Dodo

• 1. Introduction

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  The dodo, Raphus cucullatus, was a flightless bird found nowhere on Earth except Mauritius, an island east of Madagascar in the Indian Ocean. Today it stands as a shorthand for extinction and obsolescence. Millions of readers first met it in Alice's Adventures in Wonderland. Yet almost everything about the real animal is uncertain. Its appearance in life rests only on drawings, paintings and written accounts from the 17th century, and the artists and writers disagreed with one another. Its closest relative was another extinct flightless bird, the Rodrigues solitaire. Its closest living relative is a pigeon. How did a pigeon become a heavy ground-dweller that could not fly? Why did the first naturalists who studied it take it for an ostrich or a vulture? What actually wiped it out, and what survives of it in museums today? The answers are stranger than the cartoon.

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• 2. A Pigeon Revealed

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  Early scientists variously declared the dodo a small ostrich, a rail, an albatross or a vulture. Nobody had a living bird to examine, so every guess was made from fragments.

  In 1842 the Danish zoologist Johannes Theodor Reinhardt proposed that dodos were ground pigeons. He based this on a dodo skull he had discovered in the collection of the Natural History Museum of Denmark. His idea was met with ridicule. Support arrived in 1848, when the English naturalists Hugh Edwin Strickland and Alexander Gordon Melville published a monograph titled The Dodo and Its Kindred, which tried to separate myth from reality. They dissected the preserved head and foot of the Oxford specimen and compared them with the few known remains of the Rodrigues solitaire, Pezophaps solitaria. They concluded that the two birds were closely related.

  Strickland and Melville listed a run of features that the dodo shared only with pigeons. The horny part of the beak was very short, and its base was long, slender and bare. The nostril sat low on the middle of the beak, ringed by skin, beneath a high forehead. Bare skin around the eyes reached almost to the beak, as in other pigeons. The legs matched those of ground-living pigeons in both scales and skeleton. Pictures of a large crop pointed the same way, and so did the report of a clutch of a single egg. The dodo differed mainly in having tiny wings and a beak that was huge compared with its cranium.

  Classification stayed tangled for decades. During the 19th century several species were lumped with the dodo, including the Rodrigues solitaire and the Réunion solitaire, under names such as Didus solitarius and Raphus solitarius. In 1852 Abraham Dee Bartlett named a new species, Didus nazarenus, from an odd 17th-century description and bones found on Rodrigues. Those bones belonged to the solitaire, so the name is now a synonym. Crude drawings of the red rail of Mauritius were also misread as dodo species, called Didus broeckii and Didus herberti. For many years the dodo and the solitaire sat in a family of their own, the Raphidae, because their place among the pigeons was unresolved. Studies of bones and DNA eventually dissolved that family. In 2024 a new subtribe, Raphina, was created to hold only the dodo and the solitaire.

  The first genetic test came in 2002, when the American geneticist Beth Shapiro and colleagues analysed dodo DNA for the first time. They compared mitochondrial cytochrome b and 12S rRNA sequences taken from a tarsal bone of the Oxford specimen and a femur of a Rodrigues solitaire. The sequences confirmed the close kinship of the two birds and placed them inside the pigeon family, Columbidae. The Southeast Asian Nicobar pigeon, Caloenas nicobarica, emerged as the closest living relative. Next came the crowned pigeons of New Guinea, then the tooth-billed pigeon of Samoa, whose scientific name refers to its dodo-like beak. A similar tree published in 2007 swapped the order of two of these groups and added the pheasant pigeon and the thick-billed ground pigeon at its base. There is a caveat. The DNA came from the degraded Oxford specimen, and no usable DNA has been pulled from subfossil bones, so the results still need independent checking. In 2014 DNA from the only known specimen of the recently extinct spotted green pigeon, Caloenas maculata, showed that it too was a close relative of the Nicobar pigeon, and therefore of the dodo.

  The 2002 study suggested that the dodo and solitaire lineages split about 23.03 million years ago, near the Paleogene-Neogene boundary. The Mascarene Islands, which are Mauritius, Réunion and Rodrigues, are volcanic and less than 10 million years old. So the ancestors of both birds probably kept the power of flight for a long time after their lineage divided. The Nicobar and spotted green pigeons sit at the base of the line leading to the dodo group, which implies flying, semi-terrestrial island dwellers. That supports the idea that the ancestors reached the Mascarenes by hopping between islands from South Asia. With no mammalian herbivores competing for food, the solitaire and the dodo grew very large and lost flight. Even so, many features of the dodo skeleton stayed close to those of smaller flying pigeons.

  The dodo group was not the only giant of its kind. In 2001 researchers described the Viti Levu giant pigeon, Natunaornis gigoura, from subfossil material found in Fiji. It was only slightly smaller than the dodo and the solitaire, and it too is thought to be a relative of the crowned pigeons.

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• 3. What's in a Name

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  The earliest name for the dodo was a Dutch insult. "Walghvoghel" first appears in the journal of Vice admiral Wybrand van Warwijck, who visited Mauritius in 1598 during the Second Dutch Expedition to Indonesia. Walghe means tasteless, insipid or sickly, and voghel means bird. Jakob Friedlib rendered it in German as Walchstök or Walchvögel. The original Dutch report, Waarachtige Beschryving, was lost, but an English translation survives. It says the crew found great fowl twice as big as swans and enjoyed them at first. Then they found plenty of pigeons and parrots and scorned the large birds, calling them Wallowbirds, which the translator glossed as "lothsome or fulsome birdes."

  A second account from that voyage, perhaps the first to mention the dodo, says the Portuguese called the birds penguins. That name may not come from the penguin at all, since the Portuguese called those birds "fotilicaios" at the time. It may instead come from pinion, a nod to the small wings. In 1602 the crew of the Dutch ship Gelderland used "Dronte", meaning swollen, a name still used in some languages. The same crew also said "griff-eendt" and "kermisgans", comparing the birds to fowl fattened for the Kermesse festival in Amsterdam, which had been held the day after the ship anchored on Mauritius.

  The word dodo itself has a murky origin. Some link it to the Dutch dodoor, meaning sluggard. A likelier source is Dodaars, which means either fat-arse or knot-arse and refers to the knot of feathers at the rear. Dodaars is first recorded in Captain Willem Van West-Zanen's journal in 1602. Sir Thomas Herbert was the first to print the word dodo, in his 1634 travelogue, where he said the Portuguese used it after visiting Mauritius in 1507. Another Englishman, Emmanuel Altham, had written it in a 1628 letter and also credited the Portuguese. A variant, dodar, entered English at the same time but was used only until the 18th century. As far as is known, the Portuguese never mentioned the bird. Even so, some sources still derive dodo from the Portuguese doudo, meaning fool or crazy. Another suggestion is that it imitates the bird's call, a two-note pigeon-like sound resembling doo-doo.

  The Latin name has its own trail. In 1635 the Spanish Jesuit Juan Eusebio Nieremberg coined cucullatus, meaning hooded, in the combination Cygnus cucullatus. He was describing Carolus Clusius's 1605 depiction of a dodo. In the tenth edition of Systema Naturae, Carl Linnaeus kept cucullatus as the species name but paired it with Struthio, the ostrich. Mathurin Jacques Brisson coined the genus name Raphus in 1760, referring to the bustards, which produced the modern Raphus cucullatus. In 1766 Linnaeus tried again with Didus ineptus, meaning inept dodo. Because of the rule of priority that name lost out to the older one and became a synonym, though the reputation it expressed lingered in popular pictures for far longer.

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• 4. Portraits and Weights

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  The Dutch ship Gelderland kept a travel journal from 1601 to 1603 that was rediscovered in the 1860s. It holds the only known sketches of living or recently killed dodos drawn on Mauritius itself. They are attributed to the professional artist Joris Joostensz Laerle, who also drew other now-extinct Mauritian birds, and to a second, less refined hand. Of the roughly twenty 17th-century illustrations of dodos, it is unknown how many were drawn from live birds and how many from stuffed ones. That uncertainty matters, because paintings and written accounts are the main evidence for what the bird looked like.

  The Dutch painter Roelant Savery was the most prolific and influential dodo illustrator, with at least twelve depictions, often tucked into the lower corners of his scenes. His 1626 painting, now called Edwards's Dodo because the ornithologist George Edwards once owned it, hangs in the Natural History Museum in London. It shows a particularly fat bird and has become the standard image of a dodo. Many later illustrations copy it.

  A very different bird turned up in 1955, when Aleksander Iwanow rediscovered an Indian Mughal painting in the Hermitage Museum in St. Petersburg. It shows a slimmer, brownish dodo among native Indian birds, which are clearly identifiable and correctly coloured. Iwanow and the British palaeontologist Julian Hume regarded it as one of the most accurate depictions of the living dodo. It is believed to date from the 17th century and has been attributed to the Mughal painter Ustad Mansur. The bird probably lived in the menagerie of the Mughal Emperor Jahangir in Surat, where the English traveller Peter Mundy claimed to have seen two dodos between 1628 and 1633.

  Scientists today suspect that the fat, clumsy dodo of tradition is exaggerated. Many old European pictures were probably based on overfed captive birds or on crudely stuffed specimens. Another suggestion is that some images show dodos with puffed-up feathers during a display. In 2014 a newly reported Indian illustration turned out to copy an 1836 German picture. All depictions after 1638 appear to be based on earlier ones, around the time that reports of dodos became rare. Hume argued that the living dodo had slit nostrils, as seen in the Gelderland sketches, in Cornelis Saftleven's work, in Savery's Crocker Art Gallery sketch and in the Mansur painting. In his view, the gaping nostrils in many paintings suggest that taxidermy specimens served as models. Most pictures show the wings held out, like those of an ostrich or a kiwi and unlike those of flying pigeons.

  Written descriptions come mostly from the logs and journals of Dutch East India Company ships that stopped at Mauritius while the Dutch Empire ruled it. Few are reliable, since many copy earlier accounts and none was written by a scientist. Van Warwijck described birds "larger than our swans", with huge heads half covered in skin as if hooded, and a tail of a few soft ash-coloured feathers. Herbert's 1634 account is among the most detailed. He compared the bird for rareness to the Phoenix of Arabia and wrote that "her body is round and fat, few weigh less than fifty pound." He judged it more a wonder than a meal.

  Most representations agree on greyish or brownish plumage, a tuft of curly light feathers high on the rear, and a bare grey head. The beak was green, black and yellow, and the stout yellowish legs ended in black claws. A study of the few feathers left on the Oxford head found them pennaceous rather than downy, most like those of other pigeons. Details such as beak markings, tail form and colouring vary so much from account to account that nobody can say whether they show age, sex or invention.

  The bird stood about 62.6-75 cm tall and was sexually dimorphic, with larger males that had proportionally longer beaks. Weight is a long-running dispute. In 1993 Bradley C. Livezey proposed 21 kg for males and 17 kg for females. In the same year Andrew C. Kitchener blamed high estimates, and the roundness of dodos in European pictures, on birds overfed in captivity. He put wild weights at 10.6-17.5 kg and fattened birds at 21.7-27.8 kg. A 2011 estimate by Angst and colleagues gave an average as low as 10.2 kg, which has also been questioned. A 2016 study using CT scans of composite skeletons arrived at 10.6 to 14.3 kg.

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• 5. Life on Mauritius

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  A 1601 map in the Gelderland journal marks a small island off the coast of Mauritius where dodos were caught. Julian Hume has suggested it was l'île aux Bénitiers in Tamarin Bay, on the west coast. Old descriptions place the dodo in the woods of the drier coastal areas of the south and west. That fits the Mare aux Songes swamp, where most dodo remains have been dug up, since it lies close to the sea in the south-east. Subfossil bones have also turned up in caves in highland areas, so the bird once ranged onto the mountains. Around the swamp the vegetation was dominated by tambalacoque and Pandanus trees and endemic palms. Before people arrived, forest covered all of Mauritius, and very little of it remains. The dodo lived alongside the flightless red rail, the broad-billed parrot, the Mauritius blue pigeon and the saddle-backed Mauritius giant tortoise, all now gone.

  Very little is known about the dodo's behaviour, because the descriptions are brief. Tests of the strength of its leg bones suggest it could run quite fast, and the sturdy legs also made it agile in dense forest. The wings were small, but scars for well-developed muscles show they were not useless. They may have served for display and balance, as in living pigeons. Unlike the Rodrigues solitaire, the dodo shows no sign of using its wings to fight its own kind. It may have used its large hooked beak in territorial disputes instead. Mauritius gets more rain and has less seasonal swing than Rodrigues, so the dodo had less reason to evolve aggression, and the solitaire was probably the fiercer bird. In 2016 researchers made the first 3D endocast of a dodo brain. Its size relative to the body matched that of modern pigeons, so the dodo was probably about as intelligent.

  The only account of the dodo's diet is a Dutch letter from 1631, long thought lost and rediscovered in 2017. It uses word-play, with the dodos presumably standing in for wealthy mayors, and says "their war weapon was the mouth, with which they could bite fiercely." Their food, it says, was raw fruit. Beyond fallen fruit, the bird probably ate nuts, seeds, bulbs and roots, and perhaps crabs and shellfish like its relatives the crowned pigeons. Anthonie Cornelis Oudemans suggested that, because Mauritius has marked dry and wet seasons, the dodo fattened on ripe fruit at the end of the wet season to survive the dry one. In 1996 the Mauritian ornithologist France Staub proposed that it fed mainly on palm fruits and tried to match its fat cycle to the fruiting of the palms. The upper jaw was rhynchokinetic, meaning its parts could move relative to each other, which helps living fruit pigeons handle large items. The beak could also take heavy loads, which points to hard food. The brain endocast showed a comparatively large olfactory bulb, so the dodo may have smelled out fruit.

  Sir Hamon L'Estrange saw a live bird in London around 1638 and called it "somewhat bigger than the largest Turkey cock." Its keeper showed him a heap of large pebbles at the end of a chimney, some as big as nutmegs, which the bird swallowed to help digestion. L'Estrange was confident that she later cast them all up again. Depictions show a large crop, probably used for food storage and for making crop milk, the way related pigeons feed their young. It has been proposed that the amount of crop milk they could produce limited how big the dodo and solitaire could grow.

  The tambalacoque, or dodo tree, gave rise to a famous idea. In 1973 it was thought to be dying out on Mauritius, with only 13 specimens left, all about 300 years old. Stanley Temple hypothesised that its seeds germinated only after passing through a dodo's gut, so the tree was nearly coextinct with the bird. Temple overlooked reports from the 1940s showing that seeds germinated, if very rarely, without being abraded in digestion. In 1991 the ecologists Wendy Strahm and Anthony Cheke noted that the tree had germinated since the dodo vanished and numbered several hundred. Others suggested that giant tortoises of the genus Cylindraspis, fruit bats or the broad-billed parrot also spread the seeds. In 1997 Carlos Yamashita suggested the broad-billed parrot itself depended on dodos and tortoises to eat and excrete palm fruits.

  Breeding is known mostly from the 1651 account of François Cauche. He described birds bigger than a swan with "a cry like a gosling" and a single white egg beside which they placed a white stone. His account also gives the bird three toes and no tongue, unlike a dodo. Some took that to mean he had seen a separate species, and others think his description was mixed with that of a cassowary. An egg held at the East London Museum in South Africa is claimed to be a dodo's. It was donated by Marjorie Courtenay-Latimer, whose great aunt had received it from a captain who said he found it in a swamp on Mauritius. In 2010 the museum's curator proposed genetic tests, and it may turn out to be an aberrant ostrich egg.

  A single-egg clutch and a large body suggest the dodo was K-selected, producing few young that needed parental care until maturity. No juvenile dodos have been found in the Mare aux Songes. A 2017 study of thin-sectioned bones, local ecology and old accounts proposed that dodos bred around August, after fattening. The chicks grew fast to near adult size and reached sexual maturity before the Austral summer and the cyclone season. Adults that had just bred moulted around March, replacing wing and tail feathers first, and finished by the end of July. Birds at different stages of moult may explain why the old descriptions of plumage disagree.

  A 2023 study by Julia H. Heinen and colleagues concluded that losing the dodo, which probably preferred large seeds and fruits, hurt seed dispersal for native plants. Introduced species handle seeds differently, so they do not restore the function.

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• 6. Sailors and Specimens

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  Portuguese ships called at Mauritius between 1507 and 1513, and Arab vessels had visited in the Middle Ages, but neither group settled. No record of dodos from these visits is known, although the Portuguese name for the island, Cerne (swan) Island, may refer to the birds. The Dutch Empire acquired Mauritius in 1598, renamed it after Maurice of Nassau, and used it to supply the trade vessels of the Dutch East India Company. The earliest dodo accounts come from Dutch travellers on the expedition led by admiral Jacob van Neck in 1598. They appear in reports published in 1601, which also carry the first published illustration of the bird.

  The first sailors had been at sea a long time, so their interest in the big birds was mostly culinary. West-Zanen's 1602 journal from the ship Bruin-Vis records 24-25 dodos hunted for food, so large that two could scarcely be eaten at one meal. The leftovers were preserved with salt. An illustration made for the 1648 edition of the journal shows dodos being killed, along with a dugong and possibly Mascarene grey parakeets. Its Dutch caption poem, in Strickland's translation, says the seamen "tap the palms, and round-rumped dodos they destroy." Opinions on the taste varied. Some early travellers found dodo meat unsavoury and preferred parrots and pigeons, while others called it tough but good. Some hunters took the birds only for their gizzards, thought the finest part. The birds were easy to catch, but hunters had to avoid their powerful beaks.

  The oddness of the dodo and the red rail pushed Peter Mundy to a remarkable guess, 230 years before Charles Darwin's theory of evolution. Neither bird was known anywhere else, although the island lay about 100 leagues from St. Lawrence, and neither could fly or swim. Mundy wondered whether strange forms came from a mixture of kinds or whether climate, air and earth altered the first shapes over a long time.

  Live dodos were sent to Europe and the East, but how many arrived alive is uncertain. From a combination of accounts, paintings and specimens, Julian Hume inferred that at least eleven reached their destinations alive. L'Estrange's London bird of 1638 is the only account that explicitly describes a live dodo in Europe. In 1626 Adriaen van de Venne drew a dodo he claimed to have seen in Amsterdam, though he did not say whether it was alive. In 1628 Emmanuel Altham wrote to his brother in England that he was sending a dodo with Mr. Perce, adding "if it live." Whether it survived is unknown, and the letter was destroyed by fire in the 19th century.

  The earliest known picture of a dodo specimen in Europe is from a collection of paintings dated about 1610, showing animals in the royal menagerie of Emperor Rudolph II in Prague. The bird, which may be a juvenile, looks dried or embalmed, and probably lived in the emperor's zoo for a while. Stuffed dodos in Europe imply that the birds arrived alive and died there, since taxidermists were unlikely to be aboard ships and spirits were not yet used to preserve specimens.

  The farthest journey ended in Nagasaki, Japan, in 1647. For a long time nobody knew whether that dodo ever arrived. Documents first published in 2014 proved the story and showed that it arrived alive. It was meant as a gift and, despite its rarity, was rated equal in value to a white deer and a bezoar stone. It is the last recorded live dodo in captivity.

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• 7. The Vanishing

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  The human population of Mauritius, an island of 1860 km2, never exceeded 50 people in the 17th century. Even so, the dodo vanished. Like many animals that evolved without strong predators, it was entirely fearless of people, and being flightless it made easy prey. Yet hunting was not the main cause of extinction, contrary to popular belief. Scattered reports describe mass killings for ship provisions, but archaeological work has 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. Those caves sat on high, broken terrain that dodos would not easily have reached.

  The settlers brought dogs, pigs, cats, rats and crab-eating macaques. These animals plundered dodo nests and competed for the limited food, while people also cleared the forest habitat. Today the effect of the introduced animals, especially the pigs and macaques, is judged more severe than hunting. Rats may have been less of a threat, since dodos would have been used to dealing with local land crabs. The dodo may also have been rare or localised before humans came, because it would be unlikely to vanish so fast had it filled every remote corner of the island. A 2005 expedition found subfossil remains of dodos and other animals killed by a flash flood, a mass death that would have added risk to a species already in danger. Yet the bird had survived hundreds of years of volcanic activity and climate change, which shows it was resilient within its ecosystem.

  The date of the last dodo is disputed. The last widely accepted record is a 1662 report by Volkert Evertsz, a shipwrecked mariner from the Dutch ship Arnhem. He described birds caught on a small islet off Mauritius, now suggested to be Amber Island. The animals stared at the men, "not knowing whether they had wings to fly away or legs to run off." When one was held by the leg and cried out, the others came running to help and were caught too. These may not have been the last dodos. Isaac Johannes Lamotius recorded a claimed sighting in 1688. In 2003 the biologists David L. Roberts and Andrew R. Solow analysed those records and estimated extinction in 1693, with a 95% confidence interval of 1688-1715. They noted that the sighting before 1662 dated from 1638, so the dodo was probably quite rare by the 1660s and a disputed 1674 report by an escaped slave should not be dismissed outright.

  Alfred Newton suggested in 1868 that the dodo's name was transferred to the red rail after the dodo died out. Cheke noted that some descriptions after 1662 use the names Dodo and Dodaers for the red rail, so he regarded the 1662 description as the last credible one. An English traveller, John Marshall, wrote in 1668 and used Dodo and Red Hen interchangeably for the red rail. Even the 1662 account has been questioned by the writer Errol Fuller, since the birds' reaction to distress cries matches what is described for the red rail. Until that explanation was proposed, a description of dodos from 1681 was thought the last, and some still defend that date. In 2014 Cheke said newly accessible Dutch manuscripts show that settlers saw no dodos in 1664-1674. In 2020 Cheke and the British researcher Jolyon C. Parish proposed that all later mentions were of red rails and that feral pigs destroyed the dodo during a gap in settlement from 1658 to 1664. Nobody noticed, because new settlers expected flightless birds and gave the old name to the red rail. Red rails probably bred more efficiently, with larger clutches, faster incubation and perhaps hidden nests. The IUCN Red List accepts Cheke's reasoning and chooses 1662. In any case the dodo was probably gone by 1700, about a century after its discovery in 1598. The Dutch left Mauritius in 1710.

  The extinction went unrecognised for generations. For religious reasons, extinction was not believed possible until Georges Cuvier proved otherwise, and many scientists doubted that the dodo had ever existed. It seemed too strange a creature, and many thought it a myth. In 1833 Penny Magazine used the bird as an example of human-induced extinction for the first time, and it has been called an icon of extinction ever since.

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• 8. Bones and Heads

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  The Oxford University Museum of Natural History holds a dried head and foot, specimen OUM 11605, the only soft tissue of a dodo left anywhere. Only a handful of remains from the 17th century survive: that head and foot, a lost foot once in the British Museum, a skull in Copenhagen and an upper jaw in Prague. The stuffed dodos seen in museums around the world are in fact made from the feathers of other birds, many by the company of the British taxidermist Rowland Ward.

  The Oxford specimen was first mentioned in the Tradescant collection in 1656 and moved to the Ashmolean Museum in 1659. In 2018 scans showed that its skin and bone contain lead shot, used to hunt birds in the 17th century. So the bird was shot either before shipping or sometime after reaching Britain. Many sources claim the Ashmolean burned the stuffed dodo around 1755 because of decay and kept only the head and leg. That deliberate destruction is now believed to be a myth, since the specimen was simply removed from display to preserve what remained. The foot is 11% smaller and more slender than the London foot, so the bird was probably a female. The head was shown at Oxford from at least the 1860s until 1998, and has mostly been kept in storage since to prevent damage.

  The Copenhagen skull, specimen ZMUC 90-806, was part of the collection of Bernardus Paludanus in Enkhuizen until 1651, when it moved to the museum at Gottorf Castle in Schleswig. After Danish forces occupied the castle in 1702, the collection joined the Royal Danish collection. J. T. Reinhardt rediscovered the skull in 1840, and it may be the oldest known surviving dodo remains brought to Europe. The Prague fragment, specimen NMP P6V-004389, was found in 1850 among the remains of the Böhmisches Museum and may come from a stuffed bird in Rudolph II's menagerie.

  Until 1860 those four incomplete specimens were all that was known. Philip Burnard Ayres found the first subfossil bones that year and sent them to Richard Owen at the British Museum, who did not publish them. In 1865 George Clark, the government schoolmaster at Mahébourg, finally found abundant dodo bones in the Mare aux Songes after a 30-year search inspired by Strickland and Melville's monograph. He explained in The Ibis that he sent workers to wade through the swamp, feeling for bones with their feet. They found few bones until they cleared the plants from the deepest part. Harry Pasley Higginson, a railway engineer from Yorkshire, reported finding the bones at the same time, and the dispute over who came first is unresolved. The swamp yielded the remains of over 300 dodos, but very few skull and wing bones.

  Owen and Newton both wanted to be first to describe the dodo's body below the skull. Owen bought a shipment of bones meant for Newton, and a rivalry began. In October 1866 Owen published Memoir on the Dodo, but he based his reconstruction on Savery's Edwards's Dodo, making it too squat and obese. When more bones arrived in 1869 he corrected the stance to a more upright one. Newton turned to the Réunion solitaire.

  Later collectors kept digging. In 1889 Théodor Sauzier was commissioned to explore the historical souvenirs of Mauritius and found more remains in the Mare aux Songes. Around 1900 the amateur naturalist Louis Étienne Thirioux found remains at several sites, including the first subfossil dodo skeleton outside the swamp. It was found in 1904 in a cave near Le Pouce mountain and is the only known complete skeleton of an individual dodo. He donated it to the Museum Desjardins, now the Natural History Museum in Port Louis. His heirs sold a second, composite mounted skeleton to the Durban Museum of Natural Science in South Africa in 1918. These two skeletons were largely forgotten until 2011, when a group of researchers sought them out. They were laser scanned, and the 3-D models became the basis of a 2016 monograph on the dodo's bones. In 2006 explorers found a complete dodo skeleton in a lava cave in Mauritius, only the second associated skeleton ever found.

  In 2005 an international team, the International Dodo Research Project, excavated part of the Mare aux Songes after a century of neglect. To prevent malaria, the British had covered the swamp with hard core, which had to be removed. The dig found bones of at least 17 dodos of varying maturity, though no juveniles, and the skeleton of one bird preserved in its natural position. Results were announced in December 2005 at the Naturalis museum in Leiden. Of the fossils, 63% belonged to turtles of the extinct genus Cylindraspis and 7.1% to dodos, laid down over several centuries about 4,000 years ago. Later digs suggested that dodos and other animals were mired in the swamp while trying to reach water during a severe drought about 4,200 years ago. Cyanobacteria thrived on the animals' droppings, and the crowd died of poisoning, thirst, trampling and miring. Worldwide, 26 museums hold significant dodo material, almost all from the Mare aux Songes. In 2011 a wooden box of Edwardian-era dodo bones was rediscovered at the Grant Museum at University College London, where it had been stored with crocodile bones.

  One more phantom bird belongs to this story, the supposed white dodo of Réunion. It is now considered a mistaken conjecture, built on reports of the Réunion ibis and on 17th-century paintings by Pieter Withoos and Pieter Holsteyn of white, dodo-like birds that surfaced in the 19th century. The confusion began with Willem Ysbrandtszoon Bontekoe, who visited Réunion around 1619 and mentioned fat flightless birds he called "Dod-eersen", without giving their colour. A white, stocky flightless bird was first listed in the island's fauna by Chief Officer J. Tatton in 1625. In 1848 Baron Edmond de Sélys Longchamps coined Raphus solitarius for these birds, believing they were a dodo species. The engraving printed with Bontekoe's journal in 1646 came from Savery's Crocker Art Gallery sketch, even though Bontekoe never mentioned colour.

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