Dodo

For other uses, see Dodo (disambiguation).

Dodo Temporal range: Late Holocene
Dodo skeleton cast and model based on modern research, at Oxford University Museum of Natural History
Conservation status
Extinct (about 1662)  (IUCN 3.1)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Order:	Columbiformes
Family:	Columbidae
Subfamily:	Raphinae
Genus:	Raphus Brisson, 1760
Species:	R. cucullatus
Binomial name
Raphus cucullatus (Linnaeus, 1758)
Former range (area in blue)
Synonyms
Struthio cucullatus Linnaeus, 1758 Didus ineptus Linnaeus, 1766

The Dodo (Raphus cucullatus) is an extinct, flightless bird that was endemic to the Mascarene island of Mauritius east of Madagascar in the Indian Ocean. It is genetically related to pigeons and doves, and its closest relative is the likewise extinct Rodrigues Solitaire, the two forming the Raphinae subfamily. The closest living relative is the Nicobar Pigeon. A white Dodo was believed to have existed on the nearby island of Réunion, but this is now known to be incorrect.

The Dodo was about one metre (3.3 ft) tall and may have weighed 10–18 kg (22–40 lb) in the wild. Its external appearance is evidenced only by paintings and written accounts from the 17th century. Because these vary considerably, and only a few sketches are known to have been drawn from life, its exact appearance remains a mystery. The same is true of its habitat and behaviour. It was depicted with brownish grey plumage, yellow feet, and a tuft of tail feathers, a grey, naked head, and a beak coloured black, yellow and green. It is assumed that the Dodo became flightless because of abundant food sources and the absence of predators on Mauritius.

The Dodo was first mentioned by Dutch sailors in 1598. Hereafter the bird was preyed upon by hungry sailors, their domesticated animals, and other invasive species introduced during that time, and the last credible sighting is from 1662. The extinction was not immediately noticed, and some considered it a mythological creature until the 19th century, when research was conducted on surviving remains of four specimens 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 a century of its discovery called attention to the previously unrecognised human involvement in extinction.

The Dodo achieved widespread recognition due to a notable role in Alice in Wonderland, and has since become a fixture in popular culture, sometimes as a symbol of extinction and obsolescence.

Taxonomy and evolution

1848 lithograph of the Oxford specimen skull

Many different affinities have historically been suggested for the Dodo, including that it was a small ostrich, a rail, an albatross, or a vulture.

In 1842, Johannes Theodor Reinhardt proposed they were ground doves, based on studies of a Dodo skull he had rediscovered in the royal Danish collection of Copenhagen. This view was met with ridicule, but later supported by Hugh Edwin Strickland and Alexander Gordon Melville in 1848, after their dissection of the preserved head and foot of the specimen at Oxford Museum, and they also found that the Dodo was closely related to the likewise extinct Rodrigues Solitaire.

The theory has recently been confirmed by mtDNA cytochrome b and 12S rRNA sequence analysis, wherein DNA extracted from a tarsal from the Dodo foot in Oxford was compared with genetic material from other birds, indicating that the ancestors of the Dodo diverged from those of its closest known relative, the Rodrigues Solitaire, around the Paleogene-Neogene boundary. The Mascarene Islands, a group of islands in the Indian Ocean east of Madagascar, which includes Mauritius and Réunion, are of volcanic origin and less than 10 million years old, so the ancestors of both birds likely remained capable of flight for some considerable time after the separation of their lineages.

The same study has been interpreted to show that the closest living relative of the Dodo is the Southeast Asian Nicobar Pigeon. Other genetically related pigeons are the Crowned Pigeons of the genus Goura, and the superficially Dodo-like Tooth-billed Pigeon from Samoa. The generic name of the latter is Didunculus, which means "little dodo", and it was also referred to as "Dodlet" by Richard Owen. However, this proposed phylogeny is problematic regarding the relationships of other taxa.

A sketch of the Oxford specimen, which is the main source for genetic studies

The following cladogram shows the Dodo's closest relationships within columbidae, based on Shapiro et al., 2002.

	Goura victoria Caloenas nicobarica Pezophaps solitaria Raphus cucullatus Didunculus strigirostris

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.

For a long time, the Dodo and the Rodrigues Solitaire were placed in a family of their own, the Raphidae, formerly "Dididae". This was because their relationships to other groups of birds, such as rails, was unresolved. Recently, it has been suggested that the group should be dissolved, and the Dodo and Solitaire placed in the existing subfamily Raphinae within the Columbidae.

Throughout the 19th century, several species were classified as being congeneric with the Dodo, including the Rodrigues Solitaire and the Réunion 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 to the erection of a new species, Didus nazarenus. 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.

Etymology

Labeled sketch from 1634 by Sir Thomas Herbert, showing a Broad-billed Parrot, a Red Rail ("cacato" and "hen"), and a Dodo

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 used the word Dodo in English in 1628, but it is unclear whether he was the first; the Portuguese had visited the island in 1507, but did not use the word. Nevertheless, some sources state that "Dodo" derives from Portuguese 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 like "doo-doo".

One of the original names for the Dodo was "walghvogel", first used in the journal of vice-admiral Wybrand van Warwijck, who visited the island with the Van Neck expedition in 1598. The description explained the meaning of the name as follows:

Landscape with Birds showing a Dodo in the lower right, by Roelant Savery, 1628

On their left hand was a little island which they named Heemskirk Island, and the bay it selve they called Warwick Bay... finding in this place great quantity of foules twice as bigge as swans, which they call Walghstocks or Wallowbirdes being very good meat. But finding an abundance of pigeons & popinnayes [parrots], they disdained any more to eat those great foules calling them Wallowbirds, that is to say lothsome or fulsome birdes.

The Dodo was referred to as "Dronte", meaning "swollen", by the crew of the Dutch ship Gelderland in 1602, a name still used in some languages. The Gelderland 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.

In his 18th century work Systema Naturae, Carl Linnaeus used the specific name cucullatus, meaning hooded, but combined it with the genus name Struthio, that of the ostrich. The name cucullatus was first used by Juan Eusebio Nieremberg in 1635, as Cygnus cucullatus, in reference to Carolus Clusius' 1605 depiction of a Dodo. Mathurin Jacques Brisson erected the genus name Raphus, referring to bustards, resulting in the current name. Linnaeus later coined the new name Didus ineptus, meaning "inept Dodo", but this has become a synonym of the earlier name because of nomenclatural priority.

Description

Indian Mughal miniature that shows a Dodo among Indian birds, possibly by Ustad Mansur 1625 CE.

No complete Dodo specimens exist to this day, 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 10.6–21.1 kg (23–47 lb) in the wild. A later estimate gives an average weight as low as 10.2 kg (22 lb). 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 plumaceous (downy) rather than vaned. The sternum was insufficient to support flight and the wings were very small. These features have been argued to indicate paedomorphosis.

Illustrations and written accounts of encounters with Dodos made between the Dodo's discovery and its extinction (1598–1662) are the primary evidence for its external appearance. According to most renditions, the Dodos 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.

Contemporary descriptions

Most descriptions are found in journals of Dutch ships that docked in Mauritius. 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.

An early account from Van Neck's journey in 1598 describes the bird thus:

Painting of a Dodo head by Cornelis Saftleven from 1638, which is the last original illustration of a Dodo

Blue parrots are very numerous there, as well as other birds; among which are a kind, conspicuous for their size, larger than our swans, with huge heads only half covered with skin as if clothed with a hood. These birds lack wings, in the place of which 3 or 4 blackish feathers protrude. The tail consists of a few soft incurved feathers, which are ash coloured. These we used to call 'Walghvogel', for the reason that the longer and oftener they were cooked, the less soft and more insipid eating they became. Nevertheless their belly and breast were of a pleasant flavour and easily masticated.

One of the most detailed descriptions is by Sir Thomas Herbert from 1634, who had visited Mauritius:

First here only and in Dygarrois [now Rodrigues, likely referring to the Solitaire] is generated the Dodo, which for shape and rareness may antagonize the Phoenix of Arabia: her body is round and fat, few weigh less than fifty pound. It is reputed more for wonder than for food, greasie stomackes may seeke after them, but to the delicate they are offensive and of no nourishment. Her visage darts forth melancholy, as sensible of Nature's injurie in framing so great a body to be guided with complementall wings, so small and impotent, that they serve only to prove her bird.

The halfe of her head is naked seeming couered with a fine vaile, her bill is crooked downwards, in midst is the trill [nostril], from which part to the end tis a light green, mixed with pale yellow tincture; her eyes are small and like to Diamonds, round and rowling; her clothing downy feathers, her train three small plumes, short and inproportionable, her legs suiting her body, her pounces sharpe, her appetite strong and greedy. Stones and iron are digested, which description will better be conceived in her representation.

Contemporary depictions

Compilation of the Gelderland sketches from 1601 of live and recently killed Dodos

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. Apart from the Gelderland sketches, it is also unknown whether any of the illustrations were drawn from life or from crudely stuffed specimens, affecting their reliability. Dodo specialist Julian Hume has argued that the nostrils of the living Dodo would have actually been slits, as seen in the Gelderland, Saftleven, Crocker Gallery and Mansur images. According to this claim, the gaping nostrils often seen in Dodo paintings indicate that dried specimens had been used as models for these.

The travel journal of the Dutch East India Company ship Gelderland (1601–1603), 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.

The famous "Edwards' Dodo", painted by Roelant Savery in 1626

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" 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 a Dodo.

Behaviour and ecology

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 enemes on Mauritius, it likely nested on the ground. It has been proposed that the Dodo was K-selected, meaning that it produced a low number of offspring, which required extensive parental care until they matured.

An account by François Cauche from 1651 is the only description of the egg and call:

I have seen in Mauritius birds bigger than a Swan, without feathers on the body, which is covered with a black down; the hinder part is round, the rump adorned with curled feathers as many in number as the bird is years old. In place of wings they have feathers like these last, black and curved, without webs. They have no tongues, the beak is large, curving a little downwards; their legs are long, scaly, with only three toes on each foot. It has a cry like a gosling, and is by no means so savoury to eat as the Flamingos and Ducks of which we have just spoken. They only lay one egg which is white, the size of a halfpenny roll, by the side of which they place a white stone the size of a hen's egg. They lay on grass which they collect, and make their nests in the forests; if one kills the young one, a grey stone is found in the gizzard. We call them Oiseaux de Nazaret. The fat is excellent to give ease to the muscles and nerves.

1601 map of a bay on Mauritius; the small D on the far right side marks where Dodos were found

However, this 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. 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.

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 other of the endemic species of Mauritius went 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 Réunion, but went extinct in both islands.

Diet

The Dodo likely subsisted on fallen fruits, nuts, seeds, bulbs and roots. A 1631 document, rediscovered in 1887 but now lost, is the only account of its diet, and also mentions that it used the beak for defense:

These Burgmeesters are superb and proud. They displayed themselves to us with stiff and stern faces, and wide-open mouths. Jaunty and audacious of gait, they would scarcely move a foot before us. Their war weapon was their mouth, with which they could bite fiercely; their food was fruit; they were not well feathered but abundantly covered with fat. Many of them were brought onboard to the delight of us all.

Dodo and its gizzard stone by Carolius Clusius from 1605, copied from an illustration in the journal of van Neck

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 shelfish, 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 and described it as follows:

About 1638, as I walked London streets, I saw the picture of a strange looking fowle hung out upon a clothe and myselfe with one or two more in company went in to see it. It was kept in a chamber, and was a great fowle somewhat bigger than the largest Turkey cock, and so legged and footed, but stouter and thicker and of more erect shape, coloured before like the breast of a young cock fesan, and on the back of a dunn or dearc colour. The keeper called it a Dodo, and in the ende of a chymney in the chamber there lay a heape of large pebble stones, whereof hee gave it many in our sight, some as big as nutmegs, and the keeper told us that she eats them (conducing to digestion), and though I remember not how far the keeper was questioned therein, yet I am confident that afterwards she cast them all again.

"Dodo tree" seeds at the Dodo exhibition at 'Naturalis'

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 tortoises to eat fruits and excrete their seeds, which would then be eaten by the parrots. This would correlate its extinction with that of the Dodo. Anodorhynchus macaws depended on the now extinct South American megafauna in the same way, but have since switched to domesticated cattle performing this function.

Relationship with humans

Copper engraving from 1601, showing Dutch activities on the shore of Mauritius, as well as the first published depiction of a Dodo (2), on the left

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. The Mascarene islands, Mauritius, Réunion and Rodrigues, were named after Pedro Mascarenhas, who was credited as having discovered them. It has been suggested that the Portuguese name for Mauritius, "Cerne (swan) Island", was a reference to the Dodos. The island was acquired by the Dutch in 1598, who renamed it 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 bird 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. According to Julian Hume, the following part of this journal may be the first mention of Dodos, referred to as penguins, a word not used for penguins at the time:

We also found large birds, with wings as large as of a pigeon, so that they could not fly and were named penguins by the Portuguese. These particular birds have a stomach so large that it could provide two men with a tasty meal and was actually the most delicious part 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.

Engraving showing the killing of Dodos (centre left, depicted as penguin-like) and other animals now extinct from Mauritius

The journal of Willem Van West-Zanen from 1602, unpublished until 1648, describes interaction with Dodos and mentions large numbers hunted for food:

They caught birds called by some Dod-aars by others Dronte. These were given the name Walghvogel during Van Neck's voyage, because even with long stewing they would hardly become tender, but stayed tough and hard with the exception of the breast and stomach which were extremely good...

The sailors brought 50 birds back to the Bruin-Vis, among them 24 or 25 Dod-aarsen, so big and heavy that scarcely two were consumed at meal time, and all that were remaining were flung into salt.

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:

For food the seamen hunt the flesh of feathered fowl,

They tap the palms, and round-rumped dodos they destroy,
The parrot's life they spare that he may peep and howl,

And thus his fellows to imprisonment decoy.

A painting from the collection of Emperor Rudolph II in Prague, by Jacob Hoefnagel, c. 1602

However, 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 "Edwads' Dodo". Two live specimens were seen by Peter Mundy in Surat, India, between 1628–1634, and one of them likely depicted in an Indian painting. One Dodo had been sent as far as Nagasaki, Japan in 1647, but it is unknown if it arrived.

In 1626, Emmanuel Altham visited Mauritius and sent a letter to his brother in England:

Adriaen van de Venne's 1626 depiction

You shall receive a jar of India ginger for my sister your wife, as also some beades for my cousins your daughters, and with all a strange fowle which I had at the island of Mauritius, called by ye portingalls a Dodo, which for rareness thereof I hope will be welcome to you.

Mauritius ye 18th June 1628.

Your most loving brother, Emmanuel Altham.

Whether the Dodo survived the journey is unknown, and the letter was destroyed by fire in the 19th century.

230 years before Darwin's theory of evolution, the appearance of the Dodo and the Red Rail led Peter Mundy to speculate:

Of these 2 sorts off fowl afforementionede, For oughtt wee yett know, Not any to bee Found out of this Iland, which lyeth aboutt 100 leagues From St. Lawrence. A question may bee demaunded how they should bee here and Not elcewhere, beeing soe Farer From other land and can Neither fly or swymme; whither by Mixture off kindes producing straunge and Monstrous formes, or the Nature of the Climate, ayer and earth in alltring the First shapes in long tyme, or how.

Extinction

Savery sketch of three Dodos from c. 1626, known as “the Crocker Art Gallery sketch”

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 humans. 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.

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.

Perseus and Andromeda with a Dodo (far right) and seashells, by Gillis d'Hondecoeter, 1627

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:

These animals on our coming up to them stared at us and remained quiet where they stand, not knowing whether they had wings to fly away or legs to run off, and suffering us to approach them as close as we pleased. Amongst these birds were those which in India they call Dod-aersen (being a kind of very big goose); these birds are unable to fly, and instead of wings, they merely have a few small pins, yet they can run very swiftly. We drove them together into one place in such a manner that we could catch them with our hands, and when we held one of them by its leg, and that upon this it made a great noise, the others all on a sudden came running as fast as they could to its assistance, and by which they were caught and made prisoners also.

The birds were caught on a small islet off Mauritius, 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.

Pieter van den Broecke's 1617 drawing of a Dodo, a one-horned sheep, and a Red Rail, a bird that was also sometimes referred to as "Dodo"

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. An example of this is the following description by the English traveller John Marshall from 1668:´

Here are also great plenty of Dodos or red hens which are larger a little than our English henns, have long beakes and no, or very little Tayles... They are good meate when roasted, tasting something like a pig, and their skin like pig skin when roosted, being hard.

Cheke therefore points to the 1662 description as the last credible observation, but even this 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. Until the Red Hen explanation was proposed, a description of "Dodos" from 1681 was long thought to be the last account. The Dutch left Mauritius in 1710, but by this time, the Dodo and most of the large, terrestrial vertebrates there had gone extinct.

Savery's 'The Paradise' from 1626 with a Dodo in the lower right corner

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 agency of man, in limiting the increase of the inferior animals, and in extirpating certain races, was perhaps never more strikingly exemplified than in the case of the Dodo. That a species so remarkable in its character should become extinct, within little more than two centuries, so that the fact of its existence at all has been doubted, is a circumstance which may well excite our surprise, and lead us to a consideration of similar changes which are still going on from the same cause.

Physical remains

17th century specimens

The leg and skull fragments in the National Museum of Prague

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 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:

Plaster casts of the Oxford head and the London foot made when they were still intact, Booth Museum of Natural History

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. The dissection resulted in the seminal 1848 monograph, The Dodo and Its Kindred, which attempted to separate Dodo myth from reality.

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 foot was 11% smaller than the Oxford foot, but is believed to have come from an adult specimen.

The skull in the Zoological Museum of Copenhagen

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 Denmark 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 Böhmisches 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.

Subfossil remains

In 1865, government schoolmaster at Mahébourg, 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:

After many fruitless visits to the spot... I resolved by sending some men into the centre of the marsh, where the water was about three feet deep and there, by feeling in the mud with their naked feet, they met with one entire tibia, a portion of another, and a tarso-metatarsus. The Dodo bones were imbedded only in the mud at the bottom of the water in the deepest part of the marsh... Encouraged by success, I employed several hands to search in the manner described, but I met with but few specimens of dodo bones till I thought of cutting away a mass of floating herbage nearly two feet in thickness, which covered the deepest part of the marsh. In the mud under this, I was rewarded by finding bones of many dodos.

The skeleton Richard Owen put together from bones found in the Mare aux Songes, at the Natural History Museum

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 Réunion Solitaire instead. The remaining bones not sold to Owen or Newton were auctioned off in London in 1866.

In 1889, Théodor Sauzier was commissioned to find more Dodo remains in the Mare aux Songes. He was successful, and also found remains of other extinct species.

Bones that were rediscovered at Grant Museum in 2011

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 no one knows 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.

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 dissociated subfossil remains. 26 museums worldwide have significant holdings of Dodo material, almost all found in the Mare aux Songes. An alleged Dodo egg is on display at the East London museum in South Africa, but genetic studies are underway to determine its authencity.

In October 2005, a part of the Mare aux Songes swamp was excavated by an international team of researchers. 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 while exploring a cave in Mauritius.

In 2011, a wooden box full of Dodo bones from the Edwardian era was discovered at the Grant Museum, during preparations for a move.

The white Dodo

Pieter Holsteyn's painting of a white Dodo

The supposed "Réunion Solitaire" or "White Dodo" of Réunion is now believed to have been an erroneous conjecture based on contemporary reports of the Réunion Sacred Ibis, combined with paintings by Pieter Withoos and Pieter Holsteyn from the 1600s of white Dodos that surfaced in the 19th century.

Willem Ysbrandtsz. Bontekoe, who visited Réunion around 1619, mentioned that it was inhabited by "Dod-eersen", though without mentioning colouration. When his journal was published in 1646, it was accompanied by a copy of Roelant Savery's "Crocker Art Gallery sketch". A white bird on Réunion was first described as follows in 1625 by Chief Officer J. Tatton:

There is store of land fowle both small and great, plenty of Doves, great Parrats, and such like; and a great fowle of the bignesse of a Turkie, very fat, and so short winged, that they cannot fly, being white, and in a manner tame: and so be all other fowles, as having not been troubled nor feared with shot. Our men did beat them down with sticks and stones. Ten men may take fowle enough to serve fortie men a day.

Pieter Withoos's painting of a white Dodo

In 1674 these white birds were again described by Sieur D. B. Dubois:

Solitaires. These birds are thus named because they always go alone. They are as big as a big goose and have white plumage, black at the extremity of the wings and of the tail. At the tail there are some feathers resembling those of the Ostrich. They have the neck long and the beak formed like that of the Woodcocks, but larger, and the legs and feet like those of Turkey-chicks. This bird betakes itself to running, only flying but very little. It is the best game on the Island.

19th century naturalists assumed these were descriptions of the white Dodo shown in the paintings, and a new species, Raphus solitarius, was erected by Baron Edmund de Sélys-Longchamps in 1848 for this bird. 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.

Others believed it was a species similar to the Rodrigues Solitaire, as it was also referred to as "Solitaire", or even that there were white species of both the Dodo and Solitaire on the island.

Roelant Savery painting from 1611 showing a whitish Dodo in the lower right corner

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 (1607–1611). 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 Réunion 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 Réunion 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 Réunion Solitaire, and this suggestion was published in 1995. The ibis was also reassigned to the genus Threskiornis, now combined with the specific epithet solitarius from the binomial Raphus solitarius. Birds of this genus are also white and black with slender beaks, which fits the old descriptions of the Réunion Solitaire. No fossil remains of Dodo-like birds have ever been found on the island.

Cultural significance

John Tenniel's 1865 illustration of Alice and the Dodo

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.

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.

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 Réunion 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

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:

The Dodo used to walk around,

And take the sun and air.
The sun yet warms his native ground –
The Dodo is not there!

The voice which used to squawk and squeak
Is now for ever dumb –
Yet may you see his bones and beak

All in the Mu-se-um.

See also

• Holocene extinction
• List of extinct birds

External links

• The Dodo – The merging of myth and reality: Half hour video interview with expert Julian Hume about the white Dodo
• Painting the dodo: Two minute video about Julian Hume's modern interpretation of Roelant Savery's Dodo
• Aves3D - Raphus cucullatus: Interactive 3D scans of various Dodo elements