Platypus

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How to read a taxoboxPlatypus[1]
Ornithorhynchidae-00.jpg
Conservation status
Status iucn2.3 LC.svg
Least concern

(IUCN) [2]

Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Monotremata
Family: Ornithorhynchidae
Genus: Ornithorhynchus
Blumenbach, 1800
Species: O. anatinus
Binomial name
Ornithorhynchus anatinus
(Shaw, 1799)
Platypus range (indicated by darker shading)[3]
Platypus range (indicated by darker shading)[3]

Platypus is the common name for a semi-aquatic, egg-laying mammal, Ornithorhynchus anatinus, endemic to eastern Australia, including Tasmania, and uniquely characterized by a snout like a duck's bill, a broad and flat beaver-like tail, and webbed feet like an otter. It is one of only five extant species of monotremes (order Monotremata), the only mammals that lay eggs instead of giving birth to live young. Also known as the duck-billed platypus and the duckbill, it is the sole living representative of its family (Ornithorhynchidae) and genus (Ornithorhynchus), though a number of related species have been found in the fossil record. The platypus is one of the few venomous mammals; the male platypus has a spur on the hind foot that delivers a venom capable of causing severe pain to humans.

The bizarre appearance of this egg-laying, venomous, duck-billed, beaver-tailed, otter-footed mammal baffled European naturalists when they first encountered descriptions and a pelt, with some considering it an elaborate hoax. Despite a limited distribution, it is one of the more well-known animals worldwide given its unique features, and it is a recognizable and iconic symbol of Australia. Its singular characteristics also make the platypus an important subject in the study of evolutionary biology. Ecologically, it has an important role in food chains, feeding on freshwater shrimp, crayfish, annelids, and insect larvae, and being consumed by snakes, birds of prey, crocodiles, and other predators.

Until the early twentieth century, the platypus was hunted for its fur, but it is now protected throughout its range. Although captive breeding programs have had only limited success and the Platypus is vulnerable to the effects of pollution, it is not under any immediate threat.

The plural usually is formed as either platypuses or platypus, with either correct, although platypi also appears on occasion.

Contents

Overview and description

A color print of platypuses from 1863

The platypus and four species of echidnas ("spiny anteaters") comprise the only extant species of monotremes, the egg-laying mammals of the order Monotremata, the only order in subclass Monotremata (or Prototheria). Monotremes are one of three subdivisions of mammals, the other two being the placentals (Placentalia or Eutheria) and the marsupials (Marsupialia or Metatheria), both of which give birth to live young. The key physiological difference between monotremes and other mammals is that in monotremes the urinary, defecatory, and reproductive systems all open into a single duct, the cloaca. Other mammal females have separate openings for reproduction, urination, and defecation. Monotremes are placed in two families, with the platypus belonging to Ornithorhynchidae and the four species of echidnas in the Tachyglossidae family. Echidnas also are found in Australia, as well as New Guinea.

The squat body and the broad, flat tail of the platypus are covered with dense brown fur that traps a layer of insulating air to keep the animal warm.[3][4] The platypus uses its tail for storage of fat reserves (an adaptation also found in animals such as the Tasmanian devil[5] and fat-tailed sheep). It has webbed feet and a large, rubbery snout; these are features that appear closer to those of a duck than to those of any known mammal. Unlike a bird's beak (in which the upper and lower parts separate to reveal the mouth), the snout of the platypus is a sensory organ with the mouth on the underside. The nostrils are located on the dorsal surface of the snout, while the eyes and ears are located in a groove set just back from it; this groove is closed when swimming.[4] The webbing is more significant on the front feet and is folded back when walking on land.[4]

Weight varies considerably from 0.7 to 2.4 kilograms (1.5 to 5.3 pounds), with males being larger than females: males average 50 centimeters (20 inches) in total length while females average 43 centimeters (17 inches)[4] There is substantial variation in average size from one region to another, and this pattern does not seem to follow any particular climatic rule and may be due to other environmental factors, such as predation and human encroachment.[6]

The platypus has an average body temperature of about of about 32°C (90°F), rather than the 37°C (99°F) typical of placental mammals.[7] Research suggests this has been a gradual adaptation to harsh environmental conditions on the part of the small number of surviving monotreme species rather than a historical characteristic of monotremes.[8][9]

The modern platypus young have three-cusped molars which they lose before or just after leaving the breeding burrow;[10][11] adults have heavily keratinized pads in their place.[4] The platypus jaw is constructed differently from that of other mammals, and the jaw-opening muscle is different.[4] 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 cynodonts and other pre-mammalian synapsids. However, the external opening of the ear still lies at the base of the jaw.[4] The platypus has extra bones in the shoulder girdle, including an interclavicle, which is not found in other mammals.[4] It has a reptilian gait, with legs that are on the sides of the body, rather than underneath.[4]

Venom

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

The male platypus has ankle spurs that produce a cocktail of venom,[12][13][14] composed largely of defensin-like proteins (DLPs); the venom is unique to the platypus.[15] Although powerful enough to kill smaller animals,[15] the venom is not lethal to humans, but is so excruciating that the victim may be incapacitated. Oedema rapidly develops around the wound and gradually spreads throughout the affected limb. Information obtained from case histories and anecdotal evidence indicates that the pain develops into a long-lasting hyperalgesia that persists for days or even months.[16][17] Venom is produced in the crural glands of the male, which are kidney-shaped alveolar glands 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.[4]

The venom appears to have a different function from those produced by non-mammalian species: Its effects are not life-threatening but nevertheless powerful enough to seriously impair the victim. Since only males produce venom and production rises during the breeding season, it is theorized that it is used as an offensive weapon to assert dominance during this period.[15]

Electrolocation

Monotremes are the only mammals known to have a sense of electroreception: They locate their prey in part by detecting electric fields generated by muscular contractions. The platypus' electroreception is the most sensitive of any monotreme.[18]

The electroreceptors are located in rostro-caudal rows in the skin of the bill, while mechanoreceptors (which detect touch) are uniformly distributed across the bill. The electrosensory area of the cerebral cortex is contained within the tactile somatosensory area, and some cortical cells receive input from both electroreceptors and mechanoreceptors, suggesting a close association between the tactile and electric senses. Both electroreceptors and mechanoreceptors in the bill dominate the somatotopic map of the platypus brain, in the same way human hands dominate the Penfield homunculus map.[19][20]

The platypus can determine the direction of an electric source, perhaps by comparing differences in signal strength across the sheet of electroreceptors. This would explain the animal's characteristic side-to-side motion of its head while hunting. The cortical convergence of electrosensory and tactile inputs suggests a mechanism for determining the distance of prey items which, when they move, emit both electrical signals and mechanical pressure pulses, which would also allow for computation of distance from the difference in time of arrival of the two signals.[18]

The platypus feeds by digging in the bottom of streams with its bill. The electroreceptors could be used to distinguish animate and inanimate objects in this situation (in which the mechanoreceptors would be continuously stimulated).[18] When disturbed, its prey would generate tiny electrical currents in their muscular contractions, which the sensitive electroreceptors of the platypus could detect. Experiments have shown that the platypus will even react to an "artificial shrimp" if a small electrical current is passed through it.[21]

Ecology and behavior

The platypus is very difficult to spot even on the surface of a river.

The platypus is semi-aquatic, inhabiting small streams and rivers over an extensive range from the cold highlands of Tasmania and the Australian Alps to the tropical rainforests of coastal Queensland as far north as the base of the Cape York Peninsula.[22] Inland, its distribution is not well known: it is extinct in South Australia (barring an introduced population on Kangaroo Island) and is no longer found in the main part of the Murray-Darling Basin, possibly due to the declining water quality brought about by extensive land clearing and irrigation schemes.[23] Along the coastal river systems, its distribution is unpredictable; it appears to be absent from some relatively healthy rivers, and yet maintains a presence in others that are quite degraded (the lower Maribyrnong, for example).[24]

Its habitat bridges rivers and the riparian zone for both a food supply of prey species and banks where it can dig resting and nesting burrows.[25] It may have a range of up to 7 kilometers (4.3 miles), with male's home ranges overlapping with those of 3 or 4 females.[26]

The platypus is generally regarded as nocturnal and crepuscular, but individuals are also active during the day, particularly when the sky is overcast.[25]

The platypus is an excellent swimmer and spends much of its time in the water foraging for food. When swimming, it can be distinguished from other Australian mammals by the absence of visible ears.[27] Uniquely among mammals, it propels itself when swimming by alternate rowing motion with the front two feet; although all four feet of the Platypus are webbed, the hind feet (which are held against the body) do not assist in propulsion, but are used for steering in combination with the tail.[28] Dives normally last around 30 seconds, but can last longer although few exceed the estimated aerobic limit of 40 seconds. Ten to twenty seconds are commonly spent in recovery at the surface.[29][30] The species is endothermic, maintaining its body temperature about 32°C (90°F), lower than most mammals, even while foraging for hours in water below 5°C (41°F).[4]

The platypus is a carnivore. It feeds on annelid worms and insect larvae, freshwater shrimps, and yabbies (freshwater crayfish) that it digs out of the riverbed with its snout or catches while swimming. It utilizes cheek-pouches to carry prey to the surface where they are eaten.[27] The platypus needs to eat about twenty percent of its own weight each day. This requires the platypus to spend an average of 12 hours each day looking for food.[29] When not in the water, the platypus retires to a short, straight resting burrow of oval cross-section, nearly always in the riverbank not far above water level, and often hidden under a protective tangle of roots.[27]

Natural predators include snakes, water rats, goannas, hawks, owls, and eagles. Low platypus numbers in northern Australia are possibly due to predation by crocodiles.[31] The introduction of red foxes as a predator for rabbits may have had some impact on its numbers on the mainland.[6]

Platypuses have been heard to emit a low growl when disturbed and a range of other vocalizations have been reported in captive specimens.[3]

Reproduction and life cycle

When the platypus was first discovered, scientists were divided over whether the female laid eggs. This was not confirmed until 1884 when W. H. Caldwell was sent to Australia where, after extensive searching assisted by a team of 150 Aborigines, he managed to discover a few eggs.[4][15] Mindful of the high cost of wiring England based on the cost per word, Caldwell famously, but tersely, wired London, "Monotremes oviparous, ovum meroblastic." That is, monotremes lay eggs, and the eggs are similar to those of reptiles in that only part of the egg divides as it develops.

The species exhibits a single breeding season; mating occurs between June and October, with some local variation taking place in populations across the extent of its range.[31] Historical observation, mark and recapture studies, and preliminary investigations of population genetics indicate the possibility of resident and transient members of populations and suggest a polygynous mating system.[32] Females are thought likely to become sexually mature in their second year, with breeding confirmed to still take place in animals over nine years old.[32]

Outside the mating season, the platypus lives in a simple ground burrow whose entrance is about 30 centimeters (12 inches) above the water level. After mating, the female constructs a deeper, more elaborate burrow up to 20 meters (66 feet) long and blocked with plugs at intervals (which may act as a safeguard against rising waters or predators, or as a method of regulating humidity and temperature).[33]

The male takes no part in caring for its young, and retreats to its yearlong burrow. The female softens the ground in the burrow with dead, folded, wet leaves and she fills the nest at the end of the tunnel with fallen leaves and reeds for bedding material. This material is dragged to the nest by tucking it underneath her curled tail.[3]

The female platypus has a pair of ovaries but only the left one is functional. It lays one to three (usually two) small, leathery eggs (similar to those of reptiles), that are about 11 millimeters (0.43 inches) in diameter and slightly rounder than bird eggs.[34] The eggs develop in utero for about 28 days with only about 10 days of external incubation (in contrast to a chicken egg, which spends about 1 day in tract and 21 days externally). After laying her eggs, the female curls around them. The incubation period is separated into three parts. In the first, the embryo has no functional organs and relies on the yolk sac for sustenance. The yolk is absorbed by the developing young.[35] During the second, the digits develop, and in the last, the egg tooth appears.[36]

The newly hatched young are vulnerable, blind, and hairless, and are fed by the mother's milk. Although possessing mammary glands, the platypus lacks teats. Instead, milk is released through pores in the skin. There are grooves on the mother's abdomen that form pools of milk, allowing the young to lap it up.[3][31] After they hatch, the offspring are suckled for three to four months. During incubation and weaning, the mother initially only leaves the burrow for short periods to forage. When doing so, she creates a number of thin soil plugs along the length of burrow, possibly to protect the young from predators; pushing past these on her return forces water from her fur and allows the burrow to remain dry.[37] 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.[31]

In captivity, platypuses have survived to 17 years of age and tagged wild specimens have been recaptured at 11 years old. Mortality rates for adults in the wild appear to be low.[4]

History, taxonomy, and etymology

When the platypus was first discovered by Europeans in 1798, a pelt and sketch were sent back to the United Kingdom by Captain John Hunter, the second Governor of New South Wales.[38] The British scientists were at first convinced that the attributes must have been a hoax.[3] George Shaw, who in 1799 produced the first description of the animal in the Naturalist's Miscellany, stated that it was impossible not to entertain doubts as to its genuine nature, and Robert Knox believed it may have been produced by some Asian taxidermist.[39] It was thought that 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.[3]

The common name, platypus, is Latin derived from the Greek words πλατύς or platys, meaning "flat" or "broad," and πους or pous, meaning "foot,"—in other words, ""flat foot."[40] Shaw assigned it as a Linnaean genus name when he initially described it, but the term was quickly discovered to already belong to the wood-boring ambrosia beetle (genus Platypus).[4] It was independently described as Ornithorhynchus paradoxus by Johann Blumenbach in 1800 (from a specimen given to him by Sir Joseph Banks)[41] and following the rules of priority of nomenclature, it was later officially recognize as Ornithorhynchus anatinus.[4] The scientific name Ornithorhynchus is derived from ορνιθόρυνχος ("ornithorhynkhos"), which literally means "bird snout" in Greek, and anatinus, which means "duck-like" in Latin.

There is no universally agreed upon plural of "platypus" in the English language. Scientists generally use "platypuses" or simply "platypus." Colloquially, "platypi" is also used for the plural, although this is pseudo-Latin;[3] the Greek plural would be "platypodes." Early British settlers called it by many names, such as watermole, duckbill, and duckmole.[3] The name "platypus" is often prefixed with the adjective "duck-billed" to form duck-billed platypus, despite there being only one species of platypus.[42]

Evolution

Platypus skeleton

The platypus and other monotremes were very poorly understood and some of the nineteenth century myths that grew up around them—for example, that the monotremes were "inferior" or quasi-reptilian—still endure.[43] In fact, modern monotremes are the survivors of an early branching of the mammal tree; a later branching is thought to have led to the marsupial and placental groups.[44][43] Although in 1947, William King Gregory had theorized 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.[43][45]

The oldest discovered fossil of the modern platypus dates back to about 100,000 years ago, during the Quaternary period. The extinct monotremes (Teinolophos and Steropodon) were closely related to the modern platypus.[45] The fossilized 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, 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.[10] The fossil is thought to be about 110 million years old, which means that the platypus-like animal was alive during the Cretaceous period, making it the oldest mammal fossil found in Australia. Monotrematum sudamericanum, another fossil relative of the Platypus, has been found in Argentina, indicating that monotremes were present in the supercontinent of Gondwana when the continents of South America and Australia were joined via Antarctica (up to about 167 million years ago).[10][46]

Because of the early divergence from the therian mammals and the low numbers of extant monotreme species, it is a frequent subject of research in evolutionary biology. In 2004, researchers at the Australian National University discovered the platypus has ten sex chromosomes, compared with two (XY) in most other mammals (for instance, a male platypus is always XYXYXYXYXY).[47] Although given the XY designation of mammals, the sex chromosomes of the platypus is more similar to the ZZ/ZW sex chromosomes found in birds.[48] It also lacks the mammalian sex-determining gene SRY, meaning that the process of sex determination in the platypus remains unknown.[49]

A draft version of the platypus genome sequence was published in Nature on May 8, 2008, revealing both reptilian and mammalian elements, as well as two genes found previously only in birds, amphibians, and fish.[48] More than 80 percent of the platypus' genes are common to the other mammals whose genomes have been sequenced.

Conservation status

A depiction of a platypus from a book for children published in Germany in 1798

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.

The current and historical abundance of the platypus, however, is less well-known and it has probably declined in numbers, although still being considered as "common" over most of its current range.[25] The species was extensively hunted for its fur until the early years of the twentieth century and, although protected throughout Australia in 1905,[37] up until about 1950 it was still at risk of drowning in the nets of inland fisheries.[23] The platypus does not appear to be in immediate danger of extinction thanks to conservation measures, but it could be impacted by habitat disruption caused by dams, irrigation, pollution, netting, and trapping.[2] The IUCN lists the Platypus on its Red List as Least Concern.[2]

Platypuses generally suffer from few diseases in the wild; however, there is widespread public concern in Tasmania about the potential impacts of a disease caused by the fungus Mucor amphibiorum. The disease (termed Mucormycosis) only affects Tasmanian platypuses, and has not been observed in platypuses in mainland Australia. Affected platypuses can develop ugly skin lesions or ulcers on various parts of the body, 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 efficiency. The Biodiversity Conservation Branch at the Department of Primary Industries and Water are collaborating with NRM north and University of Tasmania researchers to determine the impacts of the disease on Tasmanian platypus, as well as the mechanism of transmission and current spread of the disease.[50]

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. This is a difficult task, and only a few young have been successfully raised since—notably at Healesville Sanctuary in Victoria. The leading figure in these efforts was David Fleay, who established a platypussary—a simulated stream in a tank—at the Healesville Sanctuary and had a successful breeding first in 1943.

Notes

  1. C. Groves, "Order Primates," "Order Monotremata," (and select other orders). Page(s) 2 in D. E. Wilson and D. M. Reeder, eds., Mammal Species of the World, 3rd edition, Johns Hopkins University Press (2005). ISBN 0801882214.
  2. 2.0 2.1 2.2 Australasian Marsupial and Monotreme Specialist Group, "Ornithorhynchus anatinus," 2007 IUCN Red List of Threatened Species (1996). Retrieved on September 25, 2008. Database entry includes a brief justification of why this species is of least concern
  3. 3.0 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 Australian Platypus Conservancy, "Platypus facts file," Australian Platypus Conservancy. Retrieved September 25, 2008.
  4. 4.00 4.01 4.02 4.03 4.04 4.05 4.06 4.07 4.08 4.09 4.10 4.11 4.12 4.13 4.14 J. R. Grant, "Chapter 16, Ornithorhynchidae,", in D. Walton, and B. J. Richardson, (eds.), Fauna of Australia. Volume 1B, Mammalia. (Canberra: Australian Govt. Pub. Service. 1989. ISBN 0644060565). Retrieved September 26, 2008.
  5. E. R. Guiler, "Tasmanian devil," pages 27 to 28 in R. Strahan, (ed.), The Australian Museum Complete Book of Australian Mammals. (Angus & Robertson 1983). ISBN 0207144540.
  6. 6.0 6.1 S. Munks, and S. Nicol, "Current research on the platypus, Ornithorhynchus anatinus in Tasmania: Abstracts from the 1999 'Tasmanian Platypus WORKSHOP'," University of Tasmania (1999). Retrieved September 25, 2008.
  7. Department of Biology, Davidson College, ["http://www.bio.davidson.edu/courses/anphys/1999/White/thermal.htm "Thermal biology of the platypus,"] Davidson College (1999). Retrieved September 25, 2008.
  8. J. M. Watson, and J. A. M. Graves, "Monotreme cell-cycles and the evolution of homeothermy," Australian Journal of Zoology 36(1988)(issue 5): 573–584. Retrieved September 25, 2008.
  9. T. J. Dawson, T. R. Grant, and D. Fanning, "Standard metabolism of monotremes and the evolution of homeothermy," Australian Journal of Zoology 27 (4)(1979): 511–515. Retrieved September 25, 2008.
  10. 10.0 10.1 10.2 R. Pascual, F. J. Goin, L. Balarino, and D. E. Udrizar Sauthier, "New data on the Paleocene monotreme Monotrematum sudamericanum, and the convergent evolution of triangulate molars," Acta Palaeontologica Polonica 47(3) (2002): 487–492.
  11. H. Rance, "Living mammals are placentals (eutheria), marsupials, and monotremes," pages 304-306 in The Age of Mammals. Retrieved September 25, 2008.
  12. Australian Fauna, "Platypus (Ornithorhynchus anatinus)," Australianfauna.com (2006). Retrieved September 25, 2008.
  13. University of Sydney, "Platypus venom linked to pain relief," University of Sydney (2008). Retrieved September 25, 2008.
  14. Rainforest Australia, "Platypus poison,". Rainforest Australia. Retrieved September 25, 2008.
  15. 15.0 15.1 15.2 15.3 V. B. Gerritsen, "Platypus poison," Protein Spotlight 29(2002). Retrieved September 25, 2008.
  16. G. M. de Plater, P. J. Milburn, and R. L. Martin, "Venom from the platypus, Ornithorhynchus anatinus, induces a calcium-dependent current in cultured dorsal root ganglion cells," Journal of Neurophysiology 85(2001)(issue 3): 1340–1345. Retrieved September 25, 2008.
  17. B. G. Frey, "The venom of the platypus (Ornithorhynchus anatinus), Kingsnake.com. Retrieved September 25, 2008.
  18. 18.0 18.1 18.2 J. D. Pettigrew, "Electroreception in monotremes," The Journal of Experimental Biology 202(1999): 1447–1454. Retrieved September 25, 2008.
  19. J. D. Pettigrew, P. R. Manger, and S. L. Fine, "The sensory world of the platypus," Philosophical Transactions of the Royal Society of London 353(1998): 1199–1210. Retrieved September 25, 2008.
  20. R. Dawkins, The Ancestor's Tale, A Pilgrimage to the Dawn of Life (Boston: Houghton Mifflin Company, 2004). ISBN 0618005838.
  21. A. Manning, and M. S. Dawkins, An Introduction to Animal Behavior, 5th edition. (Cambridge University Press, 1998). ISBN 0521570247
  22. Department of Primary Industries and Water (DPIW), Tasmania, "Platypus," Department of Primary Industries and Water, Tasmania. Retrieved September 25, 2008.
  23. 23.0 23.1 A. Scott and T. Grant, "Impacts of water management in the Murray-Darling Basin on the platypus (Ornithorhynchus anatinus) and the water rat (Hydromus chrysogaster)," CSIRO Australia Technical Report 23/97 (1997). Retrieved September 25, 2008.
  24. Australian Platypus Conservancy, "Platypus in country areas," Australian Platypus Conservancy. Retrieved September 25, 2008.
  25. 25.0 25.1 25.2 T. G. Grant, and P. D. Temple-Smith, "Field biology of the platypus (Ornithorhynchus Anatinus): Historical and current perspectives," Philosophical Transactions: Biological Sciences 353(1998)(issue 1372): 1081–1091. Retrieved September 26, 2008.
  26. J. L. Gardner and M. Serena, "Spatial-organization and movement patterns of adult male platypus, Ornithorhynchus anatinus (Monotremata, Ornithorhynchidae)," Australian Journal of Zoology 43(1995)(issue 1): 91–103. Retrieved September 26, 2008.
  27. 27.0 27.1 27.2 Parks and Wildlife Service, Tasmania, "Platypus, Ornithorhynchus anatinus," Parks and Wildlife Service, Tasmania. Retrieved September 26, 2008.
  28. F. E. Fish, R. V. Baudinette, P. B. Frappell, and M. P. Sarre, "Energetics of swimming by the platypus Ornithorhynchus Anatinus: Metabolic effort associated with rowing," The Journal of Experimental Biology 200(1997)(issue 20): 2647–2652. Retrieved September 26, 2008.
  29. 29.0 29.1 P. Bethge, "Energetics and foraging behavior of the platypus," University of Tasmania (2002). Retrieved September 26, 2008.
  30. H. Kruuk, "The diving behaviour of the platypus (Ornithorhynchus anatinus) in waters with different trophic status," The Journal of Applied Ecology 30(1993)(issue 4): 592–598. Retrieved September 26, 2008.
  31. 31.0 31.1 31.2 31.3 Environmental Protection Agency, "Platypus (Ornithorhynchus anatinus)," Environmental Protection Agency/Queensland Parks and Wildlife Service (2008). Retrieved September 25, 2008.
  32. 32.0 32.1 T. R. Grant, M. Griffiths, and R.M.C. Leckie, "Aspects of lactation in the platypus, Ornithorhynchus anatinus (Monotremata), in waters of Eastern New South Wales," Australian Journal of Zoology 31(1983)(issue 6): 881–889. Retrieved September 26, 2008.
  33. A. B. Sorin and P. Myers, "Family Ornithorhynchidae (platypus)," Animal Diversity Web (2001). Retrieved September 26, 2008.
  34. R. L. Hughes, and L. S. Hall, "Early development and embryology of the platypus," Philosophical Transactions of the Royal Society B: Biological Sciences 353(1998)(issue 1372): 1101–1114. Retrieved September 26, 2008.
  35. A. Moyal, "The puzzling platypus," Ockhams Razor July 22, 2001. Radio National (Australia). Retrieved September 26, 2008.
  36. P. R. Manger, L. S. Hall, and J. D. Pettigrew, "The development of the external features of the platypus (Ornithorhynchus anatinus)," Philosophical Transactions: Biological Sciences 353(1998)(issue 1372): 1115–1125. Retrieved September 26, 2008.
  37. 37.0 37.1 Queensland Museum, "Egg-laying mammals," Queensland Museum. Retrieved September 26, 2008.
  38. B. K. Hall, "The paradoxical platypus," BioScience 49 (3) (1999): 211–218. Retrieved September 26, 2008.
  39. A. Boese, "Duck-billed platypus," Museum of Hoaxes. Retrieved September 26, 2008.
  40. H. G. Liddell, and R. Scott. 1987. A Lexicon Abridged From Liddell and Scott's Greek-English Lexicon. (Oxford: Clarendon Press. ISBN 0199102074)
  41. W. Horky, "Platypus paradoxes," National Library of Australia 52 (2001). Retrieved September 26, 2008.
  42. P. Bethge, "Platypus," Department of Anatomy & Physiology, University of Tasmania (1997). Retrieved September 26, 2008.
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References

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