|first4=Richard W. |journal=Journal of Morphology |volume=272 |issue=8 |pages=949–57 |pmid=21567446|last2=Schwab|s2cid=28546474}}</ref>
+{{-}}
-Although the platypus's eyes are small and not used under water, several features indicate that vision was important for its ancestors. The [[cornea]]l surface and the adjacent surface of the lens is flat, while the posterior surface of the lens is steeply curved, similar to the eyes of other aquatic mammals such as otters and sea-lions. A [[Temporal bone|temporal]] (ear side) concentration of [[retinal ganglion cell]]s, important for binocular vision, indicates a vestigial role in [[predation]], though the actual visual acuity is insufficient for such activities. Limited acuity is matched by low [[cortical magnification]], a small [[lateral geniculate nucleus]], and a large [[optic tectum]], suggesting that the [[Midbrain tectum|visual midbrain]] plays a more important role than the [[visual cortex]], as in some rodents. These features suggest that the platypus has adapted to an aquatic and nocturnal lifestyle, developing its electrosensory system at the cost of its visual system. This contrasts with the small number of electroreceptors in the [[short-beaked echidna]], which dwells in dry environments, while the [[long-beaked echidna]], which lives in moist environments, is intermediate between the other two monotremes.<ref name="sensory_platypus" />
+==Distribution and habitat==
+The platypus is native to the freshwaters of eastern Australia, from Queenland to Tasmania (including [[King Island (Tasmania)|King Island]] but not the [[Furneaux Group]]).<ref name=mammalianspecies/><ref name="DPIW">{{cite web|url=http://www.dpiw.tas.gov.au/inter.nsf/WebPages/BHAN-53573T?open |title=Platypus |publisher=Department of Primary Industries and Water, Tasmania |date=31 August 2006 |access-date=12 October 2006 |archive-url=https://web.archive.org/web/20061009211345/http://www.dpiw.tas.gov.au/inter.nsf/WebPages/BHAN-53573T?open |archive-date= 9 October 2006 }}</ref> It was believed to be extinct on the [[South Australia]]n mainland, with the last sighting recorded at [[Renmark, South Australia|Renmark]] in 1975.<ref name="Sutton">{{cite web | last=Sutton | first=Malcolm | title=Platypus 'sighting' in the Adelaide Hills sparks camera set-up to capture extinct species - ABC News | website=ABC (Australian Broadcasting Corporation) | date=3 May 2017 | url=https://www.abc.net.au/news/2017-05-04/cameras-set-up-to-confirm-platypus-sighting-in-adelaide-hills/8492400 | access-date=12 October 2020 | archive-date=26 November 2020 | archive-url=https://web.archive.org/web/20201126044708/https://www.abc.net.au/news/2017-05-04/cameras-set-up-to-confirm-platypus-sighting-in-adelaide-hills/8492400 | url-status=live }}</ref> Platypuses were captively bred at [[Warrawong Sanctuary]] in 1990-91.<ref name=Gemmell1995>{{cite journal|last1=Gemmell|first1=N. J.|last2=Grant|first2=T. R.|last3=Western|first3=P. S.|last4=Walmsley|first4=J|last5=Watson|first5=J. M.|last6=Murray|first6=N. D.|last7=Graves|first7=J. A. M.|year=1995|title=Determining Platypus Relationships|journal=Australian Journal of Zoology|volume=43|issue=3|pages=283–291|doi=10.1071/ZO9950283}}</ref> In October 2020 a nesting platypus was filmed in the wild after the previously abandoned Sanctuary reopened.<ref name="Sutton 2020">{{cite web | last=Sutton | first=Malcolm | title=V6 Commodore water pump gets the tick from nesting platypus at Warrawong | website=ABC News | publisher=Australian Broadcasting Corporation | date=1 October 2020 | url=https://www.abc.net.au/news/2020-10-02/playtpus-nesting-in-restored-swamp-at-warrawong/12719346 | access-date=7 October 2020 | archive-date=7 October 2020 | archive-url=https://web.archive.org/web/20201007035918/https://www.abc.net.au/news/2020-10-02/playtpus-nesting-in-restored-swamp-at-warrawong/12719346 | url-status=live }}</ref> There is a population on [[Kangaroo Island]]<ref name="RKI">{{cite web|url=http://www.ees.adelaide.edu.au/research/fbp/ki_res/|archive-url=https://web.archive.org/web/20040706181309/http://www.ees.adelaide.edu.au/research/fbp/ki_res/|archive-date=6 July 2004|title=Research on Kangaroo Island|publisher=University of Adelaide|date=4 July 2006| access-date = 23 October 2006}}</ref> introduced in the 1920s, said to stand at 150 individuals in the Rocky River region of [[Flinders Chase National Park]]. In the [[2019–20 Australian bushfire season]], large portions of the island burned, decimating wildlife. However, [[Department for Environment and Water (South Australia)|SA Department for Environment and Water]] recovery teams worked to restore their habitat, with a number of sightings reported by April 2020.<ref>{{cite web | title=Find out how platypuses are faring on Kangaroo Island following the bushfires | website=Department for Environment and Water | date=7 April 2020 | url=https://www.environment.sa.gov.au/goodliving/posts/2020/04/platypus-recovery-on-ki | access-date=12 October 2020 | archive-date=21 July 2021 | archive-url=https://web.archive.org/web/20210721003955/https://www.environment.sa.gov.au/goodliving/posts/2020/04/platypus-recovery-on-ki | url-status=live }}</ref> The platypus has almost disappeared from the [[Murray–Darling basin|Murray–Darling Basin]], possibly due to poor water management.<ref>{{cite news|last1=Casben|first1=Liv|date=8 Aug 2019|title=Platypus struggle to survive, with huge national decline over last 200 years, research finds|work=ABC News|url=https://www.abc.net.au/news/2019-08-09/platypus-research-indicates-huge-national-population-decline/11394502|access-date=6 July 2025}}</ref> Platypuses can be found in a variety of freshwater habitats including rivers, streams, lakes and lagoon-like pools. The surrounding terrestrial environment includes tropical rainforests and colder alpine areas.<ref name=mammalianspecies/>
-=== Biofluorescence ===
+==Ecology and behaviour==
-In 2020, research revealed that platypus fur gives a bluish-green [[biofluorescent]] glow in [[Blacklight|black light]].<ref>{{cite web|last=November 2020|first=Mindy Weisberger-Senior Writer 02|title=Platypuses glow an eerie blue-green under UV light|url=https://www.livescience.com/platypuses-glow-uv-light.html|access-date=7 November 2020|website=livescience.com|date=2 November 2020 |language=en|archive-date=5 November 2020|archive-url=https://web.archive.org/web/20201105063117/https://www.livescience.com/platypuses-glow-uv-light.html|url-status=live}}</ref>
-==Distribution, ecology, and behaviour==
-[[File:Animaldentition ornithoryncusanatinus.png|thumb|Dentition, as illustrated in Knight's ''Sketches in Natural History'']]
 [[File:Ornithorhynchus anatinus -Sydney Aquarium, Sydney, Australia -swimming-6a.ogv|thumbtime=50|right|thumb|Swimming underwater at [[Sydney Aquarium]], Australia]]
+The platypus is [[semiaquatic]] and requires permanent freshwater habitat.<ref name=mammalianspecies/> Its swimming style is unique among mammals, propelling itself by alternating strokes of each front foot, while the webbed hind feet and tail are used for steering.<ref name="Rowing">{{cite journal|journal=The Journal of Experimental Biology|volume=200|issue=20|pages=2647–52|title=Energetics of Swimming by the Platypus ''Ornithorhynchus anatinus'': Metabolic Effort Associated with Rowing|author=Fish, F.E.|author2=Baudinette, R.V.|author3=Frappell, P.B.|author4=Sarre, M.P.|year=1997|doi=10.1242/jeb.200.20.2647|url=http://jeb.biologists.org/cgi/reprint/200/20/2647.pdf|pmid=9359371|bibcode=1997JExpB.200.2647F |access-date=23 October 2006|archive-date=26 September 2009|archive-url=https://web.archive.org/web/20090926191559/http://jeb.biologists.org/cgi/reprint/200/20/2647.pdf|url-status=live}}</ref> It can maintain its relatively low body temperature when feeding in colder depths of below {{convert|5|C|F}}.<ref name="ABRS" /> In one study, dives lasted on average thirty-five seconds, with surfacing intervals averaging thirteen seconds.<ref>{{cite journal|journal=The Journal of Applied Ecology|jstor=2404239|title=The Diving Behaviour of the Platypus (''Ornithorhynchus anatinus'') in Waters with Different Trophic Status|author=Kruuk, H. |volume=30|issue=4|year=1993|pages=592–8 | doi = 10.2307/2404239|bibcode=1993JApEc..30..592K }}</ref> The species is mainly [[Nocturnality|nocturnal]] but is also active at [[dusk]] during the summer and daytime during winter. A platypus may spend half the day in water and then retreat into its burrow, which is constructed by digging into the [[bank (geography)|bank]]. These vary between simple resting banks and complex nesting/breeding burrows.<ref name=mammalianspecies/><ref name="ABRS" /><ref name="HCP">{{cite journal|journal=Philosophical Transactions: Biological Sciences|title=Field biology of the platypus (''Ornithorhynchus anatinus''): historical and current perspectives|volume=353|issue=1372|author=Grant, T.G.|author2= Temple-Smith, P.D. |year=1998|pmid=9720106|pages=1081–91|pmc=1692311 | doi = 10.1098/rstb.1998.0267}}</ref> It may have a range of up to {{convert|7|km|mi|abbr=on}}, with a male's home range overlapping those of three or four females.<ref name="AJZ4">{{cite journal|journal=Australian Journal of Zoology|title=Spatial-Organization and Movement Patterns of Adult Male Platypus, ''Ornithorhynchus anatinus'' (Monotremata, Ornithorhynchidae) |volume=43 |issue=1 |author= Gardner, J. L. |author2=Serena, M. |year=1995 |pages=91–103 | doi = 10.1071/ZO9950091}}</ref> Platypuses are not very vocal; they have been recorded to [[growling|growl]] when disturbed and squeak when feeling pain.<ref name="sensory_platypus" />
-The platypus is semiaquatic, inhabiting small streams and rivers over an extensive range from the cold highlands of [[Tasmania]] and the [[Australian Alps]] to the [[tropical rainforest]]s of coastal [[Queensland]] as far north as the base of the [[Cape York Peninsula]].<ref name="DPIW">{{cite web|url=http://www.dpiw.tas.gov.au/inter.nsf/WebPages/BHAN-53573T?open |title=Platypus |publisher=Department of Primary Industries and Water, Tasmania |date=31 August 2006 |access-date=12 October 2006 |url-status=dead |archive-url=https://web.archive.org/web/20061009211345/http://www.dpiw.tas.gov.au/inter.nsf/WebPages/BHAN-53573T?open |archive-date= 9 October 2006 }}</ref>
+The platypus is a [[carnivore]] and forages by probing along the [[benthic zone|bottom]]. It feeds on insect [[larva]]e, [[annelid]] worms, [[shrimp]], [[crayfish]], [[Bivalvia|bivalves]], [[tadpole]]s and fish eggs. It stores food in its cheek pouches for later consumption.<ref name=mammalianspecies/><ref name="HCP"/> In captivity, platypuses have survived up to thirty years, and wild specimens have been recaptured at twenty-four years old.<ref>{{cite journal|last1=Serena|first1=M|last2=Snowball|first2=G|last3=Thomas|first3=J. L.|last4=Williams|first4=G. A.|last5=Danger|first5=A|year=2024|title=Platypus longevity: a new record in the wild and information on captive life span|journal=Australian Mammalogy|volume=46|issue=2|article-number=AM23048|doi=10.1071/AM23048|doi-access=free}}</ref> They are preyed upon by eels, snakes, [[goanna]]s, [[Bird of prey|birds of prey]], crocodiles, [[Cats in Australia|feral cats]] and [[Red foxes in Australia|foxes]].<ref name=Grant89/>{{rp|49}} Parasites and viruses also affect their mortality, though platypuses appear to have a high tolerance for them. Externally, platypuses may carry fleas, mites, and ticks, the latter being more prominent in young.<ref name=mammalianspecies/> The platypus is a common host for the tick species ''[[Ixodes ornithorhynchi]]''.<ref>{{cite journal|last1=Gofton|first1=A. W.|last2=Loh|first2=S-M|last3=Barbosa|first3=A. D.|last4=Paparini|first4=A|last5=Gillett|first5=A|last6=Macgregor|first6=J|last7=Oskam|first7=C. L.|last8=Ryan|first8=U. M.|last9=Irwin|first9=P. J.|year=2018|title=A novel Ehrlichia species in blood and ''Ixodes ornithorhynchi'' ticks from platypuses (''Ornithorhynchus anatinus'') in Queensland and Tasmania, Australia|journal=Ticks and Tick Borne Dieases|volume=9|issue=2|pages=435–442|doi=10.1016/j.ttbdis.2017.12.011}}</ref> Internally, the platypus may host [[protozoa]]ns, [[trematode]]s, and [[nematode]]s.<ref name=mammalianspecies/> The fungus ''[[Mucor amphibiorum]]'' has been reported in Tasmanian platypuses, which causes the disease [[mucormycosis]], symptoms of which include skin lesions and [[Ulcer (dermatology)|ulcer]]s along the body.<ref>{{cite journal|last1=Connolly|first1=J. H.|last2=Obendorf|first2=D. L.|last3=Whittington|first3=R. J.|last4=Muir|first4=D. B.|year=1998|title=Causes of morbidity and mortality in platypus (''Ornithorhynchus anatinus'') From Tasmania, with particular reference to ''Mucor amphibiorum'' infection|journal=Australian Mammalogy|volume=20|issue=2|pages=177–187|doi=10.1071/AM98177}}</ref>
-Inland, its distribution is not well known. It was considered extinct on the [[South Australia]]n mainland, with the last sighting recorded at [[Renmark, South Australia|Renmark]] in 1975.<ref name="Sutton">{{cite web | last=Sutton | first=Malcolm | title=Platypus 'sighting' in the Adelaide Hills sparks camera set-up to capture extinct species - ABC News | website=ABC (Australian Broadcasting Corporation) | date=3 May 2017 | url=https://www.abc.net.au/news/2017-05-04/cameras-set-up-to-confirm-platypus-sighting-in-adelaide-hills/8492400 | access-date=12 October 2020 | archive-date=26 November 2020 | archive-url=https://web.archive.org/web/20201126044708/https://www.abc.net.au/news/2017-05-04/cameras-set-up-to-confirm-platypus-sighting-in-adelaide-hills/8492400 | url-status=live }}</ref> In the 1980s, [[John Wamsley]] created a platypus breeding program in [[Warrawong Sanctuary]] (see [[#Platypus in wildlife sanctuaries|below]]), which subsequently closed.<ref>{{cite web | title=Life reinstated to much-loved Warrawong Wildlife Sanctuary | first=Melissa | last=Keogh | website=The Lead SA | date=3 October 2018 | url=http://theleadsouthaustralia.com.au/industries/regional-showcase/life-reinstated-to-much-loved-warrawong-wildlife-sanctuary/ | access-date=12 October 2020 | archive-date=12 October 2020 | archive-url=https://web.archive.org/web/20201012171044/http://theleadsouthaustralia.com.au/industries/regional-showcase/life-reinstated-to-much-loved-warrawong-wildlife-sanctuary/ | url-status=live }}</ref><ref>{{cite news|last=Adams|first=Prue|title=Wamsley walks away from Earth Sanctuaries|work=Landline|publisher=Australian Broadcasting Corporation|url=https://www.abc.net.au/local/archives/landline/content/2005/s1330004.htm|date=27 March 2005|access-date=12 October 2020|archive-date=12 October 2020|archive-url=https://web.archive.org/web/20201012103440/https://www.abc.net.au/local/archives/landline/content/2005/s1330004.htm|url-status=live}}</ref> In 2017 there were some unconfirmed sightings downstream from the sanctuary,<ref name="Sutton"/> and in October 2020 a nesting platypus was filmed inside the recently reopened sanctuary.<ref name="Sutton 2020">{{cite web | last=Sutton | first=Malcolm | title=V6 Commodore water pump gets the tick from nesting platypus at Warrawong | website=ABC News | publisher=Australian Broadcasting Corporation | date=1 October 2020 | url=https://www.abc.net.au/news/2020-10-02/playtpus-nesting-in-restored-swamp-at-warrawong/12719346 | access-date=7 October 2020 | archive-date=7 October 2020 | archive-url=https://web.archive.org/web/20201007035918/https://www.abc.net.au/news/2020-10-02/playtpus-nesting-in-restored-swamp-at-warrawong/12719346 | url-status=live }}</ref>
-There is a population on [[Kangaroo Island]]<ref name="RKI">{{cite web|url=http://www.ees.adelaide.edu.au/research/fbp/ki_res/|archive-url=https://web.archive.org/web/20040706181309/http://www.ees.adelaide.edu.au/research/fbp/ki_res/|url-status=dead|archive-date=6 July 2004|title=Research on Kangaroo Island|publisher=University of Adelaide|date=4 July 2006| access-date = 23 October 2006}}</ref> introduced in the 1920s, said to stand at 150 individuals in the Rocky River region of [[Flinders Chase National Park]]. In the [[2019–20 Australian bushfire season]], large portions of the island burnt, decimating wildlife. However, [[Department for Environment and Water (South Australia)|SA Department for Environment and Water]] recovery teams worked to reinstate their habitat, with a number of sightings reported by April 2020.<ref>{{cite web | title=Find out how platypuses are faring on Kangaroo Island following the bushfires | website=Department for Environment and Water | date=7 April 2020 | url=https://www.environment.sa.gov.au/goodliving/posts/2020/04/platypus-recovery-on-ki | access-date=12 October 2020 | archive-date=21 July 2021 | archive-url=https://web.archive.org/web/20210721003955/https://www.environment.sa.gov.au/goodliving/posts/2020/04/platypus-recovery-on-ki | url-status=live }}</ref>
-The platypus is no longer found in the main [[Murray–Darling basin|Murray–Darling Basin]], possibly due to declining [[water quality]] from land clearing and [[irrigation]]<ref name="CSIRO">{{cite web|url=http://www.clw.csiro.au/publications/technical97/tr23-97.pdf|title=Impacts of water management in the Murray-Darling Basin on the platypus (''Ornithorhynchus anatinus'') and the water rat (''Hydromus chrysogaster'')|author1=Scott, Anthony|author2=Grant, Tom|publisher=CSIRO Australia|date=November 1997|access-date=23 October 2006|archive-date=15 March 2016|archive-url=https://web.archive.org/web/20160315115754/http://www.clw.csiro.au/publications/technical97/tr23-97.pdf|url-status=live}}</ref> although it is found in the [[Goulburn River]] in Victoria.<ref>{{cite web|url=https://platypus.asn.au/platypus-burrows/ |title=Platypus burrows |website=Australian Platypus Conservancy|date=6 March 2024 }}</ref> Along the coastal river systems, its distribution is unpredictable: absent in some relatively healthy rivers, but present in some quite degraded ones, for example the lower [[Maribyrnong River|Maribyrnong]].<ref name="APC2">{{cite web|url=http://www.platypus.asn.au/platypus_in_country_areas.html |archive-url=https://web.archive.org/web/20160917002720/http://www.platypus.asn.au/platypus_in_country_areas.html |archive-date=17 September 2016 |title=Platypus in Country Areas |publisher=Australian Platypus Conservancy |access-date=23 October 2006 |url-status=dead }}</ref>
-In captivity, platypuses have survived to 30 years of age, and wild specimens have been recaptured when 24 years old. [[Mortality rate]]s for adults in the wild appear to be low.<ref>{{cite news|title=Oldest-known wild platypus gives new insights into longevity of enigmatic Australian species|website=ABC Science |author=Peter de Kruijff|url=https://www.abc.net.au/news/science/2024-02-15/new-platypus-age-records-for-captive-and-wild-specimens/103450732|date=14 February 2024}}</ref> Natural predators include [[snake]]s, [[Rakali|water rats]], [[goanna]]s, [[hawk]]s, [[owl]]s, and [[eagle]]s. Low platypus numbers in northern Australia are possibly due to predation by [[crocodile]]s.<ref name="EPA">{{cite web|url=http://www.epa.qld.gov.au/nature_conservation/wildlife/az_of_animals/platypus.html |title=Platypus |publisher=Environmental Protection Agency/Queensland Parks and Wildlife Service |year=2006 |access-date=24 July 2009 |url-status=dead |archive-url=https://web.archive.org/web/20091021042522/http://www.epa.qld.gov.au/nature_conservation/wildlife/az_of_animals/platypus.html |archive-date=21 October 2009 }}</ref> The introduction of [[red fox]]es in 1845 for sport hunting may have had some impact on its numbers on the mainland.<ref name="Workshop" /> The platypus is generally [[nocturnal]] and [[crepuscular]], but can be active on overcast days.<ref name="EC">{{cite web|url=http://www.biology.iastate.edu/InternationalTrips/1Australia/04papers/CromerMonotrRepro.htm |title=Monotreme Reproductive Biology and Behavior |publisher=Iowa State University |author=Cromer, Erica |date=14 April 2004 |access-date=18 June 2009 |url-status=dead |archive-url=https://web.archive.org/web/20090313030236/http://www.biology.iastate.edu/InternationalTrips/1Australia/04papers/CromerMonotrRepro.htm |archive-date=13 March 2009 }}</ref><ref name="HCP">{{cite journal|journal=Philosophical Transactions: Biological Sciences|title=Field biology of the platypus (''Ornithorhynchus anatinus''): historical and current perspectives|volume=353|issue=1372|author=Grant, T.G.|author2= Temple-Smith, P.D. |year=1998|pmid=9720106|pages=1081–91|pmc=1692311 | doi = 10.1098/rstb.1998.0267}}</ref> Its habitat bridges [[river]]s and the [[riparian zone]], where it finds both prey and river banks to dig resting and nesting burrows.<ref name="HCP" /> It may have a range of up to {{convert|7|km|mi|abbr=on}}, with a male's home range overlapping those of three or four females.<ref name="AJZ4">{{cite journal|journal=Australian Journal of Zoology|title=Spatial-Organization and Movement Patterns of Adult Male Platypus, ''Ornithorhynchus anatinus'' (Monotremata, Ornithorhynchidae) |volume=43 |issue=1 |author= Gardner, J. L. |author2=Serena, M. |year=1995 |pages=91–103 | doi = 10.1071/ZO9950091
-}}</ref>
-The platypus is an excellent swimmer and spends much of its time in the water foraging for food. It has a swimming style unique among mammals,<ref name="dpipwe">{{cite web|url=https://dpipwe.tas.gov.au/wildlife-management/fauna-of-tasmania/mammals/echidnas-and-platypus/platypus/platypus-in-tasmania|title=Platypus in Tasmania {{!}} Department of Primary Industries, Parks, Water and Environment, Tasmania|website=dpipwe.tas.gov.au|access-date=10 April 2020|archive-date=8 March 2020|archive-url=https://web.archive.org/web/20200308221910/https://dpipwe.tas.gov.au/wildlife-management/fauna-of-tasmania/mammals/echidnas-and-platypus/platypus/platypus-in-tasmania|url-status=live}}</ref> propelling itself by alternate strokes of the front feet, while the webbed hind feet are held against the body and only used for steering, along with the tail.<ref name="Rowing">{{cite journal|journal=The Journal of Experimental Biology|volume=200|issue=20|pages=2647–52|title=Energetics of Swimming by the Platypus ''Ornithorhynchus anatinus'': Metabolic Effort Associated with Rowing|author=Fish, F.E.|author2=Baudinette, R.V.|author3=Frappell, P.B.|author4=Sarre, M.P.|year=1997|doi=10.1242/jeb.200.20.2647|url=http://jeb.biologists.org/cgi/reprint/200/20/2647.pdf|pmid=9359371|bibcode=1997JExpB.200.2647F |access-date=23 October 2006|archive-date=26 September 2009|archive-url=https://web.archive.org/web/20090926191559/http://jeb.biologists.org/cgi/reprint/200/20/2647.pdf|url-status=live}}</ref> It can maintain its relatively low body temperature of about 32{{spaces}}°C (90{{spaces}}°F) while foraging for hours in water below 5{{spaces}}°C (41{{spaces}}°F).<ref name="ABRS" /> Dives normally last around 30 seconds, with an estimated aerobic limit of 40 seconds, with 10 to 20 seconds at the surface between dives.<ref name="Bethge">{{cite thesis|url=http://eprints.utas.edu.au/2326/|title=Energetics and foraging behaviour of the platypus|publisher=University of Tasmania|author=Philip Bethge|date=April 2002|doi=10.25959/23210588.v1 |access-date=21 June 2009|archive-date=25 October 2018|archive-url=https://web.archive.org/web/20181025070921/https://eprints.utas.edu.au/2326/|type=thesis |url-status=live}}</ref><ref>{{cite journal|journal=The Journal of Applied Ecology|jstor=2404239|title=The Diving Behaviour of the Platypus (''Ornithorhynchus anatinus'') in Waters with Different Trophic Status|author=Kruuk, H. |volume=30|issue=4|year=1993|pages=592–8 | doi = 10.2307/2404239|bibcode=1993JApEc..30..592K }}</ref>
-The platypus rests in a short, straight burrow in the riverbank about {{convert|30|cm|in|0|abbr=on}} above water level, its oval entrance-hole often [[Camouflage|hidden]] under a tangle of roots.<ref name="dpipwe" /> It may sleep up to 14 hours per day, after half a day of diving.<ref>{{cite journal |author=Holland, Jennifer S. |title=40 Winks? |journal= National Geographic|volume=220 |issue=1 |date=July 2011 }}</ref>
-=== Diet ===
-The platypus is a [[carnivore]], feeding on [[annelid]] worms, [[Larva|insect larvae]], freshwater [[shrimp]], and [[Cherax|yabby]] ([[crayfish]]) that it digs out of the riverbed with its snout or catches while swimming. It carries prey to the surface in cheek-pouches before eating it.<ref name="dpipwe" /> It eats about 20% of its own weight each day, which requires it to spend an average of 12 hours daily looking for food.<ref name="Bethge" />
 === Reproduction ===
 [[File:Ornithorhynchus anatinus - nest with eggs - MUSE.JPG|thumb|Platypus's nest with eggs (replica)]]
-The species has a single [[breeding season]] between June and October, with some local variation.<ref name="EPA" /> Investigations have found both resident and transient platypuses, and suggest a [[Polygyny in animals|polygynous]] mating system.<ref name="AJZ2">{{cite journal|journal=Australian Journal of Zoology|title=Aspects of Lactation in the Platypus, ''Ornithorhynchus anatinus'' (Monotremata), in Waters of Eastern New South Wales|author=Grant, T. R. |author2=Griffiths, M. |author3=Leckie, R.M.C. |volume=31|issue=6|pages=881–9|doi=10.1071/ZO9830881|year=1983}}</ref> Females are believed to become sexually mature in their second year, with breeding observed in animals over nine years old.<ref name="AJZ2"/> During [[Copulation (zoology)|copulation]], the male grasps the female's tail with his bill, wraps his tail around her, then grips her neck or shoulder, everts his [[penis]] through his [[cloaca]], and inserts it into her [[urogenital sinus]].<ref>{{Cite book |last=Grant |first=Tom |url=https://books.google.com/books?id=bu8rIm-TdqwC&dq=platypus+copulation&pg=PA17 |title=Platypus |date=2007-10-02 |publisher=Csiro Publishing |isbn=978-0-643-10127-2 |language=en}}</ref> He takes no part in nesting, living in his year-long resting burrow. After mating, the female constructs a deep, elaborate nesting burrow up to {{convert|20|m|ft|round=5|abbr=on}} long.<ref name="ADW">{{cite web |url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Ornithorhynchidae.html |title=Family Ornithorhynchidae (platypus) |author1=Anna Bess Sorin |author2=Phil Myers |name-list-style=amp |publisher=University of Michigan Museum of Zoology |year=2001 |access-date=24 October 2006 |archive-date=10 April 2011 |archive-url=https://web.archive.org/web/20110410142645/http://animaldiversity.ummz.umich.edu/site/accounts/information/Ornithorhynchidae.html |url-status=live }}</ref> She tucks fallen leaves and reeds underneath her curled tail, dragging them to the burrow to soften the tunnel floor with folded wet leaves, and to line the nest at the end with bedding.<ref name="APC" />
+Platypuses are [[seasonal breeder]]s, with the more southerly populations breeding later in the year than those further north; those in New South Wales mate during fall and winter.<ref name="HCP"/> Research has found both resident and transient platypuses, and suggest a [[Polygyny in animals|polygynous]] mating system.<ref name="AJZ2">{{cite journal|journal=Australian Journal of Zoology|title=Aspects of Lactation in the Platypus, ''Ornithorhynchus anatinus'' (Monotremata), in Waters of Eastern New South Wales|author=Grant, T. R. |author2=Griffiths, M. |author3=Leckie, R.M.C. |volume=31|issue=6|pages=881–9|doi=10.1071/ZO9830881|year=1983}}</ref> Females are believed to become sexually mature at two years of age and continue to breed beyond nine years.<ref name="AJZ2"/> During [[Copulation (zoology)|copulation]], the male swims behind the female and grabs the tail with his bill, positioning his own tail under her from the side, then grips her neck or shoulder.<ref name=Grant2007>{{Cite book |last=Grant |first=Tom |url=https://books.google.com/books?id=bu8rIm-TdqwC&dq=platypus+copulation&pg=PA17 |title=Platypus |date=2007-10-02 |publisher=Csiro Publishing |isbn=978-0-643-10127-2 |language=en}}</ref>{{rp|17–19}} The female retreats into a nesting/breeding burrow made of grass, which can be {{convert|30|m|abbr=on}} long, to lay eggs and raise young.<ref name=mammalianspecies/><ref name="ABRS"/><ref name=Handasyde/> Early European [[natural history|naturalists]] did not believe that the female platypus lays eggs, but this was confirmed by [[William Hay Caldwell]] in 1884.<!--for wiring London and other such trivia see that article--><ref name="ABRS"/><ref name="PS"/>
-The male platypus has [[penile spines]] and an asymmetrical [[glans penis]], with the right side smaller than the left.<ref>{{Cite book |last1=Vogelnest |first1=Larry |url=https://books.google.com/books?id=4d6VDwAAQBAJ&dq=platypus&pg=PA64 |title=Current Therapy in Medicine of Australian Mammals |last2=Portas |first2=Timothy |date=2019-05-01 |publisher=Csiro Publishing |isbn=978-1-4863-0752-4 |language=en}}</ref> The female has two [[Ovary|ovaries]], but only the left one is functional.<ref>{{Cite book |last1=Vogelnest |first1=Larry |url=https://books.google.com/books?id=p8cDEAAAQBAJ&pg=PA107 |title=Medicine of Australian Mammals |last2=Woods |first2=Rupert |date=18 August 2008 |publisher=Csiro Publishing |isbn=978-0-643-09797-1 |language=en |access-date=27 April 2024 |archive-date=25 May 2024 |archive-url=https://web.archive.org/web/20240525044226/https://books.google.com/books?id=p8cDEAAAQBAJ&pg=PA107#v=onepage&q&f=false |url-status=live }}</ref><ref name="EC" /> She lays one to three (usually two) small, leathery eggs (similar to those of reptiles), about {{convert|11|mm|in|frac=32|abbr=on}} in diameter and slightly rounder than bird eggs.<ref name="BSED">{{cite journal|journal= Philosophical Transactions of the Royal Society B: Biological Sciences|title=Early development and embryology of the platypus|author=Hughes, R. L. |author2=Hall, L. S. |volume=353|issue=1372|pages=1101–14|date=28 July 1998|pmid= 9720108|pmc= 1692305 | doi = 10.1098/rstb.1998.0269}}</ref> The eggs develop ''[[Uterus|in utero]]'' for about 28 days, with only about 10 days of external [[Avian incubation|incubation]] (in contrast to a chicken egg, which spends about one day in tract and 21 days externally).<ref name="EC" /> The female curls around the incubating eggs, which develop in three phases.<ref name="RS2" /> In the first, the [[embryo]] has no functional organs and relies on the [[yolk sac]] for sustenance, until the sac is absorbed.<ref>{{cite web|title=Ockhams Razor|work=The Puzzling Platypus|url=http://www.abc.net.au/radionational/programs/ockhamsrazor/the-puzzling-platypus/3492688|access-date=2 December 2006|date=20 July 2001|archive-date=9 August 2017|archive-url=https://web.archive.org/web/20170809103030/http://www.abc.net.au/radionational/programs/ockhamsrazor/the-puzzling-platypus/3492688|url-status=live}}</ref> During the second phase, the digits develop, and in the last phase, the [[egg tooth]] appears.<ref name="RS2">{{cite journal|journal=Philosophical Transactions: Biological Sciences|title =The development of the external features of the platypus (''Ornithorhynchus anatinus'')|author=Manger, Paul R. |author2=Hall, Leslie S. |author3=Pettigrew, John D. |volume=353|issue=1372|pages=1115–25|date=29 July 1998|pmid=9720109|pmc=1692310|doi =10.1098/rstb.1998.0270}}</ref> At first, European [[natural history|naturalists]] could hardly believe that the female platypus lays eggs, but this was finally confirmed by [[William Hay Caldwell]] in 1884.<!--for wiring London and other such trivia see that article--><ref name="ABRS"/><ref name="PS"/>
-Most mammal zygotes go through [[holoblastic]] cleavage, splitting into multiple divisible daughter cells. However, [[monotremes]] like the platypus, along with [[Sauropsida|reptiles and birds]], undergo meroblastic cleavage, in which the ovum does not split completely. The cells at the edge of the yolk remain continuous with the egg's cytoplasm, allowing the yolk and embryo to exchange waste and nutrients with the egg through the cytoplasm.<ref>{{cite journal|author=Myers, P. Z.|url=http://www.nature.com/scitable/topicpage/interpreting-shared-characteristics-the-platypus-genome-44568|title=Interpreting Shared Characteristics: The Platypus Genome|journal=Nature Education|volume=1|issue=1|page=462008|year=2008|access-date=26 March 2015|archive-date=4 March 2018|archive-url=https://web.archive.org/web/20180304124410/http://www.nature.com/scitable/topicpage/interpreting-shared-characteristics-the-platypus-genome-44568|url-status=live}}</ref>
+Most mammal zygotes go through [[holoblastic]] cleavage, splitting into several divisible daughter cells. However, [[monotremes]] such as the platypus, along with [[Sauropsida|reptiles and birds]], undergo [[meroblastic]] cleavage, in which the [[ovum]] does not split completely. The cells at the edge of the [[yolk]] remain continuous with the egg's [[cytoplasm]], allowing the yolk and embryo to exchange waste and nutrients with the egg through the cytoplasm.<ref>{{cite journal|author=Myers, P. Z.|url=http://www.nature.com/scitable/topicpage/interpreting-shared-characteristics-the-platypus-genome-44568|title=Interpreting Shared Characteristics: The Platypus Genome|journal=Nature Education|volume=1|issue=1|page=462008|year=2008|access-date=26 March 2015|archive-date=4 March 2018|archive-url=https://web.archive.org/web/20180304124410/http://www.nature.com/scitable/topicpage/interpreting-shared-characteristics-the-platypus-genome-44568|url-status=live}}</ref><ref name="BSED">{{cite journal|journal= Philosophical Transactions of the Royal Society B: Biological Sciences|title=Early development and embryology of the platypus|author=Hughes, R. L. |author2=Hall, L. S. |volume=353|issue=1372|pages=1101–14|date=28 July 1998|pmid= 9720108|pmc= 1692305 | doi = 10.1098/rstb.1998.0269}}</ref> The female normally lays two small, leathery eggs about {{convert|17|mm|in|frac=32|abbr=on}} long. The eggs develop ''[[Uterus|in utero]]'' for about twenty-eight days, followed by a ten day external [[Avian incubation|incubation]].<ref name="EC" /> The female curls around the incubating eggs,<ref name=Grant89/>{{rp|34}} as the embryo continues to develop.<ref name="BSED"/> Newly hatched platypuses are vulnerable; blind and hairless, they are fed by the mother's milk, which is thicker than in placental mammals and provides all the requirements for growth and development.<ref name="ABRS"/><ref>{{cite journal|last1=Stannard|first1=Hayley J.|last2=Miller|first2=Robert D.|last3=Old|first3=Julie M.|title=Marsupial and monotreme milk – a review of its nutrients and immune properties|journal=PeerJ|year=2020|volume=8|article-number=e9335|doi=10.7717/peerj.9335 |pmid=32612884 |pmc=7319036 |doi-access=free }}</ref><ref name="RS2">{{cite journal|journal=Philosophical Transactions: Biological Sciences|title =The development of the external features of the platypus (''Ornithorhynchus anatinus'')|author=Manger, Paul R. |author2=Hall, Leslie S. |author3=Pettigrew, John D. |volume=353|issue=1372|pages=1115–25|date=29 July 1998|pmid=9720109|pmc=1692310|doi =10.1098/rstb.1998.0270}}</ref> With no teats, the milk is released through pores in the skin from which the young lap it up in her fur.<ref name="QM"/> The offspring are milk-fed for around four months; a minority of young are weaned after they exit the burrow but usually within five days.<ref name=Handasyde>{{cite journal|last1=Thomas|first1=J. L.|last2=Parrott|first2=M. L.|last3=Handasyde|first3=K. A.|last4=Temple-Smith|first4=P|year=2020|title=Maternal care of platypus nestlings (''Ornithorhynchus anatinus'')|journal=Australian Mammalogy|volume=42|issue=3|pages=283–292|doi=10.1071/AM19019}}</ref> Hatched young also have a remnant yolk sac which disappears within four days.<ref name="RS2"/>
-Young platypus are called "puggles". Newly hatched platypuses are vulnerable, blind, and hairless, and are fed by the mother's milk, that provides all the requirements for growth and development.<ref>{{cite web|last1=Stannard|first1=Hayley J.|last2=Old|first2=Julie M.|title=Wallaby joeys and platypus puggles are tiny and undeveloped when born. But their mother's milk is near-magical|url=https://theconversation.com/wallaby-joeys-and-platypus-puggles-are-tiny-and-undeveloped-when-born-but-their-mothers-milk-is-near-magical-207726|publisher=The Conversation|year=2023|access-date=25 October 2023|archive-date=10 November 2023|archive-url=https://web.archive.org/web/20231110202925/https://theconversation.com/wallaby-joeys-and-platypus-puggles-are-tiny-and-undeveloped-when-born-but-their-mothers-milk-is-near-magical-207726|url-status=live}}</ref><ref>{{cite journal|last1=Stannard|first1=Hayley J.|last2=Miller|first2=Robert D.|last3=Old|first3=Julie M.|title=Marsupial and monotreme milk – a review of its nutrients and immune properties|journal=PeerJ|year=2020|volume=8|page=e9335|doi=10.7717/peerj.9335 |pmid=32612884 |pmc=7319036 |doi-access=free }}</ref> The platypus's [[mammary gland]]s lack [[teat]]s, with milk released through pores in the skin. The milk pools in grooves on the mother's abdomen, allowing the young to lap it up.<ref name="APC" /><ref name="EPA" /> After they hatch, the offspring are milk-fed for three to four months.
+During incubation and weaning, the mother initially leaves the burrow only for short periods to forage. She leaves behind her a number of thin soil plugs along the length of the burrow; pushing past these on her return squeezes water from her fur and allows the burrow to remain dry.<ref name="QM">{{cite web|url=http://www.qm.qld.gov.au/inquiry/factsheets/leaflet0010.pdf|title=Egg-laying mammals|publisher=Queensland Museum|date=November 2000| access-date = 19 June 2009|archive-url = https://web.archive.org/web/20080722180447/http://www.qm.qld.gov.au/inquiry/factsheets/leaflet0010.pdf |archive-date = 22 July 2008}}</ref><ref name=Handasyde/> The female spends less time with her offspring after five weeks, who emerge from the burrow around four months.<ref name="EPA">{{cite web|url=http://www.epa.qld.gov.au/nature_conservation/wildlife/az_of_animals/platypus.html |title=Platypus |publisher=Environmental Protection Agency/Queensland Parks and Wildlife Service |year=2006 |access-date=24 July 2009 |archive-url=https://web.archive.org/web/20091021042522/http://www.epa.qld.gov.au/nature_conservation/wildlife/az_of_animals/platypus.html |archive-date=21 October 2009 }}</ref> By then, they are fully covered in fur and may be around 67% the weight of an adult and 80% the length.<ref name="ABRS"/> Juvenile males [[Biological dispersal|disperse]] further than females.<ref>{{cite journal|last1=Bino|first1=G|last2=Grant|first2=T|last3=Kingsford|first3=R|year=2015|title=Life history and dynamics of a platypus (''Ornithorhynchus anatinus'') population: four decades of mark-recapture surveys|journal=Scientific Reports|volume=5|article-number=16073|doi=10.1038/srep16073|pmid=26536832|pmc=4633588|bibcode=2015NatSR...516073B|hdl=1959.4/unsworks_36987|hdl-access=free}}</ref> They are fully grown at around two years.<ref name=Grant2007/>{{rp|28}}
-During incubation and weaning, the mother initially leaves the burrow only for short periods to forage. She leaves behind her a number of thin soil plugs along the length of the burrow, possibly to protect the young from predators; pushing past these on her return squeezes water from her fur and allows the burrow to remain dry.<ref name="QM">{{cite web|url=http://www.qm.qld.gov.au/inquiry/factsheets/leaflet0010.pdf|title=Egg-laying mammals|publisher=Queensland Museum|date=November 2000| access-date = 19 June 2009|archive-url = https://web.archive.org/web/20080722180447/http://www.qm.qld.gov.au/inquiry/factsheets/leaflet0010.pdf |archive-date = 22 July 2008|url-status=dead}}</ref> After about five weeks, the mother begins to spend more time away from her young, and at around four months, the young emerge from the burrow.<ref name="EPA" /> A platypus is born with teeth, but these drop out at a very early age, leaving the horny plates it uses to grind food.<ref name="Piper-2007">{{cite book |author-link=Ross Piper |first=Ross |last=Piper |title=Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals |url=https://archive.org/details/extraordinaryani0000pipe |url-access=registration |publisher=[[Greenwood Press (publisher)|Greenwood Press]] |year=2007 |isbn=978-0-313-33922-6 }}</ref>
 == Evolution ==
@@ Line 170: / Line 133: @@
 }}
-The platypus and other monotremes were very poorly understood, and some of the 19th century myths that grew up around them{{snd}}for example, that the monotremes were "inferior" or [[Reptile|quasireptilian]]{{snd}}still endure.<ref name="Rodent">{{cite journal|journal=Philosophical Transactions: Biological Sciences|title=The platypus is not a rodent: DNA hybridization, amniote phylogeny and the palimpsest theory|author=Kirsch, John A. W. |author2=Mayer, Gregory C. |volume=353|issue=1372|pages=1221–37|date=29 July 1998|pmid=9720117|pmc=1692306|doi=10.1098/rstb.1998.0278}}</ref> In 1947, [[William King Gregory]] theorised that placental mammals and marsupials may have diverged earlier, and a subsequent branching divided the monotremes and marsupials, but later research and fossil discoveries have suggested this is incorrect.<ref name="Rodent" /><ref name="JM">{{cite journal |journal=Nature |volume=416 |issue= 6877|pages=165–8 |year=2002 |doi=10.1038/416165a |title=The first Jurassic mammal from South America |author=Rauhut, O.W.M.|author2= Martin, T.|author3= Ortiz-Jaureguizar, E.|author4= Puerta, P. |pmid=11894091|bibcode=2002Natur.416..165R|hdl=11336/99461 |s2cid=4346804 |hdl-access=free }}</ref> In fact, modern monotremes are the survivors of an early branching of the mammal tree, and a later branching is thought to have led to the [[marsupial]] and placental groups.<ref name="Rodent" /><ref name="JME">{{cite journal|journal=Journal of Mammalian Evolution|title=Evolution of the Monotremes: Phylogenetic Relationship to Marsupials and Eutherians, and Estimation of Divergence Dates Based on α-Lactalbumin Amino Acid Sequences|author=Messer, M.|author2= Weiss, A.S.|author3= Shaw, D.C.|author4= Westerman, M.|volume=5|issue=1|pages=95–105|date=March 1998 | doi = 10.1023/A:1020523120739|s2cid=39638466}}</ref> [[Molecular clock]] and fossil dating suggest platypuses split from [[echidna]]s around 19–48{{spaces}}million years ago.<ref name="Phillips">{{cite journal | author = Phillips MJ|author2= Bennett TH|author3= Lee MS | year = 2009 | title = Molecules, morphology, and ecology indicate a recent, amphibious ancestry for echidnas | journal =Proc. Natl. Acad. Sci. U.S.A. | volume = 106 | issue = 40| pages = 17089–94 | doi = 10.1073/pnas.0904649106 | pmid = 19805098 | pmc = 2761324 | bibcode = 2009PNAS..10617089P |doi-access= free}}</ref>
+In separate publications in 1934, 1947, and 1951, [[William King Gregory]] theorised that placental mammals and marsupials may have diverged earlier, and a subsequent branching divided the monotremes and marsupials. Later research and fossil discoveries have suggested this is incorrect.<ref name="Rodent">{{cite journal|journal=Philosophical Transactions: Biological Sciences|title=The platypus is not a rodent: DNA hybridization, amniote phylogeny and the palimpsest theory|author=Kirsch, John A. W. |author2=Mayer, Gregory C. |volume=353|issue=1372|pages=1221–37|date=29 July 1998|pmid=9720117|pmc=1692306|doi=10.1098/rstb.1998.0278}}</ref><ref name="JM">{{cite journal |journal=Nature |volume=416 |issue= 6877|pages=165–8 |year=2002 |doi=10.1038/416165a |title=The first Jurassic mammal from South America |author=Rauhut, O.W.M.|author2= Martin, T.|author3= Ortiz-Jaureguizar, E.|author4= Puerta, P. |pmid=11894091|bibcode=2002Natur.416..165R|hdl=11336/99461 |s2cid=4346804 |hdl-access=free }}</ref> Modern monotremes are the survivors of an early branching of the mammal tree, and a later branching is thought to have led to the [[marsupial]] and placental groups.<ref name="Rodent" /><ref name="JME">{{cite journal|journal=Journal of Mammalian Evolution|title=Evolution of the Monotremes: Phylogenetic Relationship to Marsupials and Eutherians, and Estimation of Divergence Dates Based on α-Lactalbumin Amino Acid Sequences|author=Messer, M.|author2= Weiss, A.S.|author3= Shaw, D.C.|author4= Westerman, M.|volume=5|issue=1|pages=95–105|date=March 1998 | doi = 10.1023/A:1020523120739|s2cid=39638466}}</ref> Both [[molecular clock]] and fossil dating suggest that the platypus split from [[echidna]]s around 19–48{{spaces}}million years ago.<ref name="Phillips">{{cite journal | author = Phillips MJ|author2= Bennett TH|author3= Lee MS | year = 2009 | title = Molecules, morphology, and ecology indicate a recent, amphibious ancestry for echidnas | journal =Proc. Natl. Acad. Sci. U.S.A. | volume = 106 | issue = 40| pages = 17089–94 | doi = 10.1073/pnas.0904649106 | pmid = 19805098 | pmc = 2761324 | bibcode = 2009PNAS..10617089P |doi-access= free}}</ref>
 [[File:Steropodon BW.jpg|thumb|right|Reconstruction of ancient platypus relative ''Steropodon'']]
-The oldest discovered fossil of the modern platypus dates back to about 100,000 years ago during the [[Quaternary]] period, though a limb bone of ''Ornithorhynchus'' is known from [[Pliocene]]-aged strata.<ref>{{cite book|author=Musser, A.M.|editor1=Merrick, J.R.|editor2=Archer, M.|editor3=Hickey, G.M.|editor4=Lee, M.S.Y.|year=2006|chapter=26. Furry Egg-layers: Monotreme Relationships and Radiations|title=Evolution and Biogeography of Australasian Vertebrates|publisher=Auscipub|pages=523–550|isbn=978-0975779002}}</ref> The extinct monotremes ''[[Teinolophos]]'' and ''[[Steropodon]]'' from the [[Cretaceous]] were once thought to be closely related to the modern platypus,<ref name="JM" /> but are now considered more basal taxa.<ref name="Thomas H. Rich 2016">{{Cite journal |last1=Rich |first1=Thomas H. |last2=Hopson |first2=James A. |last3=Gill |first3=Pamela G. |last4=Trusler |first4=Peter |last5=Rogers-Davidson |first5=Sally |last6=Morton |first6=Steve |last7=Cifelli |first7=Richard L. |last8=Pickering |first8=David |last9=Kool |first9=Lesley |year=2016 |title=The mandible and dentition of the Early Cretaceous monotreme Teinolophos trusleri |journal=Alcheringa: An Australasian Journal of Palaeontology |volume=40 |issue=4 |pages=475–501 |doi=10.1080/03115518.2016.1180034 |bibcode=2016Alch...40..475R |s2cid=89034974 |hdl=1885/112071 |hdl-access=free }}</ref> The fossilised ''Steropodon'' was discovered in [[New South Wales]] and is composed of an opalised lower jawbone with three molar teeth (whereas the adult contemporary platypus is toothless). The molar teeth were initially thought to be [[Tribosphenic molar|tribosphenic]], which would have supported a variation of Gregory's theory, but later research has suggested, while they have three cusps, they evolved under a separate process.<ref name="MS">{{cite journal|journal=Acta Palaeontologica Polonica|url=http://www.app.pan.pl/archive/published/app47/app47-487.pdf|title=New data on the Paleocene monotreme ''Monotrematum sudamericanum'', and the convergent evolution of triangulate molars|author=Pascual, R.|author2=Goin, F.J.|author3=Balarino, L.|author4=Udrizar Sauthier, D.E.|volume=47|issue=3|pages=487–492|year=2002|access-date=18 March 2009|archive-date=9 August 2017|archive-url=https://web.archive.org/web/20170809023141/http://www.app.pan.pl/archive/published/app47/app47-487.pdf|url-status=live}}</ref> The fossil jaw of ''Teinolophos'' is thought to be about 110{{spaces}}million years old, making it the oldest mammal fossil found in Australia. Unlike the modern platypus (and [[echidna]]s), ''Teinolophos'' lacked a beak.<ref name="Thomas H. Rich 2016" />
+The oldest discovered fossil of the modern platypus dates back to about 100,000 years ago during the [[Quaternary]] period, though a limb bone of ''Ornithorhynchus'' is known from [[Pliocene]]-epoch strata.<ref>{{cite book|author=Musser, A.M.|editor1=Merrick, J.R.|editor2=Archer, M.|editor3=Hickey, G.M.|editor4=Lee, M.S.Y.|year=2006|chapter=26. Furry Egg-layers: Monotreme Relationships and Radiations|title=Evolution and Biogeography of Australasian Vertebrates|publisher=Auscipub|pages=523–550|isbn=978-0-9757790-0-2}}</ref> The extinct monotremes ''[[Teinolophos]]'', ''[[Steropodon]]'' and ''[[Kollikodon]]'' from the [[Cretaceous]] period are considered to be basal to the platypus and echidnas.<ref>{{cite journal|last1=Pain|first1=R|last2=Archer|first2=M|last3=Hand|first3=S. J.|last4=Beck|first4=R. M. D.|last5=Cody|first5=A|year=2016|title=The upper dentition and relationships of the enigmatic Australian Cretaceous mammal ''Kollikodon ritchiei''|journal=Memoirs of Museum Victoria|volume=74|pages=97–105|doi=10.24199/j.mmv.2016.74.10|hdl=1959.4/unsworks_41916|hdl-access=free}}</ref> The remains of ''Steropodon'' were discovered in [[New South Wales]], composed of an opalised lower jawbone with three molar teeth (whereas the adult contemporary platypus is toothless). The molar teeth were initially thought to be [[Tribosphenic molar|tribosphenic]], which would have supported a variation of Gregory's theory, but later research has suggested that, while they have three cusps, they evolved under a separate process.<ref name="MS">{{cite journal|journal=Acta Palaeontologica Polonica|url=http://www.app.pan.pl/archive/published/app47/app47-487.pdf|title=New data on the Paleocene monotreme ''Monotrematum sudamericanum'', and the convergent evolution of triangulate molars|author=Pascual, R.|author2=Goin, F.J.|author3=Balarino, L.|author4=Udrizar Sauthier, D.E.|volume=47|issue=3|pages=487–492|year=2002|access-date=18 March 2009|archive-date=9 August 2017|archive-url=https://web.archive.org/web/20170809023141/http://www.app.pan.pl/archive/published/app47/app47-487.pdf|url-status=live}}</ref> The fossil jaw of ''Teinolophos'' is elongated but unlike the modern platypus (and [[echidna]]s), lacks a beak.<ref name="Thomas H. Rich 2016">{{Cite journal |last1=Rich |first1=Thomas H. |last2=Hopson |first2=James A. |last3=Gill |first3=Pamela G. |last4=Trusler |first4=Peter |last5=Rogers-Davidson |first5=Sally |last6=Morton |first6=Steve |last7=Cifelli |first7=Richard L. |last8=Pickering |first8=David |last9=Kool |first9=Lesley |year=2016 |title=The mandible and dentition of the Early Cretaceous monotreme Teinolophos trusleri |journal=Alcheringa: An Australasian Journal of Palaeontology |volume=40 |issue=4 |pages=475–501 |doi=10.1080/03115518.2016.1180034 |bibcode=2016Alch...40..475R |s2cid=89034974 |hdl=1885/112071 |hdl-access=free }}</ref>
-In 2024, [[Late Cretaceous]] ([[Cenomanian]])-aged fossil specimens of actual early platypus relatives were recovered from the same rocks as ''Steropodon'', including the [[Basal (phylogenetics)|basal]] ''[[Opalios]]'' and the more derived ''[[Dharragarra]]'', the latter of which may be the oldest member of the platypus [[Stem group|stem]]-lineage, as it retains the same [[Dentition|dental formula]] found in Cenozoic platypus relatives.<ref name=":0">{{cite journal |last1=Flannery |first1=Timothy F. |last2=McCurry |first2=Matthew R. |last3=Rich |first3=Thomas H. |last4=Vickers-Rich |first4=Patricia |last5=Smith |first5=Elizabeth T. |last6=Helgen |first6=Kristofer M. |title=A diverse assemblage of monotremes (Monotremata) from the Cenomanian Lightning Ridge fauna of New South Wales, Australia |journal=Alcheringa: An Australasian Journal of Palaeontology |date=2 April 2024 |volume=48 |issue=2 |pages=319–337 |doi=10.1080/03115518.2024.2348753 |doi-access=free |bibcode=2024Alch...48..319F }}</ref> ''[[Monotrematum]]'' and ''[[Patagorhynchus]]'', two other fossil relatives of the platypus, are known from the latest Cretaceous ([[Maastrichtian]]) and the mid-[[Paleocene]] of [[Argentina]], indicating that some monotremes managed to colonize [[South America]] from Australia when the two continents were connected via [[Antarctica]]. These are also considered potential members of the platypus stem-lineage.<ref name="MS" /><ref name="patagonia">{{Cite journal|title = A platypus in Patagonia (Ancient life – 1992)|last = Folger|first = Tim| date = 1993 |journal = Discover|volume = 14|issue = 1|page = 66}}</ref><ref name=":0" />  The closest fossil relative of the platypus was ''[[Obdurodon]]'', known from the late [[Oligocene]] to the [[Miocene]] of Australia. It closely resembled the modern platypus, aside from the presence of molar teeth.<ref name=":0" /> A fossilised tooth of the giant platypus ''[[Obdurodon tharalkooschild]]'' was dated 5–15{{spaces}}million years ago. Judging by the tooth, the animal measured 1.3 metres long, making it the largest platypus on record.<ref>{{cite news|url = http://www.zmescience.com/science/geology/platypus-fossil-paleontology-05112013/|title = 'Platypus-zilla' fossil unearthed in Australia|newspaper = ZME Science|last = Mihai|first = Andrei|year = 2013|access-date = 5 November 2013|archive-date = 21 July 2021|archive-url = https://web.archive.org/web/20210721004004/https://www.zmescience.com/science/geology/platypus-fossil-paleontology-05112013/|url-status = live}}</ref>
+In 2024, [[Late Cretaceous]] ([[Cenomanian]])-aged fossil specimens of early platypus relatives were recovered from the same rocks as ''Steropodon'', including the [[Basal (phylogenetics)|basal]] ''[[Opalios]]'' and the more derived ''[[Dharragarra]]'', the latter of which may be the oldest member of the platypus family Ornithorhynchidae, as it retains the same [[Dentition|dental formula]] found in Cenozoic platypus relatives.<ref name=":0">{{cite journal |last1=Flannery |first1=Timothy F. |last2=McCurry |first2=Matthew R. |last3=Rich |first3=Thomas H. |last4=Vickers-Rich |first4=Patricia |last5=Smith |first5=Elizabeth T. |last6=Helgen |first6=Kristofer M. |title=A diverse assemblage of monotremes (Monotremata) from the Cenomanian Lightning Ridge fauna of New South Wales, Australia |journal=Alcheringa: An Australasian Journal of Palaeontology |date=2 April 2024 |volume=48 |issue=2 |pages=319–337 |doi=10.1080/03115518.2024.2348753 |doi-access=free |bibcode=2024Alch...48..319F }}</ref> ''[[Monotrematum]]'' and ''[[Patagorhynchus]]'', also fossil relatives of the platypus, are known from the latest Cretaceous ([[Maastrichtian]]) and the mid-[[Paleocene]] of [[Argentina]], indicating that some monotremes managed to colonize [[South America]] from Australia when the two continents were connected via [[Antarctica]]. These are also considered potential members of the platypus Ornithorhynchidae.<ref name="MS" /><ref name="patagonia">{{Cite journal|title = A platypus in Patagonia (Ancient life – 1992)|last = Folger|first = Tim| date = 1993 |journal = Discover|volume = 14|issue = 1|page = 66}}</ref><ref name=":0" /> The closest fossil relative of the platypus is ''[[Obdurodon]]'', known from the late [[Oligocene]] to the [[Miocene]] of Australia. It closely resembled the modern platypus, aside from the presence of molar teeth.<ref name=":0" /> A fossilised tooth of the giant platypus ''[[Obdurodon tharalkooschild]]'' was dated 5–15{{spaces}}million years ago. Judging by the tooth, the animal measured 1.3 metres long, making it the largest platypus known.<ref>{{cite news|url = http://www.zmescience.com/science/geology/platypus-fossil-paleontology-05112013/|title = 'Platypus-zilla' fossil unearthed in Australia|newspaper = ZME Science|last = Mihai|first = Andrei|year = 2013|access-date = 5 November 2013|archive-date = 21 July 2021|archive-url = https://web.archive.org/web/20210721004004/https://www.zmescience.com/science/geology/platypus-fossil-paleontology-05112013/|url-status = live}}</ref><ref>{{cite journal|last1=Pian|first1=R|last2=Archer|first2=M|last3=Hand|first3=S. J.|year=2013|title=A new, giant platypus, ''Obdurodon tharalkooschild'', sp. nov. (Monotremata, Ornithorhynchidae), from the Riversleigh World Heritage Area, Australia|journal=Journal of Vertebrate Paleontology|volume=33|issue=6|pages=1255–1259|doi=10.1080/02724634.2013.782876|bibcode=2013JVPal..33.1255P}}</ref>
-The loss of teeth in the modern platypus has long been enigmatic, as a distinctive lower molar tooth row was previously present in its lineage for over 95 million years. Even its closest relative, ''Obdurodon'', which otherwise closely resembles the platypus, retained this tooth row. More recent studies indicate that this tooth loss was a geologically very recent event, occurring only around the [[Pliocene|Plio]]-[[Pleistocene]] (around 2.5 million years ago), when the [[rakali]], a large semiaquatic [[rodent]], colonized Australia from [[New Guinea]]. The platypus, which previously fed on a wide array of hard and soft-bodied prey, was outcompeted by the rakali for hard-bodied prey such as crayfish and mussels. This competition may have selected for the loss of teeth in the platypus and their replacement by horny pads, as a way of specializing for softer-bodied prey, over which the rakali did not compete with it.<ref name=":0" />
+The loss of teeth in the modern platypus has long been enigmatic, as a distinctive lower molar tooth row was present in its lineage for over 95 million years. Even its closest relative, ''Obdurodon'', which otherwise closely resembles the platypus, retained this tooth row. More recent studies indicate that this tooth loss was geologically a very recent event, occurring only around the [[Pliocene|Plio]]-[[Pleistocene]] about 2.5 million years ago, when the [[rakali]], a large semiaquatic [[rodent]], colonized Australia from [[New Guinea]]. The platypus, which previously fed on a wide array of hard and soft-bodied prey, was outcompeted by the rakali for hard-bodied prey such as crayfish and mussels. This competition may have selected for the loss of teeth in the platypus and their replacement by horny pads, as a way of specializing for softer-bodied prey, over which the rakali did not compete.<ref name=":0" />
 === Genome ===
-[[File:Platypus skeleton Pengo.jpg|thumb|Platypus skeleton]]
+Because of the early divergence from the [[Theria|therian mammals]] and the low numbers of extant monotreme species, the platypus is a frequent subject of research in evolutionary biology. In 2004, researchers at the [[Australian National University]] discovered that the platypus has ten [[sex chromosome]]s, compared with two (XY) in most other mammals. These ten chromosomes form five unique pairs of XY in males and XX in females, i.e. males are X{{sub|1}}Y{{sub|1}}X{{sub|2}}Y{{sub|2}}X{{sub|3}}Y{{sub|3}}X{{sub|4}}Y{{sub|4}}X{{sub|5}}Y{{sub|5}}. One of the X chromosomes of the platypus has close homology to the bird [[Z chromosome]].<ref>{{Cite journal|author1=Frank Grützner|author2=Willem Rens|author3=Enkhjargal Tsend-Ayush|author4=Nisrine El-Mogharbel|author5=Patricia C. M. O'Brien|author6=Russell C. Jones|author7=Malcolm A. Ferguson-Smith|author8=Jennifer A. Marshall Graves|title=In the platypus a meiotic chain of ten sex chromosomes shares genes with the bird Z and mammal X chromosomes|journal=Nature|volume=432|issue=7019|pages=913–917|date=16 December 2004|doi=10.1038/nature03021|pmid=15502814|bibcode=2004Natur.432..913G|s2cid=4379897}}
+</ref> The platypus genome also has both reptilian and mammalian genes associated with egg fertilisation.<ref name="draft_genome">{{cite journal| journal = Nature| volume = 453| issue = 7192| pages = 175–183| title = Genome analysis of the platypus reveals unique signatures of evolution| date = 8 May 2008| doi = 10.1038/nature06936| author = Warren, Wesley C.| pmid = 18464734| pmc = 2803040 |display-authors=etal|bibcode=2008Natur.453..175W}}</ref><ref>{{cite journal |title=Beyond the Platypus Genome – 2008 Boden Research Conference |journal=Reprod Fertil Dev |volume=21 |issue=8 |pages=i–ix, 935–1027 |year=2009 |url=http://www.publish.csiro.au/nid/44/issue/4849.htm |access-date=3 March 2012 |archive-date=21 November 2015 |archive-url=https://web.archive.org/web/20151121130525/http://www.publish.csiro.au/nid/44/issue/4849.htm |url-status=live }}</ref> Though the platypus lacks the mammalian sex-determining gene [[SRY]], a study found that the mechanism of sex determination is the [[Anti-Müllerian hormone|AMH gene]] on the oldest [[Y chromosome]].<ref>{{cite journal|last1=Cortez|first1=Diego|last2=Marin|first2=Ray|last3=Toledo-Flores|first3=Deborah|last4=Froidevaux|first4=Laure|last5=Liechti|first5=Angélica|last6=Waters|first6=Paul D.|last7=Grützner|first7=Frank|last8=Kaessmann|first8=Henrik|year=2014|title=Origins and functional evolution of Y chromosomes across mammals|journal=Nature|volume=508|issue=7497|pages=488–493|doi=10.1038/nature13151|pmid=24759410|bibcode=2014Natur.508..488C|s2cid=4462870}}</ref><ref>{{cite news|last1=Salleh|first1=Anna|title=Platypus Sex 'Master Switch' Identified|url=http://www.abc.net.au/science/articles/2014/05/05/3994897.htm|publisher=Australian Broadcasting Corporation|date=5 May 2014|access-date=5 June 2014|archive-date=6 July 2016|archive-url=https://web.archive.org/web/20160706211627/http://www.abc.net.au/science/articles/2014/05/05/3994897.htm|url-status=live}}</ref> A draft version of the platypus genome sequence was published in ''[[Nature (journal)|Nature]]'' in May 2008, revealing both reptilian and mammalian elements, as well as two genes previously found only in non-mammalian vertebrates. More than 80% of the platypus's genes exist in other amniotes whose genomes have been compared.<ref name="draft_genome" /> An updated genome, the most complete on record, was published in 2021, together with the genome of the [[short-beaked echidna]].<ref>{{cite journal |last1=Zhou |first1=Yang |last2=Shearwin-Whyatt |first2=Linda |last3=Li |first3=Jing |last4=Song |first4=Zhenzhen |last5=Hayakawa |first5=Takashi |last6=Stevens |first6=David |last7=Fenelon |first7=Jane C. |last8=Peel |first8=Emma |last9=Cheng |first9=Yuanyuan |last10=Pajpach |first10=Filip |last11=Bradley |first11=Natasha |last12=Suzuki |first12=Hikoyu |last13=Nikaido |first13=Masato |last14=Damas |first14=Joana |last15=Daish |first15=Tasman |last16=Perry |first16=Tahlia |last17=Zhu |first17=Zexian |last18=Geng |first18=Yuncong |last19=Rhie |first19=Arang |last20=Sims |first20=Ying |last21=Wood |first21=Jonathan |last22=Haase |first22=Bettina |last23=Mountcastle |first23=Jacquelyn |last24=Fedrigo |first24=Olivier |last25=Li |first25=Qiye |last26=Yang |first26=Huanming |last27=Wang |first27=Jian |last28=Johnston |first28=Stephen D. |last29=Phillippy |first29=Adam M. |last30=Howe |first30=Kerstin |last31=Jarvis |first31=Erich D. |last32=Ryder |first32=Oliver A. |last33=Kaessmann |first33=Henrik |last34=Donnelly |first34=Peter |last35=Korlach |first35=Jonas |last36=Lewin |first36=Harris A. |last37=Graves |first37=Jennifer |last38=Belov |first38=Katherine |last39=Renfree |first39=Marilyn B. |last40=Grutzner |first40=Frank |last41=Zhou |first41=Qi |last42=Zhang |first42=Guojie |title=Platypus and echidna genomes reveal mammalian biology and evolution |journal=Nature |date=29 April 2021 |volume=592 |issue=7856 |pages=756–762 |doi=10.1038/s41586-020-03039-0 |pmid=33408411 |pmc=8081666 |bibcode=2021Natur.592..756Z |doi-access=free }}</ref>
-Because of the early divergence from the [[Theria|therian mammals]] and the low numbers of extant monotreme species, the platypus is a frequent subject of research in evolutionary biology. In 2004, [[research]]ers at the [[Australian National University]] discovered the platypus has ten [[sex chromosome]]s, compared with two (XY) in most other mammals. These ten chromosomes form five unique pairs of XY in males and XX in females, i.e. males are X{{sub|1}}Y{{sub|1}}X{{sub|2}}Y{{sub|2}}X{{sub|3}}Y{{sub|3}}X{{sub|4}}Y{{sub|4}}X{{sub|5}}Y{{sub|5}}.<ref name="discover">{{cite web|url=http://discovermagazine.com/2005/apr/sex-ys-platypuses0425/|title=Sex, Ys, and Platypuses|publisher=Discover|first=Jocelyn|last=Selim|date=25 April 2005|access-date=7 May 2008|archive-date=16 May 2008|archive-url=https://web.archive.org/web/20080516233225/http://discovermagazine.com/2005/apr/sex-ys-platypuses0425|url-status=live}}</ref> One of the X chromosomes of the platypus has great homology to the bird Z chromosome.<ref>{{Cite journal|author1=Frank Grützner|author2=Willem Rens|author3=Enkhjargal Tsend-Ayush|author4=Nisrine El-Mogharbel|author5=Patricia C. M. O'Brien|author6=Russell C. Jones|author7=Malcolm A. Ferguson-Smith|author8=Jennifer A. Marshall Graves|title=In the platypus a meiotic chain of ten sex chromosomes shares genes with the bird Z and mammal X chromosomes|journal=Nature|volume=432|issue=7019|pages=913–917|date=16 December 2004|doi=10.1038/nature03021|pmid=15502814|bibcode=2004Natur.432..913G|s2cid=4379897}}
-</ref> The platypus genome also has both reptilian and mammalian genes associated with egg fertilisation.<ref name="draft_genome">{{cite journal| journal = Nature| volume = 453| issue = 7192| pages = 175–183| title = Genome analysis of the platypus reveals unique signatures of evolution| date = 8 May 2008| doi = 10.1038/nature06936| author = Warren, Wesley C.| pmid = 18464734| pmc = 2803040 |display-authors=etal|bibcode=2008Natur.453..175W}}</ref><ref>{{cite journal |title=Beyond the Platypus Genome – 2008 Boden Research Conference |journal=Reprod Fertil Dev |volume=21 |issue=8 |pages=i–ix, 935–1027 |year=2009 |url=http://www.publish.csiro.au/nid/44/issue/4849.htm |access-date=3 March 2012 |archive-date=21 November 2015 |archive-url=https://web.archive.org/web/20151121130525/http://www.publish.csiro.au/nid/44/issue/4849.htm |url-status=live }}</ref> Though the platypus lacks the mammalian sex-determining gene [[SRY]], a study found that the mechanism of sex determination is the [[Anti-Müllerian hormone|AMH gene]] on the oldest [[Y chromosome]].<ref>{{cite journal|last1=Cortez|first1=Diego|last2=Marin|first2=Ray|last3=Toledo-Flores|first3=Deborah|last4=Froidevaux|first4=Laure|last5=Liechti|first5=Angélica|last6=Waters|first6=Paul D.|last7=Grützner|first7=Frank|last8=Kaessmann|first8=Henrik|year=2014|title=Origins and functional evolution of Y chromosomes across mammals|journal=Nature|volume=508|issue=7497|pages=488–493|doi=10.1038/nature13151|pmid=24759410|bibcode=2014Natur.508..488C|s2cid=4462870}}</ref><ref>{{cite news|last1=Salleh|first1=Anna|title=Platypus Sex 'Master Switch' Identified|url=http://www.abc.net.au/science/articles/2014/05/05/3994897.htm|publisher=Australian Broadcasting Corporation|date=5 May 2014|access-date=5 June 2014|archive-date=6 July 2016|archive-url=https://web.archive.org/web/20160706211627/http://www.abc.net.au/science/articles/2014/05/05/3994897.htm|url-status=live}}</ref> A draft version of the platypus genome sequence was published in ''[[Nature (journal)|Nature]]'' on 8{{spaces}}May 2008, revealing both reptilian and mammalian elements, as well as two genes found previously only in birds, amphibians, and fish. More than 80% of the platypus's genes are common to the other mammals whose genomes have been sequenced.<ref name="draft_genome" /> An updated genome, the most complete on record, was published in 2021, together with the genome of the [[short-beaked echidna]].<ref>{{cite journal |last1=Zhou |first1=Yang |last2=Shearwin-Whyatt |first2=Linda |last3=Li |first3=Jing |last4=Song |first4=Zhenzhen |last5=Hayakawa |first5=Takashi |last6=Stevens |first6=David |last7=Fenelon |first7=Jane C. |last8=Peel |first8=Emma |last9=Cheng |first9=Yuanyuan |last10=Pajpach |first10=Filip |last11=Bradley |first11=Natasha |last12=Suzuki |first12=Hikoyu |last13=Nikaido |first13=Masato |last14=Damas |first14=Joana |last15=Daish |first15=Tasman |last16=Perry |first16=Tahlia |last17=Zhu |first17=Zexian |last18=Geng |first18=Yuncong |last19=Rhie |first19=Arang |last20=Sims |first20=Ying |last21=Wood |first21=Jonathan |last22=Haase |first22=Bettina |last23=Mountcastle |first23=Jacquelyn |last24=Fedrigo |first24=Olivier |last25=Li |first25=Qiye |last26=Yang |first26=Huanming |last27=Wang |first27=Jian |last28=Johnston |first28=Stephen D. |last29=Phillippy |first29=Adam M. |last30=Howe |first30=Kerstin |last31=Jarvis |first31=Erich D. |last32=Ryder |first32=Oliver A. |last33=Kaessmann |first33=Henrik |last34=Donnelly |first34=Peter |last35=Korlach |first35=Jonas |last36=Lewin |first36=Harris A. |last37=Graves |first37=Jennifer |last38=Belov |first38=Katherine |last39=Renfree |first39=Marilyn B. |last40=Grutzner |first40=Frank |last41=Zhou |first41=Qi |last42=Zhang |first42=Guojie |title=Platypus and echidna genomes reveal mammalian biology and evolution |journal=Nature |date=29 April 2021 |volume=592 |issue=7856 |pages=756–762 |doi=10.1038/s41586-020-03039-0 |pmid=33408411 |pmc=8081666 |bibcode=2021Natur.592..756Z |doi-access=free }}</ref>
 == Conservation ==
-=== Status and threats ===
+The [[International Union for Conservation of Nature]] categorised the platypus as "near threatened" in 2016,<ref name="iucn"/> based on estimates that numbers had fallen by about thirty percent on average since [[European settlement of Australia|European settlement]]. Other biologists have shown concern that the estimates of the 2016 baseline numbers could be wrong, and numbers may have been reduced by as much as fifty percent.<ref name=natgeog/> The species was hunted for its fur until the early years of the 20th century. Although the species gained legal protections beginning in Victoria in 1890<ref name=1890act>{{cite web | title=Game Act 1890 | website=[[Australasian Legal Information Institute]] | url=http://classic.austlii.edu.au/au/legis/vic/hist_act/ga189043/ | access-date=9 August 2023 | archive-date=9 August 2023 | archive-url=https://web.archive.org/web/20230809184631/http://classic.austlii.edu.au/au/legis/vic/hist_act/ga189043/ |date=10 July 1890 | url-status=live }}</ref> and throughout Australia by 1912,<ref name="unsw">{{cite web |title=Platypus Conservation Initiative |url=https://www.unsw.edu.au/research/platypus-conservation-initiative/the-platypus |url-status=live |archive-url=https://web.archive.org/web/20230519032354/https://www.unsw.edu.au/research/platypus-conservation-initiative/the-platypus |archive-date=19 May 2023 |access-date=9 August 2023 |website=[[University of New South Wales]]}}</ref> it continues to drown in the nets of inland [[fisheries]].<ref name="CSIRO">{{cite web|url=http://www.clw.csiro.au/publications/technical97/tr23-97.pdf|title=Impacts of water management in the Murray-Darling Basin on the platypus (''Ornithorhynchus anatinus'') and the water rat (''Hydromus chrysogaster'')|author1=Scott, Anthony|author2=Grant, Tom|publisher=CSIRO Australia|date=November 1997|access-date=23 October 2006|archive-date=15 March 2016|archive-url=https://web.archive.org/web/20160315115754/http://www.clw.csiro.au/publications/technical97/tr23-97.pdf|url-status=live}}</ref> The use of "opera house traps" by recreational fishers for catching [[Common yabby|yabbies]] is banned in the ACT, South Australia, Tasmania and Victoria, and restricted in NSW and Queensland, due to the traps drowning non-targeted species including platypuses.<ref>{{cite web|url=https://kb.rspca.org.au/knowledge-base/can-enclosed-yabby-traps-harm-animals-and-are-they-legal-to-use/|title=Can enclosed yabby traps harm animals and are they legal to use?|website=RSPCA Australia|access-date=25 April 2025}}</ref> The platypus was listed as endangered is South Australia, under the ''[[National Parks and Wildlife Act 1972]]''. In November 2020 a recommendation was made to list the platypus as a vulnerable species across all states.<ref name="2020report">{{cite web | title=A national assessment of the conservation status of the platypus | website=[[Australian Conservation Foundation]] | url=https://www.acf.org.au/platypus_report | date=23 November 2020 | access-date=28 November 2020 | archive-date=28 November 2020 | archive-url=https://web.archive.org/web/20201128041842/https://www.acf.org.au/platypus_report }}</ref> In January 2021, Victoria officially adopted the vulnerable species designation, under the state's ''[[Flora and Fauna Guarantee Act 1988]]''.<ref name="riskassessment">{{cite web |title=Platypus Risk Assessment |url=https://www.environment.vic.gov.au/__data/assets/pdf_file/0023/595040/Platypus-Risk-Assessment-Report.pdf |url-status=live |archive-url=https://web.archive.org/web/20230327161147/https://www.environment.vic.gov.au/__data/assets/pdf_file/0023/595040/Platypus-Risk-Assessment-Report.pdf |archive-date=27 March 2023 |access-date=9 August 2023 |website=[[Department of Energy, Environment and Climate Action]]}}</ref> The platypus is not covered under the federal ''[[EPBC Act]]''.<ref name=natgeog>{{cite web | title=The silent decline of the platypus, Australia's beloved oddity | website=National Geographic | first=Christie | last=Wilcox | date=29 August 2019 | url=https://www.nationalgeographic.com/animals/2019/08/common-platypus-disappearing-australia/ | access-date=12 October 2020 | archive-date=12 October 2020 | archive-url=https://web.archive.org/web/20201012141601/https://www.nationalgeographic.com/animals/2019/08/common-platypus-disappearing-australia/ }}</ref><ref>{{cite web | title=EPBC Act List of Threatened Fauna | website=Species Profile and Threats Database | publisher=Australian Government. [[Department of Agriculture, Water and the Environment]] | url=https://www.environment.gov.au/cgi-bin/sprat/public/publicthreatenedlist.pl | access-date=12 October 2020 | archive-date=5 November 2020 | archive-url=https://web.archive.org/web/20201105231657/http://www.environment.gov.au/cgi-bin/sprat/public/publicthreatenedlist.pl | url-status=live }}</ref>
-Except for its loss from the state of South Australia, the platypus occupies the same general distribution as it did prior to [[European settlement of Australia]]. However, local changes and fragmentation of distribution due to human modification of its habitat are documented. Its historical abundance is unknown and its current abundance difficult to gauge, but it is assumed to have declined in numbers, although as of 1998 was still being considered as common over most of its current range.<ref name="HCP" /> The species was extensively hunted for its fur until the early years of the 20th century. Although the species gained legal protections beginning in Victoria in 1890<ref name=1890act>{{cite web | title=Game Act 1890 | website=[[Australasian Legal Information Institute]] | url=http://classic.austlii.edu.au/au/legis/vic/hist_act/ga189043/ | access-date=9 August 2023 | archive-date=9 August 2023 | archive-url=https://web.archive.org/web/20230809184631/http://classic.austlii.edu.au/au/legis/vic/hist_act/ga189043/ |date=10 July 1890 | url-status=live }}</ref> and throughout Australia by 1912,<ref name="unsw">{{cite web |title=Platypus Conservation Initiative |url=https://www.unsw.edu.au/research/platypus-conservation-initiative/the-platypus |url-status=live |archive-url=https://web.archive.org/web/20230519032354/https://www.unsw.edu.au/research/platypus-conservation-initiative/the-platypus |archive-date=19 May 2023 |access-date=9 August 2023 |website=[[University of New South Wales]]}}</ref> until about 1950 it was still at risk of drowning in the nets of inland [[fisheries]].<ref name="CSIRO" /> The use of opera house traps for catching [[Common yabby|yabbies]] by recreational fishers is banned in the ACT, South Australia, Tasmania and Victoria, and restricted in NSW and Queensland, due to causing the drowning death of platypuses and other non-target species.<ref>[https://kb.rspca.org.au/knowledge-base/can-enclosed-yabby-traps-harm-animals-and-are-they-legal-to-use/ Can enclosed yabby traps harm animals and are they legal to use?] RSPCA Australia, 2025. Retrieved 25 April 2025.</ref>
-The [[International Union for Conservation of Nature]] recategorised its status as "near threatened" in 2016.<ref name="iucn"/> The species is protected by law, but the only state in which it is listed as endangered is South Australia, under the ''[[National Parks and Wildlife Act 1972]]''. In November 2020 a recommendation was made to list the platypus as a vulnerable species across all states<ref name="2020report">{{cite web | title=A national assessment of the conservation status of the platypus | website=[[Australian Conservation Foundation]] | url=https://www.acf.org.au/platypus_report | date=23 November 2020 | access-date=28 November 2020 | archive-date=28 November 2020 | archive-url=https://web.archive.org/web/20201128041842/https://www.acf.org.au/platypus_report | url-status=live }}</ref> with a vulnerable listing being made official in Victoria under the state's ''[[Flora and Fauna Guarantee Act 1988]]'' on 10 January 2021.<ref name="riskassessment">{{cite web |title=Platypus Risk Assessment |url=https://www.environment.vic.gov.au/__data/assets/pdf_file/0023/595040/Platypus-Risk-Assessment-Report.pdf |url-status=live |archive-url=https://web.archive.org/web/20230327161147/https://www.environment.vic.gov.au/__data/assets/pdf_file/0023/595040/Platypus-Risk-Assessment-Report.pdf |archive-date=27 March 2023 |access-date=9 August 2023 |website=[[Department of Energy, Environment and Climate Action]]}}</ref>
-==== Habitat destruction ====
-The platypus is not considered to be in immediate danger of extinction, because conservation measures have been successful, but it could be adversely affected by habitat disruption caused by [[dam]]s, [[irrigation]], [[pollution]], netting, and trapping.<ref>{{cite journal | last1=Stannard | first1=H.J. | last2=Wolfenden | first2=J. | last3=Old | first3=J.M. | year=2010 | title=Evaluating the capacity of constructed wetlands to sustain a captive population of Platypus (''Ornithorhynchus anatinus'') | journal=Australasian Journal of Environmental Management | volume=17 | issue=1 | pages=27–34 | doi=10.1080/14486563.2010.9725246 | bibcode=2010AuJEM..17...27S }}</ref> Reduction of watercourse flows and water levels through excessive droughts and extraction of water for industrial, agricultural, and domestic supplies are also considered a threat. The [[IUCN]] lists the platypus on its [[Red List]] as "[[Near Threatened]]"<ref name="iucn" /> as assessed in 2016, when it was estimated that numbers had reduced by about 30 percent on average since European settlement. The animal is listed as endangered in South Australia, but it is not covered at all under the federal ''[[EPBC Act]]''.<ref name=natgeog>{{cite web | title=The silent decline of the platypus, Australia's beloved oddity | website=National Geographic | first=Christie | last=Wilcox | date=29 August 2019 | url=https://www.nationalgeographic.com/animals/2019/08/common-platypus-disappearing-australia/ | access-date=12 October 2020 | archive-date=12 October 2020 | archive-url=https://web.archive.org/web/20201012141601/https://www.nationalgeographic.com/animals/2019/08/common-platypus-disappearing-australia/ | url-status=dead }}</ref><ref>{{cite web | title=EPBC Act List of Threatened Fauna | website=Species Profile and Threats Database | publisher=Australian Government. [[Department of Agriculture, Water and the Environment]] | url=https://www.environment.gov.au/cgi-bin/sprat/public/publicthreatenedlist.pl | access-date=12 October 2020 | archive-date=5 November 2020 | archive-url=https://web.archive.org/web/20201105231657/http://www.environment.gov.au/cgi-bin/sprat/public/publicthreatenedlist.pl | url-status=live }}</ref>
-Researchers have worried for years that declines have been greater than assumed.<ref name=natgeog/> In January 2020, researchers from the [[University of New South Wales]] presented evidence that the platypus is at risk of [[extinction]], due to a combination of extraction of [[water resource]]s, [[land clearing]], [[climate change]] and severe [[drought]].<ref name="EA-20200121">{{cite web |author=University of New South Wales |title=Platypus on brink of extinction |url=https://www.eurekalert.org/pub_releases/2020-01/uons-pob011920.php |date=21 January 2020 |work=[[EurekAlert!]] |access-date=22 January 2020 |author-link=University of New South Wales |archive-date=13 May 2020 |archive-url=https://web.archive.org/web/20200513103618/https://www.eurekalert.org/pub_releases/2020-01/uons-pob011920.php |url-status=live }}</ref><ref name=brink2020>{{cite press release |title=Platypus on brink of extinction |url=https://www.sciencedaily.com/releases/2020/01/200121112922.htm |work=ScienceDaily |publisher=University of New South Wales |date=21 January 2020 }}</ref> The study predicted that, considering current threats, the animals' abundance would decline by 47–66% and [[metapopulation]] occupancy by 22–32% over 50 years, causing "extinction of local populations across about 40% of the range". Under projections of climate change projections to 2070, reduced habitat due to drought would lead to 51–73% reduced abundance and 36–56% reduced metapopulation occupancy within 50 years respectively. These predictions suggested that the species would fall under the "Vulnerable" classification. The authors stressed the need for national conservation efforts, which might include conducting more surveys, tracking trends, reduction of threats and improvement of river management to ensure healthy platypus habitat.<ref>{{cite journal |last1=Bino |first1=Gilad |last2=Kingsford |first2=Richard T. |last3=Wintle |first3=Brendan A. |title=A stitch in time – Synergistic impacts to platypus metapopulation extinction risk |journal=Biological Conservation |date=February 2020 |volume=242 |pages=108399 |doi=10.1016/j.biocon.2019.108399 |bibcode=2020BCons.24208399B }}</ref> Co-author Gilad Bino is concerned that the estimates of the 2016 baseline numbers could be wrong, and numbers may have been reduced by as much as half already.<ref name=natgeog/>
+The platypus continues to be adversely affected by habitat disruption caused by [[dam]]s, [[pollution]], [[urban expansion]], and [[urban runoff]].<ref>{{cite journal | last1=Stannard | first1=H.J. | last2=Wolfenden | first2=J. | last3=Old | first3=J.M. | year=2010 | title=Evaluating the capacity of constructed wetlands to sustain a captive population of Platypus (''Ornithorhynchus anatinus'') | journal=Australasian Journal of Environmental Management | volume=17 | issue=1 | pages=27–34 | doi=10.1080/14486563.2010.9725246 | bibcode=2010AuJEM..17...27S }}</ref> Droughts and the demands for water for human use are also considered threats.<ref name="iucn" /> In January 2020, researchers from the [[University of New South Wales]] presented evidence that the platypus is at risk of [[extinction]], due to factors such as water extraction, [[land clearing]], [[climate change]], and invasive species. The study predicted that, considering current threats, the animals' abundance would drop by 47–66% and [[metapopulation]] occupancy by 22–32% over fifty years, causing "extinction of local populations across about 40% of the range". Using climate change projections to 2070, reduced habitat due to drought would lead to 51–73% lower abundance and 36–56% lower metapopulation occupancy after at least fifty years. These predictions suggested that the species would fall under the "Vulnerable" classification. The authors stressed the need for national conservation efforts to ensure healthy platypus habitat, which may include conducting more surveys and tracking trends, as well as better river management while reducing threats.<ref>{{cite journal |last1=Bino |first1=Gilad |last2=Kingsford |first2=Richard T. |last3=Wintle |first3=Brendan A. |title=A stitch in time – Synergistic impacts to platypus metapopulation extinction risk |journal=Biological Conservation |date=February 2020 |volume=242 |article-number=108399 |doi=10.1016/j.biocon.2019.108399 |bibcode=2020BCons.24208399B }}</ref>
-A November 2020 report by scientists from the [[University of New South Wales]], funded by a research grant from the [[Australian Conservation Foundation]] in collaboration with the [[World Wildlife Fund|World Wildlife Fund Australia]] and the [[Humane Society International Australia]] revealed that that platypus habitat in Australia had shrunk by 22 per cent in the previous 30 years, and recommended that the platypus should be listed as a threatened species under the ''EPBC Act''.<ref name=2020report/><ref>{{cite web | last=Cox | first=Lisa | title=Australia's platypus habitat has shrunk 22% in 30 years, report says | website=the Guardian | date=23 November 2020 | url=http://www.theguardian.com/environment/2020/nov/23/australias-platypus-habitat-has-shrunk-22-in-30-years-report-says | access-date=28 November 2020 | archive-date=28 November 2020 | archive-url=https://web.archive.org/web/20201128065816/https://www.theguardian.com/environment/2020/nov/23/australias-platypus-habitat-has-shrunk-22-in-30-years-report-says | url-status=live }}</ref> Declines in population had been greatest in NSW, in particular in the [[Murray–Darling basin]].<ref>{{cite web | title=Platypus should be listed as a threatened species: new report | website=UNSW Newsroom | publisher=[[University of New South Wales]] | date=23 November 2020 | url=https://newsroom.unsw.edu.au/news/science-tech/platypus-should-be-listed-threatened-species-new-report | access-date=28 November 2020 | archive-date=26 November 2020 | archive-url=https://web.archive.org/web/20201126093055/https://newsroom.unsw.edu.au/news/science-tech/platypus-should-be-listed-threatened-species-new-report | url-status=live | author1=z3525972 }}</ref><ref name=2020report/>
+A November 2020 report by scientists from the [[University of New South Wales]], funded by a research grant from the [[Australian Conservation Foundation]] in collaboration with the [[World Wildlife Fund|World Wildlife Fund Australia]] and the [[Humane Society International Australia]] revealed that over the past thirty years platypus habitat in Australia has dropped by 22%, and supported listing the platypus as a threatened species under the ''EPBC Act'', as the declines have been mostly in the [[Murray–Darling basin]] and NSW in general.<ref name=2020report/>
-==== Disease ====
+=== Sanctuaries and captivity ===
-Platypuses generally suffer from few [[disease]]s in the wild; however, as of 2008 there was concern in Tasmania about the potential impacts of a disease caused by the fungus ''[[Mucor amphibiorum]]''. The disease (termed [[mucormycosis]]) affects only Tasmanian platypuses, and had not been observed in platypuses in mainland Australia. Affected platypuses can develop skin lesions or [[ulcer]]s on various parts of their bodies, including their backs, tails, and legs. Mucormycosis can kill platypuses, death arising from secondary infection and by affecting the animals' ability to maintain body temperature and forage efficiently. The Biodiversity Conservation Branch at the Department of Primary Industries and Water collaborated with NRM north and [[University of Tasmania]] researchers to determine the impacts of the disease on Tasmanian platypuses, as well as the mechanism of transmission and spread of the disease.<ref name="DPIW2">{{cite web|url=http://www.dpiw.tas.gov.au/inter.nsf/WebPages/SSKA-7AH66E?open |title=Platypus Fungal Disease |publisher=Department of Primary Industries and Water, Tasmania |date=29 August 2008 |access-date=29 February 2008 |url-status=dead |archive-url=https://web.archive.org/web/20080307100350/http://www.dpiw.tas.gov.au/inter.nsf/WebPages/SSKA-7AH66E?open |archive-date= 7 March 2008 }}</ref>
+Few platypuses have been successfully raised and bred by humans. One notable example is at [[Healesville Sanctuary]] in [[Victoria (Australia)|Victoria]] under [[David Fleay]], where breeding was successful in 1943.<ref name="Zoos Voctoria">{{cite web| url = http://www.zoo.org.au/news/fantastic-fleay-turns-20| title = Fantastic Fleay turns 20!| access-date = 4 February 2014| date = 31 October 2013| work = Zoos Victoria| archive-url = https://web.archive.org/web/20181109084854/https://www.zoo.org.au/news/fantastic-fleay-turns-20| archive-date = 9 November 2018}}</ref><ref name="DF">{{cite web|url=http://www.epa.qld.gov.au/nature_conservation/wildlife/david_fleay_wildlife_park/50_years_wild/david_fleays_achievements/|title= David Fleay's achievements|publisher=Queensland Government|date=23 November 2003| access-date = 13 September 2006|archive-url = https://web.archive.org/web/20061002043523/http://www.epa.qld.gov.au/nature_conservation/wildlife/david_fleay_wildlife_park/50_years_wild/david_fleays_achievements/ |archive-date = 2 October 2006}}</ref><ref name="catalyst"/> More platypuses were successfully bred and raised in 1998 and again in 2000 using a stream tank.<ref name="catalyst"/> Between 2008 and 2012, platypuses were bred regularly at Healesville,<ref>{{cite web| url=https://www.zoo.org.au/news/pitter-patter---platypus-twins| title=Pitter patter – Platypus twins!| publisher=Zoo Victoria| date=4 March 2013| access-date=17 August 2017| archive-url=https://web.archive.org/web/20180828052202/https://www.zoo.org.au/news/pitter-patter---platypus-twins| archive-date=28 August 2018}}</ref> including twins.<ref name=PlatConservZoo>{{cite web | url=https://platypus.asn.au/conservation/zoos/ | title=Zoos | publisher=Australian Platypus Conservancy | access-date=17 August 2017 | date=22 November 2016 | archive-url=https://web.archive.org/web/20190304055829/https://platypus.asn.au/conservation/zoos/ | archive-date=4 March 2019 }}</ref> In 1990-91, there was successful breeding of platypuses at Warrawong Sanctuary.<ref name=Gemmell1995/> [[Taronga Zoo]] in [[Sydney]] bred twins in 2003,<ref name="catalyst">{{cite web|url=http://www.abc.net.au/catalyst/stories/s988946.htm|title=Platypus|publisher=Catalyst|date=13 November 2003|access-date=13 September 2006|archive-date=23 July 2011|archive-url=https://web.archive.org/web/20110723232610/http://www.abc.net.au/catalyst/stories/s988946.htm|url-status=live}}</ref> and the facility has since bred more platypuses to be released into the wild in NSW.<ref>{{cite web|title=First ever platypus translocation nsw results breeding success
+|website=Taronga: Conservation Society Australia| url=https://taronga.org.au/news/2024-03-21/first-ever-platypus-translocation-nsw-results-breeding-success|date=21 March 2024|access-date=19 July 2025}}</ref> {{As of|2019}}, the only platypuses in captivity outside of Australia are in the [[San Diego Zoo Safari Park#Walkabout Australia|San Diego Zoo Safari Park]] in the U.S. state of [[California]].<ref>{{cite web|url=https://www.kpbs.org/news/2019/nov/22/platypus-display-san-diego/|title=Rare Platypus On Display At San Diego Zoo Safari Park|last=Anderson|first=Erik|date=22 November 2019|website=KPBS Public Media|language=en|access-date=29 December 2019|quote=The animals are the only platypuses on display outside of their native country.|archive-date=13 May 2020|archive-url=https://web.archive.org/web/20200513103631/https://www.kpbs.org/news/2019/nov/22/platypus-display-san-diego/|url-status=live}}</ref><ref>{{cite web|url=https://animals.sandiegozoo.org/animals/platypus|title=Platypus {{!}} San Diego Zoo Animals & Plants|website=animals.sandiegozoo.org|access-date=29 December 2019|archive-date=25 July 2020|archive-url=https://web.archive.org/web/20200725024623/https://animals.sandiegozoo.org/animals/platypus|url-status=live}}</ref> Three platypuses were given to the [[Bronx Zoo]] in 1947, two females and a male. One of the females; [[Penelope (platypus)|Penelope]] had a false proto-pregnancy.<ref>{{Cite magazine|title=Science: Penelope's Secret|url=https://time.com/archive/6621597/science-penelopes-secret/|magazine = [[Time (magazine)|Time]]|date=16 November 1953|access-date=19 July 2025}}</ref>
-=== Wildlife sanctuaries ===
-Much of the world was introduced to the platypus in 1939 when ''[[National Geographic Magazine]]'' published an article on the platypus and the efforts to study and raise it in captivity. The latter is a difficult task, and only a few young have been successfully raised since, notably at [[Healesville Sanctuary]] in [[Victoria (Australia)|Victoria]]. The leading figure in these efforts was [[David Fleay]], who established a platypusary (a simulated stream in a tank) at the Healesville Sanctuary, where breeding was successful in 1943.<ref name="Zoos Voctoria">{{cite web| url = http://www.zoo.org.au/news/fantastic-fleay-turns-20| title = Fantastic Fleay turns 20!| access-date = 4 February 2014| date = 31 October 2013| work = Zoos Victoria| archive-url = https://web.archive.org/web/20181109084854/https://www.zoo.org.au/news/fantastic-fleay-turns-20| archive-date = 9 November 2018| url-status = dead}}</ref> In 1972, he found a dead baby of about 50 days old, which had presumably been born in captivity, at his [[David Fleay Wildlife Park|wildlife park]] at [[Burleigh Heads, Queensland|Burleigh Heads]] on the [[Gold Coast, Queensland|Gold Coast]], Queensland.<ref name="DF">{{cite web|url=http://www.epa.qld.gov.au/nature_conservation/wildlife/david_fleay_wildlife_park/50_years_wild/david_fleays_achievements/|title= David Fleay's achievements|publisher=Queensland Government|date=23 November 2003| access-date = 13 September 2006|archive-url = https://web.archive.org/web/20061002043523/http://www.epa.qld.gov.au/nature_conservation/wildlife/david_fleay_wildlife_park/50_years_wild/david_fleays_achievements/ |archive-date = 2 October 2006|url-status=dead}}</ref> Healesville repeated its success in 1998 and again in 2000 with a similar stream tank.<ref name="catalyst"/> Since 2008, platypus has bred regularly at Healesville,<ref>{{cite web| url=https://www.zoo.org.au/news/pitter-patter---platypus-twins| title=Pitter patter – Platypus twins!| publisher=Zoo Victoria| date=4 March 2013| access-date=17 August 2017| archive-url=https://web.archive.org/web/20180828052202/https://www.zoo.org.au/news/pitter-patter---platypus-twins| archive-date=28 August 2018| url-status=dead}}</ref> including second-generation (captive born themselves breeding in captivity).<ref name=PlatConservZoo>{{cite web | url=https://platypus.asn.au/conservation/zoos/ | title=Zoos | publisher=Australian Platypus Conservancy | access-date=17 August 2017 | date=22 November 2016 | archive-url=https://web.archive.org/web/20190304055829/https://platypus.asn.au/conservation/zoos/ | archive-date=4 March 2019 | url-status=dead }}</ref> [[Taronga Zoo]] in [[Sydney]] bred twins in 2003, and breeding was again successful there in 2006.<ref name="catalyst">{{cite web|url=http://www.abc.net.au/catalyst/stories/s988946.htm|title=Platypus|publisher=Catalyst|date=13 November 2003|access-date=13 September 2006|archive-date=23 July 2011|archive-url=https://web.archive.org/web/20110723232610/http://www.abc.net.au/catalyst/stories/s988946.htm|url-status=live}}</ref>
-=== Captivity ===
-As of 2019, the only platypuses in captivity outside of Australia are in the [[San Diego Zoo Safari Park]] in the [[U.S. state]] of [[California]].<ref>{{cite web|url=https://www.kpbs.org/news/2019/nov/22/platypus-display-san-diego/|title=Rare Platypus On Display At San Diego Zoo Safari Park|last=Anderson|first=Erik|date=22 November 2019|website=KPBS Public Media|language=en|access-date=29 December 2019|quote=The animals are the only platypuses on display outside of their native country.|archive-date=13 May 2020|archive-url=https://web.archive.org/web/20200513103631/https://www.kpbs.org/news/2019/nov/22/platypus-display-san-diego/|url-status=live}}</ref><ref>{{cite web|url=https://animals.sandiegozoo.org/animals/platypus|title=Platypus {{!}} San Diego Zoo Animals & Plants|website=animals.sandiegozoo.org|access-date=29 December 2019|archive-date=25 July 2020|archive-url=https://web.archive.org/web/20200725024623/https://animals.sandiegozoo.org/animals/platypus|url-status=live}}</ref> Three attempts were made to bring the animals to the [[Bronx Zoo]], in 1922, 1947, and 1958. Of these, only two of the three animals introduced in 1947, [[Penelope (platypus)|Penelope]] and Cecil,<ref>{{Cite magazine |date=19 August 1957 |title=Animals: End of the Affair |url=http://www.time.com/time/magazine/article/0%2C9171%2C809719%2C00.html |archive-url=https://web.archive.org/web/20070616062446/http://www.time.com/time/magazine/article/0,9171,809719,00.html |archive-date=16 June 2007 |magazine=[[Time (magazine)|Time]]}}</ref> lived longer than eighteen months.<ref>{{cite book | author = Lee S. Crandall | title = The Management of Wild Mammals in Captivity | publisher = University of Chicago Press | date = 1964}}</ref>
 == Human interactions ==
 === Usage ===
-[[Aboriginal Australians]] used to hunt platypuses for food (their fatty tails being particularly nutritious), while, after colonisation, Europeans hunted them for fur from the late 19th century until 1912, when it was prohibited by law.<ref name=1912article>{{cite web | title=Protecting Wild Animals | work=[[The Advertiser (Adelaide)]] | url=https://trove.nla.gov.au/newspaper/article/5338401 | access-date=9 August 2023 | archive-date=9 August 2023 | archive-url=https://web.archive.org/web/20230809172243/https://trove.nla.gov.au/newspaper/article/5338401 |date=27 September 1912 | url-status=live }}</ref> In addition, European researchers captured and killed platypus or removed their eggs, partly in order to increase scientific knowledge, but also to gain prestige and outcompete rivals from different countries.<ref name=2020report/>
+[[Aboriginal Australians]] hunted and ate platypuses, particularly for their fatty nutritious tails, while, after colonisation, Europeans killed them for fur from the late 19th century until 1912, when it was prohibited by law. In addition, European researchers captured and killed platypus or removed their eggs, partly in order to increase scientific knowledge, but also to gain prestige and outcompete rivals from different countries.<ref name=2020report/> During the [[Second World War]], in spite of an export ban,<ref name=bbc>{{cite news|url=https://www.bbc.co.uk/news/articles/cglzl1ez283o|work=[[BBC News]]|title=The mystery of Winston Churchill's dead platypus was unsolved - until now|first=Tiffanie|last=Turnbull|date=2 August 2025|access-date=3 August 2025}}</ref> Australia gave live platypuses as diplomatic gifts to [[Allies of World War II|Allied nations]] as part of an initiative to increase military assistance.<ref>{{cite journal|last1=Cushing|first1=N|last2=Markwell|first2=K|year=2009|title=Platypus diplomacy: animal gifts in international relations|journal=Journal of Australian Studies|volume=33|issue=3|pages=255–271|doi=10.1080/14443050903079664}}</ref> One of which was intended as a gift to [[Winston Churchill]] died from neglect while en route.<ref name=bbc/>
 === Cultural references ===
-The platypus has been a subject in the [[Dreamtime]] stories of [[Aboriginal Australians]], some of whom believed the animal was a hybrid of a duck and a [[Rakali|water rat]].<ref name=Gadi>{{cite book|author1=McKay, Helen F. |author2=McLeod, Pauline E. |author3=Jones, Francis F. |author4=Barber, June E. |year=2001|title=Gadi Mirrabooka: Australian Aboriginal Tales from the Dreaming|publisher=Libraries Unlimited|isbn=978-1563089237}}</ref>{{rp|57–60}}
+The platypus has been a subject in the [[Dreamtime]] stories of Aboriginal Australians, some of whom believed the animal was a hybrid of a duck and a [[Rakali|water rat]].<ref name=Gadi>{{cite book|author1=McKay, Helen F. |author2=McLeod, Pauline E. |author3=Jones, Francis F. |author4=Barber, June E. |year=2001|title=Gadi Mirrabooka: Australian Aboriginal Tales from the Dreaming|publisher=Libraries Unlimited|isbn=978-1-56308-923-7}}</ref>{{rp|57–60}} Aboriginals from the upper [[Darling River]] region have a story of a large water-rat called Biggoon who kidnaps a duck what wandered too far from its tribe. After managing to escape, she returned and laid two eggs which hatched the first platypuses. They were all exiled and went to live in the mountains. In another story from the upper Darling, the major animal groups, the land animals, water animals and birds, all competed for the platypus to join their respective groups, but the platypus ultimately decided to not join any of them, feeling that he did not need to be part of a group to be special,<ref name=Gadi/>{{rp|83–85}} and wished to remain friends with all of those groups.<ref name=2020report/>
-According to one story of the upper [[Darling River]],<ref name=2020report/> the major animal groups, the land animals, water animals and birds, all competed for the platypus to join their respective groups, but the platypus ultimately decided to not join any of them, feeling that he did not need to be part of a group to be special,<ref name=Gadi/>{{rp|83–85}} and wished to remain friends with all of those groups.<ref name=2020report/> Another Dreaming story emanate of the upper Darling tells of a young duck which ventured too far, ignoring the warnings of her tribe, and was kidnapped by a large water-rat called Biggoon. After managing to escape after some time, she returned and laid two eggs which hatched into strange furry creatures, so they were all banished and went to live in the mountains.<ref name=2020report/>
-The platypus is also used by some Aboriginal peoples as a [[totem]], which is to them "a natural object, plant or animal that is inherited by members of a clan or family as their spiritual emblem", and the animal holds special meaning as a totem animal for the [[Wadi Wadi]] people, who live along the [[Murray River]]. Because of their cultural significance and importance in [[connection to country]], the platypus is protected and conserved by these Indigenous peoples.<ref name=2020report/>
-The platypus has often been used as a symbol of Australia's [[cultural identity]]. In the 1940s, live platypuses were given to allies in the [[Second World War]], in order to strengthen ties and boost morale.<ref name=2020report/>
-Platypuses have been used several times as mascots: Syd the platypus was one of the three mascots chosen for the [[2000 Summer Olympics|Sydney 2000 summer Olympics]] along with an echidna and a [[kookaburra]],<ref>{{cite web|url=http://en.beijing2008.com/31/87/article211928731.shtml |title=A Brief History of the Olympic and Paralympic Mascots |publisher=Beijing2008 |date=5 August 2004 |access-date=25 October 2006 |url-status=dead |archive-url=https://web.archive.org/web/20080621113930/http://en.beijing2008.com/31/87/article211928731.shtml |archive-date=21 June 2008 }}</ref> Expo Oz the platypus was the mascot for [[World Expo 88]], which was held in [[Brisbane]] in 1988,<ref>{{cite web|url=http://www.foundationexpo88.org/aboutcontents.html |title=About World Expo '88 |publisher=Foundation Expo '88 |year=1988 |access-date=17 December 2007 |url-status=dead |archive-url=https://web.archive.org/web/20131219232431/http://www.foundationexpo88.org/aboutcontents.html |archive-date=19 December 2013 }}</ref> and Hexley the platypus is the mascot for the [[Darwin (operating system)|Darwin]] operating system, the [[BSD]]-based core of [[macOS]] and other operating systems from [[Apple Inc.]]<ref>{{cite web|url=http://www.hexley.com/|title=The Home of Hexley the Platypus|access-date=25 October 2006|archive-date=13 February 2011|archive-url=https://web.archive.org/web/20110213095652/http://www.hexley.com/|url-status=live}}</ref>
+The platypus is also featured as a [[totem]] for some Aboriginal peoples, which is to them "a natural object, plant or animal that is inherited by members of a clan or family as their spiritual emblem", and the animal holds special meaning for the [[Wadi Wadi]] people at the [[Murray River]]. Because of their cultural significance and importance in [[connection to country]], the platypus is protected and conserved by these Indigenous peoples.<ref name=2020report/> The platypus has often represented Australia's [[cultural identity]] and its image has also been used for stamps and currency and as a mascot in the [[2000 Summer Olympics]] in Sydney.<ref name=2020report/>
-Since the introduction of [[decimal currency]] to [[Australia]] in 1966, the embossed image of a platypus, designed and sculpted by [[Stuart Devlin]], has appeared on the reverse (tails) side of the [[Australian twenty-cent coin|20-cent coin]].<ref>{{cite web|url=https://www.ramint.gov.au/twenty-cents|title=Circulating coins: Twenty Cents|date=8 January 2016 |publisher=Royal Australian Mint|access-date=12 September 2020|archive-date=19 March 2021|archive-url=https://web.archive.org/web/20210319031414/https://www.ramint.gov.au/twenty-cents|url-status=live}}</ref> The platypus has frequently appeared in Australian postage stamps, most recently the 2015 "Native Animals" series and the 2016 "Australian Animals Monotremes" series.<ref>{{cite web|url=https://australiapostcollectables.com.au/stamp-issues/2015-01-13_native-animals.html|title=Native Animals - Issue Date 13 January 2015|publisher=Australia Post Collectables|access-date=12 September 2020|archive-date=25 January 2021|archive-url=https://web.archive.org/web/20210125223959/https://australiapostcollectables.com.au/stamp-issues/2015-01-13_native-animals.html|url-status=live}}</ref><ref>{{cite web|url=https://australiapostcollectables.com.au/stamp-issues/australian-animals-monotremes.html|title=Australian Animals Monotremes – Issue Date 26 September 2016|publisher=Australia Post Collectables|access-date=12 September 2020|archive-date=26 January 2021|archive-url=https://web.archive.org/web/20210126001019/https://australiapostcollectables.com.au/stamp-issues/australian-animals-monotremes.html|url-status=live}}</ref>
-In the American animated series ''[[Phineas and Ferb]]'', the [[title character]]s own a pet bluish-green platypus named [[Perry the Platypus|Perry]] who, unknown to them, is a secret agent. Such choices were inspired by media underuse, as well as to exploit the animal's striking appearance;<ref name="QA">{{cite news|url=https://www.hollywoodreporter.com/news/disney-ferb-pickup-major-push-85063|title=Disney gives 'Ferb' pickup, major push – Q&A: Dan Povenmire|date=7 June 2009|work=The Hollywood Reporter|access-date=5 March 2017|archive-date=20 June 2018|archive-url=https://web.archive.org/web/20180620203720/https://www.hollywoodreporter.com/news/disney-ferb-pickup-major-push-85063|url-status=live}}</ref> additionally, show creator [[Dan Povenmire]], who also wrote the character's theme song, said that its opening lyrics are based on the introductory sentence of the Platypus article on Wikipedia, copying the "semiaquatic egg-laying mammal" phrase word for word and appending the phrase "of action";<ref>{{Citation|title="Perry the Platypus" Live at Musi-Cal| date=9 August 2016 |url=https://www.youtube.com/watch?v=dnGX-wucxQs|language=en|access-date=23 March 2021|archive-date=2 January 2021|archive-url=https://web.archive.org/web/20210102002447/https://www.youtube.com/watch?v=dnGX-wucxQs|url-status=live}}</ref> however, the article did not include "egg-laying mammal" in the lead sentence until 2014, several years after the song released. As a character, Perry has been well received by both fans and critics.<ref name="star">{{cite web|url=https://variety.com/2009/digital/features/phineas-star-perry-makes-mark-on-auds-1118011690/|title='Phineas' star Perry makes mark on auds|author=Littleton, Cynthia|date=20 November 2009|work=[[Variety (magazine)|Variety]]|access-date=26 November 2009|archive-date=2 December 2009|archive-url=https://web.archive.org/web/20091202110736/http://www.variety.com/article/VR1118011690.html?categoryid=1050&cs=1|url-status=live}}</ref><ref>{{cite magazine|url=https://www.pastemagazine.com/blogs/lists/2009/03/7-reasons-why-phineas-and-ferb-is-the-best-kids-sh.html|title=Five Reasons Why ''Phineas and Ferb'' is the Best Kids Show on TV|magazine=[[Paste (magazine)|Paste]]|author=Jackson, John|date=31 March 2009|access-date=25 November 2009|archive-date=3 October 2019|archive-url=https://web.archive.org/web/20191003091539/https://www.pastemagazine.com/blogs/lists/2009/03/7-reasons-why-phineas-and-ferb-is-the-best-kids-sh.html|url-status=live}}</ref> Coincidentally, real platypuses show a similar cyan colour when seen under ultraviolet lighting.<ref>{{cite journal |last1=Anich |first1=Paula Spaeth |last2=Anthony |first2=Sharon |last3=Carlson |first3=Michaela |last4=Gunnelson |first4=Adam |last5=Kohler |first5=Allison M. |last6=Martin |first6=Jonathan G. |last7=Olson |first7=Erik R. |title=Biofluorescence in the platypus (''Ornithorhynchus anatinus'') |journal=Mammalia |date=26 March 2021 |volume=85 |issue=2 |pages=179–181 |doi=10.1515/mammalia-2020-0027 }}</ref>
 <gallery mode="packed" heights="180px">
 Platypus cape unknown tasmania.jpg|Platypus fur cape made in 1890, now in the National Gallery of Victoria
 "Nam Sang Woo Safety Matches" platypus matchbox label art - from, Collectie NMvWereldculturen, TM-6477-76, Etiketten van luciferdoosjes, 1900-1949 (cropped).jpg|Early 20th-century [[matchbox]] label
-Australianstamp 1551.jpg|9d postage stamp from 1937
+Australianstamp 1551.jpg|9[[Australian pound|d]] postage stamp from 1937
 </gallery>
@@ Line 242: / Line 182: @@
 == Citations ==
 {{Reflist}}
-== References ==
-=== Books ===
-* {{cite encyclopedia |last=Augee |first=Michael L. |encyclopedia=World Book Encyclopedia |title=Platypus |year=2001 }}
-* {{cite book |author=Burrell, Harry |title=The Platypus |publisher=Rigby |location=Adelaide SA |year=1974 |isbn=978-0-85179-521-8 }}
-* {{cite book |author=Fleay, David H. |title=Paradoxical Platypus: Hobnobbing with Duckbills |publisher=Jacaranda Press |year=1980 |isbn=978-0-7016-1364-8 |url=https://books.google.com/books?id=IaIKAQAAMAAJ}}
-* {{cite book |author=Grant, Tom |title=The platypus: a unique mammal |publisher=University of New South Wales Press |location=Sydney |year=1995 |isbn=978-0-86840-143-0 |url=https://books.google.com/books?id=FASJWgDhxIsC}}
-* {{cite book |author=Griffiths, Mervyn |title=The Biology of the Monotremes |publisher=Academic Press |year=1978 |isbn=978-0-12-303850-0 |url=https://books.google.com/books?id=1KTwAAAAMAAJ}}
-* {{cite encyclopedia |editor1=Hutch, Michael |editor2=McDade, Melissa C. |encyclopedia=Grzimek's Animal Life Encyclopedia |title= Grzimek's Animal Life Encyclopedia: Lower metazoans and lesser deuterosomes|year=2004 |publisher=Gale |volume=12: ''Mammals III'' |oclc=1089554968 |isbn=9780787657772}}
-* {{cite book |author=Moyal, Ann Mozley |author-link=Ann Moyal |title=Platypus: The Extraordinary Story of How a Curious Creature Baffled the World |publisher=The Johns Hopkins University Press |location=Baltimore |year=2004 |isbn=978-0-8018-8052-0 |url=https://books.google.com/books?id=5DkezNMhSTYC}}
-* {{cite book|first1=Ronald |last1=Strahan|first2=Steve |last2=Van Dyck|title=Mammals of Australia |edition=3rd |url=https://books.google.com/books?id=N3tFAQAAIAAJ |date=April 2006|publisher=New Holland|isbn=978-1-877069-25-3}}
-=== Documentaries ===
-* {{Cite episode|title=Southern Exposure |url=http://www.abc.net.au/storm/exposure/default.htm |series=Eye of the Storm |network=[[Australian Broadcasting Corporation]] |year=2000 |url-status=dead |archive-url=https://web.archive.org/web/20130507073318/http://www.abc.net.au/storm/exposure/default.htm |archive-date=7 May 2013 }} DVD [[EAN 13|EAN]] 9398710245592
-* {{Cite episode|title=El Niño |url=http://www.abc.net.au/storm/nino/default.htm |series=Eye of the Storm  |year=2000 |url-status=dead |archive-url=https://web.archive.org/web/20130228154105/http://www.abc.net.au/storm/nino/default.htm |archive-date=28 February 2013 }}
 == External links ==

Platypus: Difference between revisions

Latest revision as of 21:57, 12 November 2025

Taxonomy and naming

Description

Senses

Venom

Distribution and habitat

Ecology and behaviour

Reproduction

Evolution

Genome

Conservation

Sanctuaries and captivity

Human interactions

Usage

Cultural references

See also

Footnotes

Citations

External links

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