Dodo

As no complete dodo specimens exist, its external appearance, such as plumage and colouration, is hard to determine.

The skull of the dodo differed much from those of other pigeons, especially in being more robust, the bill having a hooked tip, and in having a short cranium compared to the jaws. The upper bill was nearly twice as long as the cranium, which was short compared to those of its closest pigeon relatives. The openings of the bony nostrils were elongated along the length of the beak, and they contained no bony septum. The cranium (excluding the beak) was wider than it was long, and the

The dodo had about nineteen presynsacral vertebrae (those of the neck and

Dodo among Indian birds (left), by Ustad Mansur, c. 1625; perhaps the most accurate depiction of a live dodo, and dodo head (right) by Cornelis Saftleven from 1638, probably the last original depiction of the species

Most contemporary descriptions of the dodo are found in

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.[45]

One of the most detailed descriptions is by Herbert in A Relation of Some Yeares Travaille into Afrique and the Greater Asia from 1634:

First here only and in Dygarrois [Rodrigues] 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 thrill [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.^[46]

Contemporary depictions

The travel journal of the Dutch 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, and to a second, less refined artist.^[47] Apart from these sketches, it is unknown how many of the twenty or so 17th-century illustrations of the dodos were drawn from life or from stuffed specimens, which affects their reliability.^[23] Since dodos are otherwise only known from limited physical remains and descriptions, contemporary artworks are important to reconstruct their appearance in life. While there has been an effort since the mid-19th century to list all historical illustrations of dodos, previously unknown depictions continue to be discovered occasionally.^[48]

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.^[49] It has also been suggested that the images might show dodos with puffed feathers, as part of display behaviour.^[41] The Dutch painter Roelant Savery was the most prolific and influential illustrator of the dodo, having made at least twelve depictions, often showing it in the lower corners. A famous painting of his from 1626, now called Edwards's Dodo as it was once owned by the ornithologist George Edwards, has since become the standard image of a dodo. It is housed in the Natural History Museum, London. The image shows a particularly fat bird and is the source for many other dodo illustrations.^[50][51]

An Indian

The preferred habitat of the dodo is unknown, but old descriptions suggest that it inhabited the woods on the drier coastal areas of south and west Mauritius. This view is supported by the fact that the

The mayors are superb and proud. They presented themselves with an unyielding, stern face and wide open mouth, very jaunty and audacious of gait. They did not want to budge before us; their war weapon was the mouth, with which they could bite fiercely. Their food was raw fruit; they were not dressed very well, but were rich and fat, therefore we brought many of them on board, to the contentment of us all.[66]

In addition to fallen fruits, the dodo probably subsisted on nuts, seeds, bulbs, and roots.

Several contemporary sources state that the dodo used Gastroliths (gizzard stones) to aid digestion. The English writer Sir Hamon L'Estrange witnessed a live bird in London 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.^[69]

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 to produce 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 could produce for their young during early growth.^[70]

In 1973, the

The Brazilian ornithologist Carlos Yamashita suggested in 1997 that the broad-billed parrot may have depended on dodos and Cylindraspis tortoises to eat palm fruits and excrete their seeds, which became food for the parrots. Anodorhynchus macaws depended on now-extinct South American megafauna in the same way, but now rely on domesticated cattle for this service.^[75]

Reproduction and development

As it was flightless and terrestrial and there were no

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.^[6]

growth series

Diagram showing life history events of a dodo based on histology

and accounts

Cauche's account is problematic, since it also mentions that the bird he was describing had three toes and no tongue, unlike dodos. This led some to believe that Cauche was describing a new species of dodo ("Didus nazarenus"). The description was most probably mingled with that of a cassowary, and Cauche's writings have other inconsistencies.^[77] A mention of a "young ostrich" taken on board a ship in 1617 is the only other reference to a possible juvenile dodo.^[78] An egg claimed to be that of a dodo is stored in the East London Museum in South Africa. It was donated by the South African museum official Marjorie Courtenay-Latimer, whose great aunt had received it from a captain who claimed to have found it in a swamp on Mauritius. In 2010, the curator of the museum proposed using genetic studies to determine its authenticity.^[79] It may instead be an aberrant ostrich egg.^[32]

Because of the possible single-egg clutch and the bird's large size, it has been proposed that the dodo was

Mauritius had previously been visited by

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.[87]

Some early travellers found dodo meat unsavoury, and preferred to eat parrots and pigeons; others described it as tough, but good. Some hunted dodos only for their gizzards, as this was considered the most delicious part of the bird. Dodos were easy to catch, but hunters had to be careful not to be bitten by their powerful beaks.^[88]

The appearance of the dodo and the red rail led Peter Mundy to speculate, 230 years before Charles Darwin's theory of evolution:

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.^[27]

Dodos transported abroad

Emperor Rudolph II in Prague, by Jacob Hoefnagel, early 1600s

Adriaen van de Venne

's 1626 depiction of a dodo he claimed to have seen

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. Based on a combination of contemporary accounts, paintings, and specimens, Julian Hume has inferred that at least eleven transported dodos reached their destinations alive.[89]

Hamon L'Estrange's description of a dodo that he saw in London in 1638 is the only account that specifically mentions a live specimen in Europe. In 1626 Adriaen van de Venne drew a dodo that he claimed to have seen in Amsterdam, but he did not mention if it was alive, and his depiction is reminiscent of Savery's Edwards's Dodo. Two live specimens were seen by Peter Mundy in Surat, India, between 1628 and 1634, one of which may have been the individual painted by Mansur around 1625.^[23] In 1628, Emmanuel Altham visited Mauritius and sent a letter to his brother in England:

Right wo and lovinge brother, we were ordered by ye said councell to go to an island called Mauritius, lying in 20d. of south latt., where we arrived ye 28th of May; this island having many goates, hogs and cowes upon it, and very strange fowles, called by ye portingalls Dodo, which for the rareness of the same, the like being not in ye world but here, I have sent you one by Mr. Perce, who did arrive with the ship William at this island ye 10th of June. [In the margin of the letter] Of Mr. Perce you shall receive a jarr of ginger for my sister, some beades for my cousins your daughters, and a bird called a Dodo, if it live.^[90]

Savery paintings featuring dodos in various corners (one with a lobster's body, lower right), painted in Europe approximately between 1625 and 1629

Whether the dodo survived the journey is unknown, and the letter was destroyed by fire in the 19th century.^[91] The earliest known picture of a dodo specimen in Europe is from a c. 1610 collection of paintings depicting animals in the royal menagerie of

One dodo was reportedly sent as far as Nagasaki, Japan, in 1647, but it was long unknown whether it arrived.^[75] Contemporary documents first published in 2014 proved the story, and showed that it had arrived alive. It was meant as a gift, and, despite its rarity, was considered of equal value to a white deer and a bezoar stone. It is the last recorded live dodo in captivity.^[92]

Extinction

Like many animals that evolved in isolation from significant predators, the dodo was entirely

It has been suggested that the dodo may already have been rare or localised before the arrival of humans on Mauritius, since it would have been unlikely to become extinct so rapidly if it had occupied all the remote areas of the island.[60] 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.^[96] Yet the fact that the dodo survived hundreds of years of volcanic activity and climatic changes shows the bird was resilient within its ecosystem.^[62]

Some controversy surrounds the date of its extinction. The last widely accepted record of a 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 Amber Island:

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.^[97]

The dodos on this islet may not necessarily have been the last members of the species.

Pieter van den Broecke's 1617 drawing of a dodo, a one-horned sheep, and a red rail; after the dodo became extinct, visitors may have confused it with the red rail (1907 restoration of that bird at right by Frederick William Frohawk)

The British ornithologist Alfred Newton suggested in 1868 that the name of the dodo was transferred to the red rail after the former had gone extinct.^[100] Cheke also pointed out that some descriptions after 1662 use the names "Dodo" and "Dodaers" when referring to the red rail, indicating that they had been transferred to it.^[101] He therefore pointed to the 1662 description as the last credible observation. A 1668 account by English traveller John Marshall, who used the names "Dodo" and "Red Hen" interchangeably for the red rail, mentioned that the meat was "hard", which echoes the description of the meat in the 1681 account.^[102] Even the 1662 account has been questioned by the writer Errol Fuller, as the reaction to distress cries matches what was described for the red rail.^[103] Until this explanation was proposed, a description of "dodos" from 1681 was thought to be the last account, and that date still has proponents.^[104]

Cheke stated in 2014 that then recently accessible Dutch manuscripts indicate that no dodos were seen by settlers in 1664–1674.

It is unlikely the issue will ever be resolved, unless late reports mentioning the name alongside a physical description are rediscovered.[95] The IUCN Red List accepts Cheke's rationale for choosing the 1662 date, taking all subsequent reports to refer to red rails. In any case, the dodo was probably extinct by 1700, about a century after its discovery in 1598.^[2][102] The Dutch left Mauritius in 1710, but by then the dodo and most of the large terrestrial vertebrates there had become extinct.^[38]

Even though the rareness of the dodo was reported already in the 17th century, its extinction was not recognised until the 19th century. This was partly because, for religious reasons, extinction was not believed possible until later proved so by Georges Cuvier, and partly because many scientists doubted that 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 in 1833,^[107] and has since been referred to as an "icon" of extinction.^[108]

Physical remains

17th-century specimens