Scientific Name
Raphus cucullatus
Extinct Year
about 1662
Conservation Status
Extinct (EX)

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Wikipedia Article

The Dodo (Raphus cucullatus) is an extinct, flightless bird that was endemic to the Mascarene island of Mauritius (located east of Madagascar in the Indian Ocean). Genetically nested within pigeons and doves, its closest genetic relative was the (likewise extinct) Rodrigues Solitaire, the two forming the subfamily Raphinae. The closest living relative of the Dodo is the Nicobar Pigeon. While a white Dodo was at one time believed to have existed on the nearby island of Runion, this is now known to be incorrect. The Dodo was about one meter (about 3.3 ft) tall, and may have weighed approximately in the wild. Its external appearance is now evidenced only by paintings and written accounts from the 17th century. Because these vary considerably, and because only a few sketches are known to have been drawn from specimens in the wild, its true physical appearance remains a mystery. Similarly, little-to-nothing about its habitat and behavior is known with certainty. It has been depicted with brownish-gray plumage, yellow feet, a tuft of tail feathers, a gray-colored, naked head, and a black, yellow, and green beak. It used gizzard stones to help digest its food, which is thought to have included fruits, and its main habitat is believed to have been the woods in the drier coastal areas of Mauritius. It is presumed that the Dodo became flightless because of the ready availability of abundant food sources and a relative absence of predators on Mauritius. The first recorded mention of the Dodo was made by Dutch sailors in 1598. After that time, the bird was preyed upon by hungry sailors, their domesticated animals, and other invasive species introduced during that time. The last credible recorded sighting of a Dodo dates from 1662. Its extinction was not immediately noticed, and some considered it to be a mythological creature. In the 19th century, research was conducted on a small amount of remains of four specimens that had been brought to Europe in the early 17th century. Since then, a large amount of subfossil material has been collected from Mauritius, mostly from the Mare aux Songes swamp. The extinction of the Dodo within only about a century of its discovery called attention to the problem of human involvement in the complete disappearance of entire species, which until that time was a previously unrecognised issue. The Dodo achieved widespread recognition from its notable role in the story of Alice in Wonderland, and it has since become a fixture in popular culture, often as a symbol of extinction and obsolescence. It is frequently used as a mascot on the island of Mauritus.
Many different explanations have historically been offered to determine the affinities of the Dodo, including that it was a small ostrich, a rail, an albatross, or a vulture. In 1842, Johannes Theodor Reinhardt proposed Dodos were actually ground pigeons, based on studies of a Dodo skull he had discovered in the royal Danish collection at Copenhagen, Denmark. This view was met with ridicule, but later supported by Hugh Edwin Strickland and Alexander Gordon Melville in their seminal 1848 monograph, The Dodo and Its Kindred, which attempted to separate Dodo myth from reality. After dissecting the preserved head and foot of the specimen at Oxford Museum, they also found that the Dodo was closely related to the likewise extinct Rodrigues Solitaire (Pezophaps solitaria). The Dodo was anatomically similar to pigeons in many features. Strickland pointed to the keratinous portion of the beak being very short, with the basal part being long, slender, and naked. Other pigeons also have naked skin around their eyes, almost reaching their beak, as in Dodos. The forehead was high in relation to the beak, and the nostril was located low on the middle of the beak and was surrounded by skin, a combination of features only shared with pigeons. The legs of the Dodo were generally more similar to terrestrial pigeons than other birds, both in the scales and in their skeletal features. Depictions of the large crop hinted at a relation, as this feature is more developed in pigeons than other birds. Pigeons generally have very small clutch sizes, which agrees with the single egg said to have been laid by the Dodo. Also like pigeons, the Dodo lacked the vomer and septum of the nostrils, and it shared details in the lower jaw, the zygomatic bone, the palate, and the hallux. The Dodo differed from other pigeons mainly in the small size of the wings, and the large size of the beak in proportion to the rest of the cranium. For many years the Dodo and the Rodrigues Solitaire were placed in a family of their own, the Raphidae (formerly Dididae). This was because their exact relationships to other pigeons were unresolved. They were also placed in a monotypic family each (Raphidae and Pezophapidae, respectively) due to the suggestion that they had evolution evolved their similar features independently. Recently, the family Raphidae was dissolved and the Dodo and Solitaire were placed in their own subfamily, Raphinae, nested within the Columbidae. Comparison of mitochondrial cytochrome b and 12S rRNA sequences isolated from the tarsal of a Dodo and a femur of a Rodrigues Solitaire confirmed their close relationship, and their placement within the Columbidae. The genetic evidence was interpreted to show that the Southeast Asian Nicobar Pigeon is their closest living relative, followed by the Crowned Pigeons of New Guinea, and the superficially Dodo-like Tooth-billed Pigeon from Samoa. The generic name of the latter is Didunculus ("little Dodo")this bird was also referred to as "Dodlet" by Richard Owen. The following cladogram, from Shapiro and colleagues (2002), shows the Dodo's closest relationships within Columbidae, a clade consisting of generally ground-dwelling island endemics. {{Clade|style=font-size:90% |1={{Clade |1={{Clade |1=Goura victoria (Victoria Crowned Pigeon) |2={{Clade |1={{Clade |1= }} }} }} |2=Didunculus strigirostris (Tooth-billed Pigeon) }} }} A similar cladogram was published in 2007, differing only in the inverted placement of Goura and Dicunculus, as well as in the inclusion of the Pheasant Pigeon and the Thick-billed Ground Pigeon at the base of the clade. The 2002 study indicated that the ancestors of the Solitaire and the Dodo diverged around the Paleogene-Neogene boundary. The Mascarene Islands, a group of islands in the Indian Ocean east of Madagascar that include Mauritius and Runion, are of volcanic origin and are less than 10 million years old. Therefore, the ancestors of both birds likely remained capable of flight for some considerable time after the separation of their lineage. The lack of mammalian herbivores competing for resources on these islands allowed the Solitaire and the Dodo to attain very large sizes. Throughout the 19th century, several species were classified as being congeneric with the Dodo, including the Rodrigues Solitaire and the Runion Solitaire, as Didus solitarius and Raphus solitarius, respectively. An atypical 17th century description of a Dodo and bones found on Rodrigues, now known to have belonged to the Rodrigues Solitaire, led Abraham Dee Bartlett to name a new species, Didus nazarenus, in 1852. Being based on Solitaire remains, it is now a synonym of that species. Crude drawings of the Red Rail of Mauritius were also misinterpreted as several species of Dodos, Didus broeckii and Didus herberti. Another large, flightless pigeon, the Viti Levu Giant Pigeon, was described in 2001 from subfossil material from Fiji. It was only slightly smaller than the Dodo and the Solitaire, and it is also thought to have been related to the Crowned Pigeons.
One of the original names for the Dodo was "walghvogel" (Dutch), first used in the journal of Vice Admiral Wybrand van Warwijck, who visited the island during the Second Dutch Expedition to Indonesia in 1598. His description, translated into English in 1599, explained the meaning of the name as follows: Another account from the same voyage, perhaps the first mentioning the bird, states the Portuguese referred to them as "penguins". The meaning may not have been derived from penguin, but from "pinion", a reference to the small wings. The crew of the Dutch ship Gelderland referred to the bird as "Dronte" (meaning "swollen") in 1602, a name that is still used in some languages. This crew also called them "griff-eendt" and "kermisgans", in reference to fowl fattened for the Kermesse festival in Amsterdam, which was held the day after they anchored on Mauritius. The etymology of the word Dodo is unclear. Some ascribe it to the Dutch word dodoor for "sluggard", but it is more likely related to Dodaars, which means either "fat-arse" or "knot-arse", referring to the knot of feathers on the hind end. The first record of the word Dodaars is in captain Willem Van West-Zanen's journal in 1602. The English form "Dodar" was used until the 18th century. Sir Thomas Herbert was the first to use the word Dodo in print in his 1634 travelogue, and claimed it was referred to as such by the Portuguese, who had visited the island in 1507. Another Englishman, Emmanuel Altham, had used the word in a 1628 letter, wherein he also claimed the origin was Portuguese. Insofar as is known, the Portuguese did not use the word. Nevertheless, some sources still state that the word "Dodo" derives from the Portuguese word doudo (currently doido), meaning "fool" or "crazy". David Quammen suggested that Dodo was an onomatopoeic approximation of the bird's own call, a two-note pigeony sound resembling "doo-doo". The Latin name cucullatus ("hooded") was first used by Juan Eusebio Nieremberg in 1635 as Cygnus cucullatus, in reference to Carolus Clusius' 1605 depiction of a Dodo. In his 18th century classic work Systema Naturae, Carl Linnaeus used cucullatus as the specific name, but combined it with the genus name Struthio (that of the ostrich). Mathurin Jacques Brisson coined the genus name Raphus (referring to bustards) in 1760, which resulted in the current name Raphus cucullatus. In 1766, Linnaeus coined the new binomial Didus ineptus (meaning "inept Dodo"). This has become a synonym of the earlier name because of nomenclatural priority.
No complete Dodo specimens exist, making its external appearance, such as plumage and colouration, hard to determine. Subfossil remains and remnants of the birds that were brought to Europe in the 17th century show that they were very large birds, one metre (3.3 ft) tall, and possibly weighing up to 23 kg (50 pounds). The higher weights have been attributed to birds in captivity, and estimated to have been in the range of in the wild. A later estimate gives an average weight as low as . This has been questioned, and there is still some controversy over the actual weight. It has been suggested that its weight was seasonal, however, and that individuals were fat during cool seasons, but slim during hot seasons. The bird was sexually dimorphic, with males being the largest, and having proportionally longer beaks. The beak was up to 23-centimetres (9-inch) long and had a hooked point. A study of the few remaining feathers on the Oxford specimen head showed that they were pennaceous rather than plumaceous (downy), and most similar to those of other pigeons. The Dodo shared several traits with the Rodrigues Solitaire, its closest relative, such as features in the skull, pelvis, and sternum, as well as their large size. It differed in other aspects, such as being more robust and shorter than the Solitaire, having a larger skull and beak, a rounded skull roof, and smaller orbits. The Dodo's neck and legs were proportionally shorter, and it did not possess an equivalent to the knob present on the wrists of the Solitaire. Many of the skeletal features of the Dodo and the Solitaire that are unique among pigeons have been attributed to their flightlessness. The pelvic elements were thicker than those of flighted pigeons to support the higher weight, and the pectoral region and the small wings were paedomorphic, meaning they were underdeveloped and retained juvenile features. However, the skull, trunk and pelvic limbs were peramorphic, meaning that they changed considerably with age. Illustrations and written accounts of encounters with Dodos made between the Dodo's discovery and its extinction (15981662) are the primary evidence for its external appearance. According to most representations, the Dodo had greyish or brownish plumage, with lighter primary feathers, and a tuft of curly light feathers high on its rear end. The head was grey and naked, the beak green, black and yellow, and the legs were stout and yellowish, with black claws.
Most contemporary descriptions of the Dodo are found in journals of Dutch East India Company ships that docked in Mauritius when the island was ruled by the Dutch Empire. These records were used as guides for future voyages. However, few contemporary accounts are reliable, as many seem based on earlier accounts, and none were written by scientists. One of the earliest accounts, from van Warwijck's 1598 journal, describes the bird thus: One of the most detailed descriptions is by Sir Thomas Herbert in A Relation of Some Yeares Travaille into Afrique and the Greater Asia from 1634:
About 20 17th century illustrations of the Dodo are known that could have been based on either live or stuffed specimens. All post-1638 depictions appear to be based on earlier images, which is also around the time reports mentioning Dodos became rarer. Differences in the depictions have led authors such as Anthonie Cornelis Oudemans and Masauji Hachisuka to speculate regarding sexual dimorphism, ontogenic traits, seasonal variation, and even the existence of different species, but these theories are not accepted today. Because details such as markings of the beak, the form of the tail feathers, and colouration vary from account to account, it is impossible to determine the exact morphology of these features, whether they indicate different age or sex, or if they even reflect reality. Dodo specialist Julian Hume has argued that the nostrils of the living Dodo would have actually been slits, as seen in the Gelderland, Cornelis Saftleven, Crocker Art Gallery and Ustad Mansur images. According to this claim, the gaping nostrils often seen in Dodo paintings indicate that dried out taxidermy specimens had been used as models. The travel journal of the Dutch ship Gelderland (16011603), rediscovered in the 1860s, contains the only known sketches of living or recently killed specimens drawn on Mauritius. They have been attributed to the professional artist Joris Joostensz Laerle, who also drew other now-extinct Mauritian birds, as well as a second, less refined artist. Apart from these sketches, it is unknown how many of the other illustrations were drawn from life or from stuffed specimens, affecting their reliability. The traditional image of the Dodo is of a very fat and clumsy bird, but this view may be exaggerated. The general opinion of scientists today is that many old European depictions were based on overfed captive birds, or crudely stuffed specimens. The Dutch painter Roelant Savery was the most prolific and influential illustrator of the Dodo, having made at least ten depictions, often showing it in the lower corners. A famous painting of his from 1626 now called "Edwards' Dodo" (as it was once owned by George Edwards) in the British Museum has since become the standard image of a Dodo. The image shows a particularly fat bird, and is the source for many other Dodo illustrations. An Indian Mughal painting that was rediscovered in St. Petersburg in the 1950s shows a Dodo along with native Indian birds. It depicts a slimmer, brownish bird, and is regarded as one of the most accurate depictions of the living Dodo by Professor A. Iwanow, who discovered it, and Dodo-specialist Julian Hume; the surrounding birds are clearly identifiable and depicted in their natural colours. It is believed to be from the 17th century and has been attributed to the artist Ustad Mansur. The bird depicted likely lived in the menagerie of Mughal Emperor Jahangir, located in Surat, where the English traveller Peter Mundy also claimed to have seen Dodos.
Preening Dodo.jpg|left|thumb|alt=Painting of a Dodo preening its foot|Dodo preening itself on a Savery painting from 1626 Little is known of the behaviour of the Dodo, as most contemporary descriptions are very brief. Studies of the cantilever strength of its leg bones indicate that it was able to run quite fast. Being flightless and terrestrial, due to the lack of mammalian predators and other kinds of natural enemies on Mauritius, it likely nested on the ground. The account by Franois Cauche from 1651 is the only description of the egg and call: Due to the clutch being described as consisting of a single egg, and the large size of the bird itself, it has been proposed that the Dodo was K-selected, meaning that it produced a low number of altricial offspring, which required extensive parental care until they matured. However, Cauche's account is problematic, since it also mentions the bird had three toes and no tongue, unlike Dodos, which led some to believe it referred to a new species of Dodo ("Didus nazarenus"). The description was most likely mingled with that of a cassowary, however, and Cauche's writings have other inconsistencies. A mention of a "young ostrich" taken on board a ship in 1617 is the only other reference to a possible juvenile Dodo. Some evidence indicates that the bird may have had a protracted development period, including the large size, and the fact that tropical and frugivorous birds have slower growth rates. The preferred habitat of the Dodo is unknown, but old descriptions suggest it inhabited the woods on the drier coastal areas of south and west Mauritius. The Mare aux Songes swamp, where most Dodo remains have been found, is close to the sea in south eastern Mauritius, which also supports this notion. Such a limited distribution across the island would likely had contributed to its extinction. A 1601 map from the journal of the ship Gelderland shows a small island off the coast of Mauritius where Dodos were caught. It has been suggested this island was in Tamarin Bay, on the west coast of Mauritius. However, subfossil bones have also been found inside caves in highland areas, indicating that it once occurred on mountains. Work the Maure aux Songes swamp has shown that its habitat was dominated by tambalacoque and Pandanus trees, as well as endemic palms. Many endemic species of Mauritius became extinct after the arrival of man, so the ecosystem of the island is heavily damaged, and hard to reconstruct. Before humans arrived, Mauritius was entirely covered in forests, but very little remains today due to deforestation. The surviving endemic fauna is still seriously threatened. The Dodo lived alongside other recently extinct Mauritian birds such as the flightless Red Rail, the Broad-billed Parrot, Thirioux's Grey Parrot, the Mauritius Blue Pigeon, the Mauritius Owl, the Mascarene Coot, the Mauritian Shelduck, the Mauritian Duck, and the Mauritius Night Heron. Extinct Mauritian reptiles include the Saddle-backed Mauritius giant tortoise, the Domed Mauritius giant tortoise, the Mauritian Giant Skink, and the Round Island Burrowing Boa. The Small Mauritian Flying Fox and the snail Tropidophora carinata lived on Mauritius and Runion, but vanished from both islands.
A 1631 Dutch document, rediscovered in 1887 but now lost, is the only account of its diet, and also mentions that it used the beak for defense: In addition to fallen fruits, the Dodo likely subsisted on nuts, seeds, bulbs and roots. Anthonie Oudemans suggested that as Mauritius has marked dry and wet seasons, the Dodo probably fattened itself on ripe fruits at the end of the wet season to survive the dry season, when food was scarce; contemporary reports describe the bird's "greedy" appetite. France Staub suggested they mainly fed on palm fruits, and attempted to correlate the fat-cycle of the Dodo with the fruiting regime of the palms. It has also been suggested that the Dodo might have eaten crabs and shellfish, like their relatives the Crowned Pigeons. Its feeding habits must have been versatile, since captive specimens were likely given a wide range of food on the long ship journeys. Several contemporary sources state that the Dodo used gizzard stones to aid digestion. The English writer Sir Hamon L'Estrange witnessed a live bird in London and described it as follows: It is not known how the young were fed, but related pigeons provide crop milk. Contemporary depictions show a large crop, which was probably used to add space for food storage, and for producing crop milk. It has been suggested that the maximum size attained by the Dodo and the Solitaire was limited by the amount of crop milk they were able to produce for their young during early growth. In 1973, the tambalacoque, also known as the "Dodo tree", was thought to be dying out on Mauritus, which it was endemic to. There were supposedly only 13 specimens left, all estimated to be about 300 years old. The true age could not be determined because Tambalacoque have no growth rings. Stanley Temple hypothesised that it depended on the Dodo for its propagation, and that its seeds would germinate only after passing through the bird's digestive tract; he claimed that the tambalacoque was now nearly extinct because of the disappearance of the Dodo. He force-fed seventeen tambalacoque fruits to wild turkeys and three germinated. Temple did not try to germinate any seeds from control fruits not fed to turkeys, so the significance of his findings is unclear. Temple also overlooked reports on tambalacoque seed germination from the 1940s, which found the seeds germinated, albeit very rarely, without being abraded during digestion. Temple's hypothesis has been contested. Others have suggested the decline of the tree was exaggerated, or seeds were also distributed by other extinct animals such as Cylindraspis tortoises, fruit bats or the Broad-billed Parrot. According to Wendy Strahm and Anthony Cheke, two experts in Mascarene ecology, the tree while rare has germinated since the demise of the Dodo and numbers several hundred, not 13 as claimed by Temple, hence discrediting Temple's view as to the Dodo and the tree's sole survival relationship. It has also been suggested that the Broad-Billed Parrot may have depended on Dodos and Cylindraspis tortoises to eat palm fruits and excrete their seeds, which would then be eaten by the parrots. Anodorhynchus macaws depended on the now extinct South American megafauna in the same way, but have since switched to domesticated cattle performing this function.
Though Mauritius had previously been visited by Arab vessels and Portuguese sailors, they did not settle on the island, and none of them left any known records of encounters with Dodos. However, it has been suggested that the Portuguese name for Mauritius, "Cerne (swan) Island", was a reference to the Dodos. The Dutch Empire acquired the island in 1598, which was renamed after Maurice of Nassau, and thereafter it was used for the provisioning of Dutch East India Company trade vessels during their journeys. The earliest known descriptions of the Dodo were made by Dutch travellers during the Second Dutch Expedition to Indonesia, led by Admiral Jacob van Neck in 1598. They appear in reports published in 1601, which also contain the first published illustration of the bird. Since the first sailors who visited Mauritius had been at sea for a long time, their interest in these large birds was mainly culinary. Although many later writings describe the meat as unsavoury, early journals state that it was tough but good, though not as delectable as the abundantly available pigeons. The journal of Willem Van West-Zanen of the ship 'Bruin-Vis', written in 1602 but unpublished until 1648, describes interaction with Dodos and mentions large numbers hunted for food: An illustration made for the published version of the Van West-Zanen journal showing the killing of Dodos, as well as a now locally extinct seacow, and possibly Thirioux's Grey Parrot, was captioned with the following Dutch poem, here in Errol Fuller's 2002 translation: 230 years before Darwin's theory of evolution, the appearance of the Dodo and the Red Rail led Peter Mundy to speculate:
The Dodo was found interesting enough that living specimens were sent to Europe and the East. The number of transported Dodos that reached their destinations alive is uncertain, and it is unknown how they relate to contemporary depictions and the few non-fossil remains in European museums. Hamon L'Estrange's description of a Dodo he saw in London in 1638 is the only account that specifically mentions a live specimen in Europe. Adriaen van de Venne drew a Dodo in 1626 he claimed to have seen in Amsterdam, but did not mention if it was alive, and his depiction is remiscent of Savery's "Edwards' Dodo". Two live specimens were seen by Peter Mundy in Surat, India, between 1628 and 1634, one of which was perhaps the individual painted by Ustad Mansur around 1625. One Dodo had been sent as far as Nagasaki, Japan in 1647, but it is unknown if it arrived. In 1628, Emmanuel Altham visited Mauritius and sent a letter to his brother in England: Whether the Dodo survived the journey is unknown, and the letter was destroyed by fire in the 19th century. That whole stuffed Dodos were present in Europe indicates they had been brought alive and died there; it is unlikely that taxidermists were on board the visiting ships, and spirits were not yet used to pickle biological specimens. Most tropical specimens were preserved as dried heads and feet. Based on a combination of contemporary accounts, paintings and specimens, Julian Hume has inferred that at least eleven transported Dodos reached their destinations alive.
Like many animals that evolved in isolation from significant predators, the Dodo was entirely fearless of humans. This fearlessness along with its inability to fly made the Dodo easy prey for visiting sailors. Although some scattered reports describe mass killings of Dodos for ship provisions, archaeological investigations have found scant evidence of human predation. Bones of at least two Dodos were found in caves at Baie du Cap, which sheltered fugitive slaves and convicts in the 17th century, and were not easily accessible to Dodos due to being isolated in high, broken terrain. The human population on Mauritius never exceeded 50 people in the 17th century, on an island of 1,860 km2, but they introduced other animals, including dogs, pigs, cats, rats, and Crab-eating Macaques, which plundered the Dodo nests, and competed for the limited food resources. At the same time, humans destroyed the Dodo's habitat forests; the impact these introduced animals, especially the pigs and macaques, had on the Dodo population is currently considered as more severe than the impact of hunting. Rats were perhaps not much of a threat to the nests, since Dodos would have been used to dealing with local land crabs. It has been suggested that the Dodo may already have been rare or localised before the arrival of humans on Mauritius, since it is unlikely it would have become extinct so rapidly if it had occupied all remote areas of Mauritius. A 2005 expedition found subfossil remains of Dodos and other animals killed by a flash flood. Such mass mortalities would have further jeopardised a species already in danger of becoming extinct. There is some controversy surrounding the extinction date of the Dodo. The last widely accepted Dodo sighting is the 1662 report by shipwrecked mariner Volkert Evertsz of the Dutch ship Arnhem, who described birds caught on a small islet off Mauritius, now suggested to be Ile d'Ambre: Statistical analysis of the hunting records of Isaac Johannes Lamotius by Roberts & Solow gave a new estimated extinction date of 1693, with a 95% confidence interval of 1688 to 1715; the last reported sighting is from these 1688 hunting records. They also pointed out that because the sighting before 1662 was in 1638, the Dodo was likely already very rare by the 1660s, and thus a disputed report from 1674 by an escaped slave cannot be dismissed out-of-hand. However, Anthony Cheke has pointed out that by this time, some descriptions specifically use the names "Dodo" and "Dodaers" when referring to the Red Rail, indicating that they had been transferred to it after the disappearance of the actual Dodos. Cheke therefore points to the 1662 description as the last credible observation. Until this explanation was proposed, a description of "Dodos" from 1681 was long thought to be the last account. A 1668 account by the English traveller John Marshall, who used the names "Dodo" and "Red Hen" interchangeably for the Red Rail, mentioned the meat was "hard", which mirrors the description of the meat in the 1681 account. However, even the 1662 account has been questioned by Errol Fuller, as the reaction to distress cries matches what was described for the Red Rail. The IUCN Red List accepts Cheke's rationale for 1662 being the date, with all subsequent reports referring to Red Rails, and the Dodo was likely extinct by 1700 in any case, about a century after the discovery of the species in 1598. The Dutch left Mauritius in 1710, but by this time, the Dodo and most of the large, terrestrial vertebrates there had gone extinct. Even though the rareness of the Dodo was reported already in the 17th century, its extinction was not realised until the 19th century. This was partially because, for religious reasons, extinction was not believed possible until later proven by Georges Cuvier, and also because many scientists doubted the Dodo had ever existed. It seemed altogether too strange a creature, and many believed it a myth. The bird was first used as an example of human induced extinction in Penny Magazine, 1833.
The only existing remains of Dodos taken to Europe in the 17th century are a dried head and foot in the Oxford University Museum of Natural History, a foot once housed in the British Museum but now lost, a skull in the University of Copenhagen Zoological Museum, and an upper jaw and leg bones in the National Museum, Prague. The last two were rediscovered and identified as Dodo remains in the mid 1800s. Other stuffed Dodos were mentioned in old museum inventories, but none of these are known to have survived. The only known soft tissue remains, the Oxford head (specimen OUM 11605) and foot, belonged to the last known stuffed Dodo, which was first mentioned as part of the Tradescant collection in 1656, and moved to the Ashmolean Museum in 1659. It has been suggested that this might be the remains of the bird Hamon L'Estrange saw in London. It is a commonly held belief that the museum burned the stuffed Dodo because of severe decay around 1755, only saving the head and leg, due to status eight of the museum, which states "That as any particular grows old and perishing the keeper may remove it into one of the closets or other repository; and some other to be substituted." Its deliberate destruction is now believed a myth. In fact, it was only removed from exhibition to preserve what remained of the specimen. This remaining soft tissue has since degraded further, as the head was dissected by Strickland and Melville in the mid 19th century, separating the skin from the skull in two halves, and the foot is in a skeletal state, with only scraps of skin and tendons. Only very few feathers remain on the head. It has been interpreted as having been a female specimen, as the foot is 11% smaller and more gracile than that of the London specimen, yet appears to be fully grown. The dried London foot, first mentioned in 1665, was long displayed in the British Museum next to Roelant Savery's "Edwards' Dodo" painting, and was also dissected by Strickland and Melville. By 1896 it was mentioned as being without its integuments, and it is thought only the bones remain today. Its present whereabouts are unknown. The Copenhagen skull (specimen ZMUC 90-806) is known to have been part of the collection of Bernardus Paludanus in Enkhuizen until 1651, when it was moved to the museum in Gottorf Castle, Schleswig. After the castle was occupied by Danish forces in 1702, the museum collection was assimilated into the Royal Danish collection, and the skull was rediscovered by J. T. Reinhardt in 1840. It is 13 mm shorter than the Oxford skull, and may have belonged to a female. It is thought to have been mummified originally, but is now devoid of soft tissue. Its sclerotic ring is also preserved. The front part of a skull (specimen NMP P6V-004389) and some leg bones in the National Museum of Prague were found in 1850 among the remains of the Bhmisches Museum. It has been suggested it may be what remains of one of the stuffed Dodos known to have been at the menagerie of Emperor Rudolph II, and perhaps the specimen painted by Hoefnagel or Savery there. Apart from these remains, a dried foot was mentioned by Carolus Clusius in 1605, but is now lost. It belonged to the Dutch professor Pieter Pauw, and its provenance is unknown. It has been suggested it was collected during the Van Neck voyage.
In 1865, government schoolmaster at Mahbourg, George Clark, finally found an abundance of subfossil Dodo bones in the swamp of Mare aux Songes in Southern Mauritius, after searching for many years, having been inspired by Strickland & Melville's monograph. In 1866, Clark explained his procedure to The Ibis, an ornithology journal: Remains of over 300 Dodos were found in the swamp, but only very few skull and wing bones among them, which may be explained by the upper bodies having been washed away or scavenged while the lower body was trapped, which is similar to the way many Moa remains have been found in New Zealand marshes. Clark's reports about the finds rekindled interest in the bird. Sir Richard Owen and Alfred Newton both wanted to be first to describe the post-cranial anatomy of the Dodo, and Owen bought a shipment of Dodo bones originally meant for Newton. Owen described the Dodo bones in Memoir on the Dodo in October 1866, but had erroneously based his Dodo reconstruction on the "Edwards' Dodo" painting by Savery, making it too squat and obese. In 1869, upon receiving more bones, he corrected its stance, making it more upright. Newton moved his focus to the Runion Solitaire instead. The remaining bones not sold to Owen or Newton were auctioned off in London in 1866. In 1889, Thodor Sauzier was commissioned to explore the "historical souvenirs" of Mauritius and find more Dodo remains in the Mare aux Songes. He was successful, and also found remains of other extinct species. A barber named Louis Etienne Thirioux also found many Dodo remains around 1900, including the first remains of a juvenile, which are now lost. He also found the first articulated specimen, but it is unknown exactly where he made his finds, except that some were found in a cave. It has been suggested the cave was located on the slopes of the Le Pouce Mountain. 26 museums worldwide have significant holdings of Dodo material, almost all found in the Mare aux Songes. The Natural History Museum, American Museum of Natural History, Cambridge University Museum of Zoology, the Senckenberg Museum and others, have almost complete specimens assembled from these dissociated subfossil remains. An alleged Dodo egg is stored at the East London Museum in South Africa. It was donated by Marjorie Courtenay-Latimer, whose great aunt had received it from a captain who claimed he had found it in a swamp on Mauritius. Genetic studies are under way to determine its authenticity. In October 2005, after a hundred years of neglect, a part of the Mare aux Songes swamp was excavated by an international team of researchers. To prevent malaria, the British had covered the swamp with hard core during their rule over Mauritius, which had to be removed. Many remains were found, including bones of birds in various stages of maturity, and several bones obviously from the skeleton of one individual bird, which have been preserved in their natural position. These findings were made public in December 2005 in the Naturalis museum in Leiden. 63% of the fossils found in the swamp belonged to turtles of the extinct Cylindraspis genus, and 7.1% belonged to Dodos, which had been deposited within several centuries, 4000 years ago. Subsequent excavations suggested that Dodos, along with other animals, became mired in the Mare aux Songes while trying to reach water during a long period of severe drought about 4,200 years ago. In June 2007, adventurers discovered the most complete and best-preserved Dodo skeleton ever found while exploring a cave in Mauritius. The specimen was nicknamed "Fred" after the finder. In 2011, a wooden box containing Dodo bones from the Edwardian era was rediscovered at the Grant Museum, during preparations for a move.
The supposed "White Dodo" (or "Solitaire") of Runion is now believed to have been an erroneous conjecture based on contemporary reports of the Runion Sacred Ibis, combined with paintings by Pieter Withoos and Pieter Holsteyn from the 1600s of white Dodos that surfaced in the 19th century. The confusion began when Willem Ysbrandtsz. Bontekoe, who visited Runion around 1619, mentioned birds with Dodo like features and with a name similar to "Dodaars" in his journal, though without mentioning colouration: When the journal was published in 1646, it was accompanied by a copy of Roelant Savery's "Crocker Art Gallery sketch". A white, flightless bird on Runion was first described by Chief Officer J. Tatton in 1625. In 1674, such birds were again described by Sieur D. B. Dubois: Baron Edmund de Slys-Longchamps coined the name Raphus solitarius for these birds in 1848, as he believed the accounts referred to a species of Dodo. When 17th century paintings of white Dodos were discovered by 19th century naturalists, it was assumed they depicted these birds. Walter Rothschild suggested that the reason the painted specimens had yellow wing-tips instead of black as in the old descriptions might have been albinism. It was also suggested that differences in paintings were due to sexual dimorphism. Others believed it was a species similar to the Rodrigues Solitaire, as it was referred to by the same name, or even that there were white species of both the Dodo and Solitaire on the island. The Pieter Withoos painting, which was discovered first, appears based on an earlier painting by Pieter Holsteyn, three versions of which are known to have existed. According to Julian Hume and Anthony Cheke, it appears that all depictions of white Dodos were based on a single painting or copies of it, showing a whitish specimen, made by Roelant Savery in ca. 1611 called "Landscape with Orpheus and the animals". This was apparently based on a stuffed specimen then in Prague; a walghvogel described as having a "dirty off-white colouring" was mentioned in an inventory of specimens in the Prague collection of the Holy Roman Emperor Rudolf II to whom Savery was contracted at the time (16071611). Savery's several later images all show grayish birds, possibly because he had by then seen another specimen. It has also been suggested that the light plumage was a juvenile trait, a result of bleaching of old taxidermied specimens, or simply due to artistic license. Since Runion was not visited by Europeans until 1635, the 1611 painting could not have shown a bird from there. In 1987, fossils of a recently extinct species of ibis from Runion with a relatively short beak, Borbonibis latipes, were described, before a connection to the Solitaire reports had been made. Anthony Cheke suggested to one of the authors, Francois Moutou, that the fossils may have been of the Runion Solitaire, and this suggestion was published in 1995. The ibis was also reassigned to the genus Threskiornis, now combined with the specific epithet la from the binomial Raphus solitarius. Birds of this genus are also white and black with slender beaks, which fits the old descriptions of the Runion Solitaire. No fossil remains of Dodo-like birds have ever been found on the island.
The Dodo's significance as one of the best-known extinct animals and its singular appearance has led to its use in literature and popular culture as symbol of an outdated concept or object, as in the expression "dead as a Dodo," which has come to mean unquestionably dead. Similarly, the phrase "to go the way of the Dodo" means to become extinct or obsolete, to fall out of common usage or practice or to become a thing of the past. In 1865, the same year that George Clark started to publish his reports about excavated Dodo fossils, the newly vindicated bird was featured as a character in Lewis Carroll's Alice's Adventures in Wonderland. It is thought he included the Dodo because he identified with it, and had adopted the name as a nickname for himself due to his stammer, which made him accidentally introduce himself as "Do-do-dodgson", his real surname. With the book's popularity, the Dodo became a well-known icon of extinction. Today, the Dodo regularly appears in works of popular fiction, and is used as a mascot for many kinds of products, especially in Mauritius. The Dodo appears as a supporter on the coat of arms of Mauritius. A smiling Dodo is the symbol of the Brasseries de Bourbon, a popular brewer on Runion Island, in reference to the white species once thought to have lived there. The Dodo is used to promote the protection of endangered species by many environmental organisations, such as the Durrell Wildlife Conservation Trust and the Durrell Wildlife Park. In 2011, the nephilid spider Nephilengys dodo, which inhabits the same woods as the Dodo once had, was named after the bird to raise awareness of the urgent need for protection of the Mauritius biota. In 2009, a previously unpublished 17th century Dutch illustration of a Dodo went for sale at Christie's and was expected to sell for 6,000. It is unknown whether the illustration was based on a specimen or on a previous image. It was sold for 44,450. The poet Hilaire Belloc included the following poem about the Dodo in his The Bad Child's Book of Beasts from 1896: