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 island of Mauritius, east of Madagascar in the Indian Ocean. Genetically grouped within pigeons and doves, its closest genetic relative was the (likewise extinct) Rodrigues Solitaire, the two forming the subfamily Raphinae. The closest living relative of the Dodo is the Nicobar Pigeon. A white Dodo was at one time incorrectly thought to have existed on the nearby island of Réunion.

The Dodo was about one metre (about 3.3 ft) tall, and may have weighed approximately 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 because only a few sketches are known to have been drawn from live specimens, its exact appearance remains a mystery. Similarly, little about its habitat and behaviour is known with certainty. It has been depicted with brownish-grey plumage, yellow feet, a tuft of tail feathers, a grey-coloured, naked head, and a black, yellow, and green beak. It used gizzard stones to help digest its food, which is thought to have included fruits, and its main habitat is believed to have been the woods in the drier coastal areas of Mauritius. It is presumed that the Dodo became flightless because of the ready availability of abundant food sources and a relative absence of predators on Mauritius.

The first recorded mention of the Dodo was by Dutch sailors in 1598. In the following years, the bird was preyed upon by hungry sailors, their domesticated animals, and other invasive species introduced during that time. The last credible recorded sighting of a Dodo was in 1662. Its extinction was not immediately noticed, and some considered it to be a mythological creature. In the 19th century, research was conducted on a small quantity of remains of four specimens that had been brought to Europe in the early 17th century. Since then, a large amount of subfossil material has been collected from Mauritius, mostly from the Mare aux Songes swamp. The extinction of the Dodo within only about a century of its discovery called attention to the problem of human involvement in the disappearance of entire species, which until that time had not been recognised.

The Dodo achieved widespread recognition from its role in the story of Alice in Wonderland, and it has since become a fixture in popular culture, often as a symbol of extinction and obsolescence. It is frequently used as a mascot on Mauritius.

Taxonomy and evolution

1848 lithograph of the Oxford specimen skull

The Dodo has been variously declared a small ostrich, a rail, an albatross, and a vulture. In 1842, Johannes Theodor Reinhardt proposed that Dodos were ground pigeons, based on studies of a Dodo skull he had discovered in the royal Danish collection at Copenhagen. This view was met with ridicule, but was later supported by Hugh Edwin Strickland and Alexander Gordon Melville in their 1848 monograph, The Dodo and Its Kindred, which attempted to separate myth from reality. After dissecting the preserved head and foot of the specimen at Oxford Museum, comparing it with the few remains of the likewise extinct Rodrigues Solitaire (Pezophaps solitaria) then available, they found that the two were closely related. Strickland stated that although not identical, these birds shared many distinguishing features in the leg bones otherwise only known in pigeons.

The Dodo was anatomically similar to pigeons in many features. Strickland pointed to the keratinous portion of the beak being very short, with a long, slender, naked basal part. Other pigeons also have bare skin around their eyes, almost reaching their beak, as in Dodos. The forehead was high in relation to the beak, and the nostril was located low on the middle of the beak, surrounded by skin, a combination of features shared only with pigeons. The legs of the Dodo were generally more similar to terrestrial pigeons than other birds, both in scale and in skeletal features. Depictions of the large crop hinted at a relation, as this feature is more developed in pigeons than other birds. Pigeons generally have very small clutch sizes, which is similar to the single egg said to have been laid by the Dodo. Like pigeons, the Dodo lacked the vomer and septum of the nostrils, and it shared details in the mandible, the zygomatic bone, the palate and the hallux. The Dodo differed from other pigeons mainly in the small size of the wings, and the large size of the beak in proportion to the rest of the cranium.

Foot of the Oxford specimen, used as source for the genetic study

For many years the Dodo and the Rodrigues Solitaire were placed in a family of their own, the Raphidae (formerly Dididae). This was because their exact relationships to other pigeons were unresolved. They were also placed in a monotypic family each (Raphidae and Pezophapidae, respectively) due to the suggestion that they had evolved their similar features independently. Osteological and molecular data has since led to the dissolution of the family Raphidae, and the Dodo and Solitaire are now placed in their own subfamily, Raphinae, grouped within the Columbidae.

Comparison of mitochondrial cytochrome b and 12S rRNA sequences isolated from the tarsal of a Dodo and a femur of a Rodrigues Solitaire confirmed their close relationship, and their placement within the Columbidae. The genetic evidence was interpreted to show that the Southeast Asian Nicobar Pigeon is their closest living relative, followed by the Crowned Pigeons of New Guinea, and the superficially Dodo-like Tooth-billed Pigeon from Samoa. The generic name of the latter is Didunculus ("little Dodo")—this bird was also referred to as "Dodlet" by Richard Owen. The following cladogram, from Shapiro and colleagues (2002), shows the Dodo's closest relationships within Columbidae, a clade consisting of generally ground-dwelling island endemics.

	Goura victoria (Victoria Crowned Pigeon) Caloenas nicobarica (Nicobar Pigeon) Pezophaps solitaria (Rodrigues Solitaire) Raphus cucullatus (Dodo) Didunculus strigirostris (Tooth-billed Pigeon)

Sketch of the Oxford specimen head prior to dissection

A similar cladogram was published in 2007, inverting the placement of Goura and Dicunculus, as well as including the Pheasant Pigeon and the Thick-billed Ground Pigeon at the base of the clade. The 2002 study indicated that the ancestors of the Solitaire and the Dodo diverged around the Paleogene-Neogene boundary. The Mascarene Islands (Mauritius, Réunion, and Rodrigues), are of volcanic origin and are less than 10 million years old. Therefore, the ancestors of both birds likely remained capable of flight for a considerable time after the separation of their lineage. The lack of mammalian herbivores competing for resources on these islands allowed the Solitaire and the Dodo to attain very large sizes.

Throughout the 19th century, several species were classified as 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 Abraham Dee Bartlett to name a new species, Didus nazarenus, in 1852. Based on Solitaire remains, it is now a synonym of that species. Crude drawings of the Red Rail of Mauritius were also misinterpreted as Dodo species, Didus broeckii and Didus herberti. Another large, flightless pigeon, the Viti Levu Giant Pigeon, was described in 2001 from subfossil material from Fiji. It was only slightly smaller than the Dodo and the Solitaire, and it is also thought to have been related to the Crowned Pigeons.

Etymology

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

One of the original names for the Dodo was "walghvogel" (Dutch), first used in the journal of Vice Admiral Wybrand van Warwijck, who visited the island during the Second Dutch Expedition to Indonesia in 1598. His description, translated into English in 1599, explained the meaning of the name as follows:

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.

Labelled sketch from 1634 by Sir Thomas Herbert, showing a Broad-billed Parrot, a Red Rail ("Cacato" and "Hen"), and a Dodo

Another account from that voyage, perhaps the first to mention the bird, states that the Portuguese referred to them as "penguins". The meaning may not have been derived from penguin, but from "pinion", a reference to the small wings. The crew of the Dutch ship Gelderland referred to the bird as "Dronte" (meaning "swollen") in 1602, a name that is still used in some languages. This crew also called them "griff-eendt" and "kermisgans", in reference to fowl fattened for the Kermesse festival in Amsterdam, which was held the day after they anchored on Mauritius.

The etymology of the word Dodo is unclear. Some ascribe it to the Dutch word dodoor for "sluggard", but it is more likely related to Dodaars, which means either "fat-arse" or "knot-arse", referring to the knot of feathers on the hind end. The first record of the word Dodaars is in captain Willem Van West-Zanen's journal in 1602. The English form "Dodar" was used until the 18th century. Sir Thomas Herbert was the first to use the word Dodo in print in his 1634 travelogue, and claimed it was referred to as such by the Portuguese, who had visited the island in 1507. Another Englishman, Emmanuel Altham, had used the word in a 1628 letter, wherein he also claimed the origin was Portuguese. As far as is known, the Portuguese did not use the word. Nevertheless, some sources still state that the word "Dodo" derives from the Portuguese word doudo (currently doido), meaning "fool" or "crazy". David Quammen suggested that Dodo was an onomatopoeic approximation of the bird's call, a two-note pigeon-like sound resembling "doo-doo".

The Latin name cucullatus ("hooded") was first used by Juan Eusebio Nieremberg in 1635 as Cygnus cucullatus, in reference to Carolus Clusius' 1605 depiction of a Dodo. In his 18th century classic work Systema Naturae, Carl Linnaeus used cucullatus as the specific name, but combined it with the genus name Struthio (ostrich). Mathurin Jacques Brisson coined the genus name Raphus (referring to bustards) in 1760, which resulted in the current name Raphus cucullatus. In 1766, Linnaeus coined the new binomial Didus ineptus (meaning "inept Dodo"). This has become a synonym of the earlier name because of nomenclatural priority.

Description

Dodo among Indian birds, by Ustad Mansur, ca 1625

No complete Dodo specimens exist, making its external appearance, such as plumage and colouration, hard to determine. Subfossil remains and remnants of the birds that were brought to Europe in the 17th century show that they were very large birds, one metre (3.3 ft) tall, and possibly weighing up to 23 kilograms (51 lb). 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 depended on season, and that individuals were fat during cool seasons, but slim during hot. The bird was sexually dimorphic. Males were larger and had proportionally longer beaks. The length of the beak was up to 23 centimetres (9.1 in), and it had a hooked point. A study of the few remaining feathers on the Oxford specimen head showed that they were pennaceous rather than plumaceous (downy) and most similar to those of other pigeons.

Many of the skeletal features of the Dodo and the Rodrigues Solitaire, its closest relative, that distinguish them from pigeons have been attributed to their flightlessness. The pelvic elements were thicker than those of flighted pigeons to support the higher weight, and the pectoral region and the small wings were paedomorphic, meaning they were underdeveloped and retained juvenile features. The skull, trunk and pelvic limbs were peramorphic, meaning that they changed considerably with age. The Dodo shared several other traits with the Rodrigues Solitaire, such as features in the skull, pelvis, and sternum, as well as their large size. It differed in other aspects, such as being more robust and shorter than the Solitaire, having a larger skull and beak, a rounded skull roof, and smaller orbits. The Dodo's neck and legs were proportionally shorter, and it did not possess an equivalent to the knob present on the Solitaire's wrists.

Illustrations and written accounts of encounters with the Dodo between its discovery and its extinction (1598–1662) are the primary evidence for its external appearance. According to most representations, the Dodo had greyish or brownish plumage, with lighter primary feathers and a tuft of curly light feathers high on its rear end. The head was grey and naked, the beak green, black and yellow, and the legs were stout and yellowish, with black claws.

Contemporary descriptions

Most contemporary descriptions of the Dodo are found in journals of Dutch East India Company ships that docked in Mauritius when the Dutch Empire ruled the island. These records were used as guides for future voyages. Few contemporary accounts are reliable, as many seem based on earlier accounts, and none were written by scientists.

One of the earliest accounts, from van Warwijck's 1598 journal, describes the bird thus:

Painting of a Dodo head by Cornelis Saftleven from 1638, the last original depiction

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 in A Relation of Some Yeares Travaille into Afrique and the Greater Asia from 1634:

First here only and in Dygarrois [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

Some twenty 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, around the time reports mentioning Dodos became rarer. Differences in the depictions 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 signal age or sex, or if they even reflect reality. Dodo specialist Julian Hume argued that the nostrils of the living Dodo would have been slits, as seen in the Gelderland, Cornelis Saftleven, Crocker Art Gallery and Ustad Mansur images. According to this claim, the gaping nostrils often seen in paintings indicate that taxidermy specimens were used as models.

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, as well as a second, less refined artist. Apart from these sketches, it is unknown how many of the other illustrations were drawn from life or from stuffed specimens, affecting their reliability.

The 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. Dutch painter Roelant Savery was the most prolific and influential illustrator of the Dodo, having made at least ten depictions, often showing it in the lower corners. A famous painting of his from 1626 now called "Edwards' Dodo" (as it was once owned by George Edwards) in the British Museum has since become the standard image of a Dodo. The image shows a particularly fat bird, and is the source for many other Dodo illustrations.

An Indian Mughal painting rediscovered in St. Petersburg in the 1950s shows a Dodo along with native Indian birds. It depicts a slimmer, brownish bird, and its discoverer A. Iwanow and Dodo specialist Julian Hume regard it as one of the most accurate depictions of the living Dodo; the surrounding birds are clearly identifiable and depicted with appropriate colouring. It is believed to be from the 17th century and has been attributed to artist Ustad Mansur. The bird depicted likely lived in the menagerie of Mughal Emperor Jahangir, located in Surat, where English traveller Peter Mundy also claimed to have seen Dodos.

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 enemies on Mauritius, it likely nested on the ground.

The 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

Due to the single-egg clutch, and its large size, it has been proposed that the Dodo was K-selected, meaning that it produced a low number of altricial offspring, which required parental care until they matured. Cauche's account is problematic, since it also mentions the bird had three toes and no tongue, unlike Dodos, which led some to believe it referred to a new species of Dodo ("Didus nazarenus"). The description was most likely mingled with that of a cassowary, and Cauche's writings have other inconsistencies. A mention of a "young ostrich" taken on board a ship in 1617 is the only other reference to a possible juvenile Dodo. Some evidence indicates that the bird may have had a protracted development period, including the large size, and the fact that tropical and frugivorous birds have slower growth rates.

The preferred habitat of the Dodo is unknown, but old descriptions suggest it inhabited the woods on the drier coastal areas of south and west Mauritius. The Mare aux Songes swamp, where most Dodo remains have been found, is close to the sea in south-eastern Mauritius, which also supports this notion. Such a limited distribution across the island would likely have contributed to its extinction. A 1601 map from the Gelderland journal 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. Subfossil bones have also been found inside caves in highland areas, indicating that it once occurred on mountains. Work at the Maure aux Songes swamp has shown that its habitat was dominated by tambalacoque and Pandanus trees, as well as endemic palms.

Many endemic species of Mauritius became extinct after the arrival of man, so the ecosystem of the island is heavily damaged, and hard to reconstruct. Before humans arrived, Mauritius was entirely covered in forests, but very little remains today due to deforestation. The surviving endemic fauna is still seriously threatened. The Dodo lived alongside other recently extinct Mauritian birds such as the flightless Red Rail, the Broad-billed Parrot, Thirioux's Grey Parrot, the Mauritius Blue Pigeon, the Mauritius Owl, the Mascarene Coot, the Mauritian Shelduck, the Mauritian Duck, and the Mauritius Night Heron. Extinct Mauritian reptiles include the Saddle-backed Mauritius giant tortoise, the Domed Mauritius giant tortoise, the Mauritian Giant Skink, and the Round Island Burrowing Boa. The Small Mauritian Flying Fox and the snail Tropidophora carinata lived on Mauritius and Réunion, but vanished from both islands.

Diet

Savery sketch of three Dodos from c. 1626, Crocker Art Gallery

A 1631 Dutch document, rediscovered in 1887 but now lost, is the only account of its diet, and also mentions that it used the beak for defence:

These mayors 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.

In addition to fallen fruits, the Dodo likely subsisted on nuts, seeds, bulbs and roots. Anthonie Oudemans suggested that as Mauritius has marked dry and wet seasons, the Dodo probably fattened itself on ripe fruits at the end of the wet season to survive the dry season, when food was scarce; contemporary reports describe the bird's "greedy" appetite. France Staub suggested they mainly fed on palm fruits, and attempted to correlate the fat-cycle of the Dodo with the fruiting regime of the palms. It has also been suggested that the Dodo might have eaten crabs and shellfish, like their relatives the Crowned Pigeons. Its feeding habits must have been versatile, since captive specimens were likely given a wide range of food on the long ship journeys.

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

Several contemporary sources state that the Dodo used gizzard stones to aid digestion. The English writer Sir Hamon L'Estrange witnessed a live bird in London and described it as follows:

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.

It is not known how the young were fed, but related pigeons provide crop milk. Contemporary depictions show a large crop, which was probably used to add space for food storage, and for producing crop milk. It has been suggested that the maximum size attained by the Dodo and the Solitaire was limited by the amount of crop milk they were able to produce for their young during early growth.

"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, to which it was endemic. There were supposedly only 13 specimens left, all estimated to be about 300 years old. 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 coextinct because of the disappearance of the Dodo. Temple overlooked reports from the 1940s, which found that tambalacoque seeds germinated, albeit very rarely, without being abraded during digestion. Others have contested his hypothesis and suggested the decline of the tree was exaggerated, or seeds were also distributed by other extinct animals such as Cylindraspis tortoises, fruit bats or the Broad-billed Parrot. According to Wendy Strahm and Anthony Cheke, two experts in Mascarene ecology, the tree while rare has germinated since the demise of the Dodo and numbers several hundred, not 13 as claimed by Temple, hence discrediting Temple's view as to the Dodo and the tree's sole survival relationship.

It has also been suggested that the Broad-Billed Parrot may have depended on Dodos and Cylindraspis tortoises to eat palm fruits and excrete their seeds, which became food for the parrots. Anodorhynchus macaws depended on the now extinct South American megafauna in the same way, but now rely on domesticated cattle for this service.

Relationship with humans

1601 engraving 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 in the Middle Ages and Portuguese ships between 1507 and 1513, they did not settle on the island, and no known records of Dodo encounters survive. However, the Portuguese name for Mauritius, "Cerne (swan) Island", may have been a reference to the Dodos. The Dutch Empire acquired the island in 1598, renaming it after Maurice of Nassau, and thereafter it was used for the provisioning of Dutch East India Company trade vessels. The earliest known descriptions of the Dodo were made by Dutch travellers during the Second Dutch Expedition to Indonesia, led by Admiral Jacob van Neck in 1598. They appear in reports published in 1601, which also contain the first published illustration of the bird. Since the first sailors who visited Mauritius had been at sea for a long time, their interest in these large birds was mainly culinary. Although many later writings describe the meat as unsavoury, early journals state that it was tough but good, though not as delectable as the abundant pigeons.

The journal of Willem Van West-Zanen of the ship 'Bruin-Vis', written in 1602 and published in 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.

1648 engraving showing the killing of Dodos (centre left, depicted as penguin-like) and other animals now extinct from Mauritius

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.

230 years before Charles 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.

Dodos transported abroad

Painting of a specimen in the collection of Emperor Rudolph II in Prague, by Jacob Hoefnagel, c. 1602

The Dodo was found interesting enough that living specimens were sent to Europe and the East. The number of transported Dodos that reached their destinations alive is uncertain, and it is unknown how they relate to contemporary depictions and the few non-fossil remains in European museums. Hamon L'Estrange's description of a Dodo he saw in London in 1638 is the only account that specifically mentions a live specimen in Europe. Adriaen van de Venne drew a Dodo in 1626 he claimed to have seen in Amsterdam, but did not mention if it was alive, and his depiction is remiscent of Savery's "Edwards' Dodo". Two live specimens were seen by Peter Mundy in Surat, India, between 1628 and 1634, one of which was perhaps the individual painted by Ustad Mansur around 1625. One Dodo had been sent as far as Nagasaki, Japan in 1647, but it is unknown if it arrived.

Adriaen van de Venne's 1626 depiction of a Dodo

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.

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

That whole stuffed Dodos were present in Europe indicates they had been brought alive and died there; it is unlikely that taxidermists were on board the visiting ships, and spirits were not yet used to pickle biological specimens. Most tropical specimens were preserved as dried heads and feet. Based on a combination of contemporary accounts, paintings and specimens, Hume inferred that at least eleven transported Dodos reached their destinations alive.

Extinction

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

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 sailors. Although some scattered reports describe mass killings of Dodos for ship provisions, archaeological investigations have found scant evidence of human predation. Bones of at least two Dodos were found in caves at Baie du Cap that sheltered fugitive slaves and convicts in the 17th century, and were not easily accessible to Dodos due to the high, broken terrain. The human population on Mauritius (an area of 1,860 km²) never exceeded 50 people in the 17th century, but they introduced other animals, including dogs, pigs, cats, rats, and crab-eating Macaques, which plundered Dodo nests, and competed for the limited food resources. At the same time, humans destroyed the Dodo's habitat forests; the impact these introduced animals, especially the pigs and macaques, had on the Dodo population is currently considered as more severe than the impact of hunting. Rats were perhaps not much of a threat to the nests, since Dodos would have been used to dealing with local land crabs.

It has been suggested that the Dodo may already have been rare or localised before the arrival of humans on Mauritius, since it is unlikely it would have become extinct so rapidly if it had occupied all remote areas of Mauritius. A 2005 expedition found subfossil remains of Dodos and other animals killed by a flash flood. Such mass mortalities would have further jeopardised a species already in danger of becoming extinct.

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

Some controversy surrounds their extinction date. The last widely accepted Dodo sighting is the 1662 report by shipwrecked mariner Volkert Evertsz of the Dutch ship Arnhem, who described birds caught on a small islet off Mauritius, now suggested to be Ile d'Ambre:

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.

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"

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 quite rare by the 1660s, and thus a disputed report from 1674 by an escaped slave cannot be dismissed out-of-hand.

Anthony Cheke pointed out that by this time, some descriptions specifically use the names "Dodo" and "Dodaers" when referring to the Red Rail, indicating that they had been transferred to it after the disappearance of the actual Dodos. Cheke therefore points to the 1662 description as the last credible observation. Until this explanation was proposed, a description of "Dodos" from 1681 was thought to be the last account. A 1668 account by English traveller John Marshall, who used the names "Dodo" and "Red Hen" interchangeably for the Red Rail, mentioned the meat was "hard", which mirrors the description of the meat in the 1681 account. Even the 1662 account has been questioned by Errol Fuller, as the reaction to distress cries matches what was described for the Red Rail. The IUCN Red List accepts Cheke's rationale for choosing the 1662 date, with all subsequent reports referring to Red Rails, and the Dodo was likely extinct by 1700 in any case, about a century after the discovery of the species in 1598. The Dutch left Mauritius in 1710, but by this time the Dodo and most of the large, terrestrial vertebrates there had gone extinct.

Even though the rareness of the Dodo was reported already in the 17th century, its extinction was not realised until the 19th century. This was partially because, for religious reasons, extinction was not believed possible until later proven by Georges Cuvier, and also because many scientists doubted the Dodo had ever existed. It seemed altogether too strange a creature, and many believed it a myth. The bird was first used as an example of human induced extinction in Penny Magazine, 1833.

Physical remains

17th century specimens

Leg and skull fragments in the National Museum of Prague

The only extant 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-19th century. Other stuffed Dodos were mentioned in old museum inventories, but none of these are known to have survived.

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

The only known soft tissue remains, the Oxford head (specimen OUM 11605) and foot, belonged to the last known stuffed Dodo, which was first mentioned as part of the Tradescant collection in 1656, and moved to the Ashmolean Museum in 1659. It has been suggested that this might be the remains of the bird Hamon L'Estrange saw in London. It is a commonly held belief that the museum burned the stuffed Dodo because of severe decay around 1755, only saving the head and leg, due to status eight of the museum, which states "That as any particular grows old and perishing the keeper may remove it into one of the closets or other repository; and some other to be substituted." Its deliberate destruction is now believed a myth. It was removed from exhibition to preserve what remained of the specimen. This remaining soft tissue has since degraded further; the head was dissected by Strickland and Melville, separating the skin from the skull in two halves. The foot is in a skeletal state, with only scraps of skin and tendons. Only very few feathers remain on the head. It has been interpreted as a female, as the foot is 11% smaller and more gracile than that of the London specimen, yet appears to be fully grown. By 1896 it was mentioned as being without its integuments, and it is thought only the bones remain today. Its present whereabouts are unknown.

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 Danish forces in 1702, the museum collection was assimilated into the Royal Danish collection, and the skull was rediscovered by J. T. Reinhardt in 1840. It is 13 mm shorter than the Oxford skull, and may have belonged to a female. It is thought to have been mummified originally, but is now devoid of soft tissue. Its sclerotic ring is also preserved.

The front part of a skull (specimen NMP P6V-004389) and some leg bones in the National Museum of Prague were found in 1850 among the remains of the 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, but is now lost. It belonged to the Dutch professor Pieter Pauw, and its provenance is unknown. It has been suggested it was collected during the Van Neck voyage.

Subfossil specimens

Until 1860, the only Dodo remains known were the four incomplete 17th century specimens. Philip Burnard Ayres found the first sub fossil bones, which were sent to Richard Owen at the British Museum, who did not publish the findings. In 1863, Owen requested that Mauritian Bishop Vincent Ryan spread word that he should be informed if any Dodo bones were found. In 1865, government schoolmaster at Mahébourg, George Clark, finally found an abundance of sub fossil Dodo bones in the swamp of Mare aux Songes in Southern Mauritius, after searching for thirty years, inspired by Strickland & Melville's monograph. In 1866, Clark explained his procedure to The Ibis, an ornithology journal; he had sent his coolies to wade through the centre of the swamp, while feeling for bones with their feet. Initially, they only found a few bones, until they cut away herbage that covered the deepest part of the swamp, where they found an abundance of fossils. Remains of over 300 Dodos were found in the swamp, but only very few skull and wing bones among them, possibly because the upper bodies were 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.

The skeleton Richard Owen assembled from bones found in the Mare aux Songes

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, which led to rivalry between the two. Owen described the Dodo bones in Memoir on the Dodo in October 1866, but 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 or donated to museums.

In 1889, Théodor Sauzier was commissioned to explore the "historical souvenirs" of Mauritius and find more Dodo remains in the Mare aux Songes. He was successful, and also found remains of other extinct species. A barber named Louis Etienne Thirioux also found many Dodo remains around 1900, including the first remains of a juvenile, which are now lost. He also found the first articulated specimen, but it is unknown exactly where he made his finds, except that some were found in a cave. It has been suggested the cave was located on the slopes of the Le Pouce Mountain.

Replica of an alleged Dodo egg in a reconstructed nest, East London Museum

Worldwide, 26 museums have significant holdings of Dodo material, almost all found in the Mare aux Songes. The Natural History Museum, American Museum of Natural History, Cambridge University Museum of Zoology, the Senckenberg Museum and others, have almost complete specimens assembled from these dissociated subfossil remains. An alleged Dodo egg is stored at the East London Museum in South Africa. It was donated by Marjorie Courtenay-Latimer, whose great aunt had received it from a captain who claimed he had found it in a swamp on Mauritius. Genetic studies are under way to determine its authenticity. It may instead be an aberrant ostrich egg.

In October 2005, after a hundred years of neglect, a part of the Mare aux Songes swamp was excavated by an international team of researchers. To prevent malaria, the British had covered the swamp with hard core during their rule over Mauritius, which had to be removed. Many remains were found, including bones of Dodos 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, 4,000 years ago. Subsequent excavations suggested that Dodos, along with other animals, became mired in the Mare aux Songes while trying to reach water during a long period of severe drought about 4,200 years ago.

In June 2007, adventurers discovered the most complete and best-preserved Dodo skeleton ever found while exploring a cave in Mauritius. The specimen was nicknamed "Fred" after the finder. In 2011, a wooden box containing Dodo bones from the Edwardian era was rediscovered at the Grant Museum, during preparations for a move.

The White Dodo

Pieter Holsteyn's painting of a white Dodo

The supposed "White Dodo" (or "Solitaire") of Réunion is now believed to have been an erroneous conjecture based on contemporary reports of the Réunion Sacred Ibis, combined with 17th century paintings of white Dodos by Pieter Withoos and Pieter Holsteyn that surfaced in the 19th century.

The confusion began when Willem Ysbrandtsz. Bontekoe, who visited Réunion around 1619, mentioned birds with Dodo like features and with a name similar to "Dodaars" in his journal, though without mentioning colouration:

There were also some Dod-eersen which had small wings but could not fly. They were so fat that they could scarcely walk, for when they did so their bellies dragged along the ground.

When the journal was published in 1646, it was accompanied by a copy of Roelant Savery's "Crocker Art Gallery sketch".

Pieter Withoos's painting of a white Dodo among various birds

A white, stocky, and flightless bird was first mentioned as part of the Réunion fauna by Chief Officer J. Tatton in 1625. In 1674, such 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.

Baron Edmund de Sélys-Longchamps coined the name Raphus solitarius for these birds in 1848, as he believed the accounts referred to a species of Dodo. When 17th century paintings of white Dodos were discovered by 19th century naturalists, it was assumed they depicted these birds. Walter Rothschild suggested that the reason the painted specimens had yellow wing-tips instead of black as in the old descriptions might have been albinism. It has also been suggested that differences in paintings were due to sexual dimorphism, that the light plumage was a juvenile trait, a result of bleaching of old taxidermied specimens, or was simply artistic license. Some authors also believed it was a species similar to the Rodrigues Solitaire instead, as it was referred to by the same name, or even that there were white species of both the Dodo and Solitaire on the island.

Roelandt Savery painting with a whitish Dodo in the lower right, 1611

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 Hume and 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 greyish birds, possibly because he had by then seen another specimen. Since Réunion was not visited by Europeans until 1635, the 1611 painting could not have shown a bird from that island.

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. Cheke suggested to one of the authors, Francois Moutou, that the fossils might have been Réunion Solitaire, as published in 1995. The ibis was reassigned to the genus Threskiornis, now combined with the specific epithet solitarius from the binomial R. 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

The Dodo's significance as one of the best-known extinct animals and its singular appearance 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 or obsolete. Similarly, the phrase "to go the way of the Dodo" means to become extinct or obsolete, to fall out of common usage or practice or to become a thing of the past.

In 1865, the same year that George Clark started to publish 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 legal surname. The book's popularity made the Dodo a well-known icon of extinction.

Today, the Dodo regularly appears in works of popular fiction, and is used as a mascot for many kinds of products, especially in Mauritius. The Dodo appears as a supporter on the coat of arms of Mauritius. A smiling Dodo is the symbol of the Brasseries de Bourbon, a popular brewer on 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.

Previously unpublished 17th century illustration of a Dodo sold 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 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

References

Footnotes

Sources

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• BBC (20 November 2003). "Scientists pinpoint dodo's demise". BBC News. London. Retrieved 7 September 2006.
• Belloc, H. (1896). The Bad Child's Book of Beasts. London: Duckworth. {{cite book}}: Invalid |ref=harv (help)
• Template:IUCN
• Cheke, Anthony S. (2004). "The Dodo's last island" (PDF). Royal Society of Arts and Sciences of Mauritius. Retrieved 12 May 2012. {{cite web}}: Invalid |ref=harv (help)
• Cheke, Anthony S.; Hume, Julian Pender (2008). Lost Land of the Dodo: an Ecological History of Mauritius, Réunion & Rodrigues. New Haven and London: T. & A. D. Poyser. ISBN 978-0-7136-6544-4. {{cite book}}: Invalid |ref=harv (help)
• Christie's (2009). "Dutch School, 17th Century; A dodo". Christies.com. Retrieved 12 May 2012. {{cite web}}: Invalid |ref=harv (help)
• Cocks, T. (2006). "Natural disaster may have killed dodos". Reuters. Retrieved 30 August 2006. {{cite web}}: Invalid |ref=harv (help)
• Dissanayake, R. (2004). "What did the dodo look like?" (PDF). The Biologist. 51 (3). Society of Biology: 165–168. Retrieved 14 September 2011. {{cite journal}}: Invalid |ref=harv (help)
• Fryer, J. (2002). "Bringing the dodo back to life". BBC News. London. Retrieved 7 September 2006. {{cite news}}: Invalid |ref=harv (help)
• Fuller, Errol (2001). Extinct Birds (revised ed.). New York: Comstock. ISBN 978-0-8014-3954-4. {{cite book}}: Invalid |ref=harv (help)
• Fuller, Errol (2002). Dodo – From Extinction To Icon. London: HarperCollins. ISBN 978-0-00-714572-0. {{cite book}}: Invalid |ref=harv (help)
• Fuller, Errol (2003). The Dodo – Extinction In Paradise (first ed.). USA: Bunker Hill Publishing Inc. ISBN 978-1-59373-002-4. {{cite book}}: Invalid |ref=harv (help)
• Herhey, D. R. (2004). "Plant Science Bulletin, Volume 50, Issue 4". Botany.org. Retrieved 12 May 2012. {{cite web}}: Invalid |ref=harv (help)
• Hill, A. W. (1941). "The genus Calvaria, with an account of the stony endocarp and germination of the seed, and description of the new species". Annals of Botany. 5: 587–606. {{cite journal}}: Invalid |ref=harv (help)
• Hume, Julian Pender (2005). "Contrasting taphofacies in ocean island settings: the fossil record of Mascarene vertebrates". Proceedings of the International Symposium "Insular Vertebrate Evolution: the Palaeontological Approach". Monografies de la Societat d'Història Natural de les Balears. 12: 129–144. {{cite journal}}: Invalid |ref=harv (help)
• Hume, Julian Pender; Prys-Jones, R. P. (2005). "New discoveries from old sources, with reference to the original bird and mammal fauna of the Mascarene Islands, Indian Ocean" (PDF). Zoologische Mededelingen. 79 (3): 85–95. {{cite journal}}: Invalid |ref=harv (help)
• Hume, Julian Pender; Datta, Anna; Martill, David M. (2006). "Unpublished drawings of the Dodo Raphus cucullatus and notes on Dodo skin relics" (PDF). Bull. B. O. C. 126A: 49–54. ISSN 0007-1595. Retrieved 14 September 2011. {{cite journal}}: Invalid |ref=harv (help)
• Hume, Julian Pender; Walters, M. (2012). Extinct Birds. London: A & C Black. ISBN 978-1-4081-5725-1. {{cite book}}: Invalid |ref=harv (help)
• Jamieson, Alastair (22 June 2009). "Uncovered: 350-year-old picture of Dodo before it was extinct". The Daily Telegraph. London. Retrieved 8 September 2012. {{cite news}}: Invalid |ref=harv (help)
• Kallio, H. (2004). The Dodo and Mauritius Island. England: Dewi Lewis Publishing. {{cite book}}: Invalid |ref=harv (help)
• Kennedy, M. (2011). "Half a Dodo found in museum drawer". The Guardian. Retrieved 12 May 2012. {{cite news}}: Invalid |ref=harv (help)
• Kitchener, Andrew C. (1993). "Justice at last for the dodo". New Scientist: 24. {{cite journal}}: Unknown parameter |month= ignored (help)
• Laing, A. (27 August 2010). "Last surviving Dodo egg could be tested for authenticity". The Daily Telegraph. London. {{cite news}}: Invalid |ref=harv (help)
• Macmillan, A. (2000). Mauritius Illustrated: Historical and Descriptive, Commercial and Industrial Facts, Figures, & Resources. New Dheli: Asian Educational Services. p. 83. ISBN 978-81-206-1508-3. Retrieved 12 May 2012. {{cite book}}: Invalid |ref=harv (help)
• Mason, A. S. (1992). George Edwards: The Bedell and His Birds. London: Royal College of Physicians of London. ISBN 978-1-873240-48-9. Retrieved 12 May 2012. {{cite book}}: Invalid |ref=harv (help)
• Mayell, H. (28 February 2002). "Extinct Dodo Related to Pigeons, DNA Shows". National Geographic News. Retrieved 19 January 2009. {{cite web}}: Invalid |ref=harv (help)
• Mourer-Chauviré, C.; Moutou, F. (1987). "Découverte d'une forme récemment éteinte d'ibis endémique insulaire de l'île de la Réunion Borbonibis latipes n. gen. n. sp". Comptes rendus de l'Académie des sciences. Série D (in French). 305 (5): 419–423. {{cite journal}}: Invalid |ref=harv (help); Unknown parameter |trans_title= ignored (|trans-title= suggested) (help)
• Oudemans, A. C. (1917). Dodo-studien, naar aanleidingvan de vondst van een gerelsteen met Dodo-beeld van 1561 to Vere. 19 (in Dutch). Amsterdam: Verhandelingen der Koninklijke Akademic van Wetenschappen. pp. 1–140. {{cite book}}: Invalid |ref=harv (help)
• "The Oxford Dodo" (PDF). Learning More. The Oxford University Museum of Natural History. 29 March 2006. Retrieved 13 September 2011.
• Quammen, David (1996). The Song of the Dodo: Island Biogeography in an Age of Extinction. New York: Touchstone. ISBN 978-0-684-82712-4. {{cite book}}: Invalid |ref=harv (help)
• Ravilious, K. (2007). "Dodo Skeleton Found on Island, May Yield Extinct Bird's DNA". National Geographic. {{cite web}}: Invalid |ref=harv (help)
• Reinhardt, Johannes Theodor (1842–1843). "Nøjere oplysning om det i Kjøbenhavn fundne Drontehoved". Nat. Tidssk. Krøyer. IV.: 71–72. 2.{{cite journal}}: CS1 maint: ref duplicates default (link)
• Rothschild, Walter (1907). Extinct Birds (PDF). London: Hutchinson & Co. {{cite book}}: Invalid |ref=harv (help)

• Staub, France (1996). "Dodo and solitaires, myths and reality". Proceedings of the Royal Society of Arts & Sciences of Mauritius. 6: 89–122. {{cite journal}}: Invalid |ref=harv (help)
• Strickland, Hugh Edwin; Melville, A. G. (1848). The Dodo and Its Kindred; or the History, Affinities, and Osteology of the Dodo, Solitaire, and Other Extinct Birds of the Islands Mauritius, Rodriguez, and Bourbon. London: Reeve, Benham and Reeve. {{cite book}}: Invalid |ref=harv (help)
• Unhook, D. P. (2006). "Mauritius: Footprints From the Past". Expresser's. Retrieved 26 September 2006. {{cite web}}: Invalid |ref=harv (help) (subscription required)

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