| Dodo Temporal range: Late Holocene |
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|---|---|
| Dodo reconstruction reflecting modern research, at Oxford University Museum of Natural History[1] | |
| Conservation status | |
| 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) |
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| Former range (in red) | |
| Synonyms | |
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The dodo (Raphus cucullatus) was a flightless bird endemic to the Indian Ocean island of Mauritius. Related to pigeons and doves, it stood about a meter (3.3 feet) tall, weighing about 20 kilograms (44 lb).
The dodo lost the power of flight because food was abundant and predators were absent. By 1681, all dodos had been killed by settlers or their domesticated animals.[3] The extinction of the bird, within 80 years of its discovery, made people realise for the first time that humans could induce the extinction of plants and animals.[4]
The bird is mainly represented by sub-fossil material today, and its external features are only known from paintings and written accounts from the 17th century, but since these vary considerably, and only a few sketches are known to have been drawn from life, mystery remains over its exact appearance. The same is true of its behaviour.
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Many different affinities were suggested for the dodo, including to ostriches and vultures, until Johannes Theodor Reinhardt proposed they were ground doves, based on studies of a dodo skull in Copenhagen.[5] This view was later supported by Strickland and Melville after a dissection of the preserved head at Oxford Museum.
The theory has recently been confirmed by mtDNA cytochrome b and 12S rRNA sequence[6] analysis, wherein DNA extracted from a tarsal from the dodo-foot in Oxford was compared with genetic material from other birds, that suggests that the ancestors of the dodo diverged from those of its closest known relative, the likewise extinct Rodrigues Solitaire, around the Paleogene-Neogene boundary. As the Mascarenes are of volcanic origin and less than 10 million years old, the ancestors of both these birds likely remained capable of flight for considerable time after the separation of their lineages. The same study has been interpreted to show that the Southeast Asian Nicobar Pigeon is the closest living relative of the dodo and the Rodrigues Solitaire.[7] The generic name of the dodo-like Tooth-billed Pigeon from Samoa is Didunculus, which means "little Dodo"[8], and it was also shown to be quite close to the dodo in the study.
However, the proposed phylogeny is problematical regarding the relationships of other taxa.[9] All that can be presently said with certainty is that the ancestors of the didine birds were pigeons from Southeast Asia or the Wallacea, which agrees with the origin of most of the Mascarenes' birds.
For a long time, the dodo and the Rodrigues Solitaire, collectively termed "didines", were placed in a family of their own, the Raphidae. This was because their relationships to other groups of birds, such as rails, was unresolved. Recently, it has been suggested that the didine group should be dissolved and the dodo and solitaire placed in the existing subfamily Raphinae within the Columbidae[10]
The etymology of the word dodo is unclear. Some ascribe it to the Dutch word dodoor for "sluggard", but it more likely is related to dodaars, which means either "fat-arse" or "knot-arse," referring to the knot of feathers on the hind end. The first recording of the word dodaerse is in captain Willem van Westsanen's journal in 1602.[11] Sir Thomas Herbert used the word dodo in 1627,[12] but it is unclear whether he was the first; the Portuguese had visited the island in 1507, but, as far as is known, did not use the word. Nevertheless, according to the Encarta Dictionary and Chambers Dictionary of Etymology, "dodo" derives from Portuguese doudo (currently doido) meaning "fool" or "crazy".[13][14] However, the present Portuguese name for the bird, dodô, is taken from the internationally used word dodo.
David Quammen considered the idea that dodo was an onomatopoeic approximation of the bird's own call, a two-note pigeony sound like "doo-doo".[15]
The bird was also referred to as "dronte" by the Dutch, meaning "swollen," a name still used in some languages.
The phrase "dead as a dodo" has come to mean undoubtedly and unquestionably dead, while 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, both in reference to the untimely extinction of the bird.
No complete dodo specimens exist to this day, making the external appearance, such as plumage and coloration, hard to determine. But from sub-fossil remains and remnants of the birds that were brought to Europe in the 17th century, it is known that they were very large birds, possibly weighing up to 23 kg (50 pounds), although some estimations give a weight of about 10.6-17.5 kg.[16] The sternum was insufficient to support flight and the wings were very small; these ground-bound birds evolved to take advantage of an island ecosystem with no predators. It had a 23-centimeter (9-inch) bill with a hooked point. A study of the few remaining feathers on the Oxford head showed that they were plumaceous (downy) rather than vaned.[17]
About 15 illustrations were made while interaction with the dodo was possible (1598-1640), and they are the primary evidence for its external appearance, along with various written accounts of ecounters with dodos on Mauritius, and with a captive bird in London.[18]
According to most renditions, 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, with a green, black and yellow beak, and the legs were stout and yellowish, with black claws.
An early account from Van Neck's journey in 1598 describes the bird thus:
"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."[19]
One of the most detailed descriptions is by Sir Thomas Herbert from 1634:
"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 i 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.[20]"
Due to the fact that 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 simply after crudely stuffed specimens, which would also affect their reliability. Julian Hume has argued that the nostrils of the dodo would had been slits in life, as seen in the Gelderland, Saftleven, Crocker Gallery and Mansur images. The gaping nostrils often seen in dodo paintings were instead an artifact of drying.
The traditional image of the dodo is of a very fat, clumsy bird, hence the synonym Didus ineptus, but this view may be exaggerated. The general opinion of scientists today is that the old European drawings showed overfed captive specimens.[21][22] The Dutch painter Roelant Savery was the most prolific and influential illustrator of the dodo, and depicted it at least six times. A famous painting of his from 1626 now in the British Museum has since became the standard image of a dodo. The image shows a particularly fat bird, and is the source of many other dodo restorations.[23][24]
A 17th century painting attributed to the Mughal artist Ustad Mansur which was rediscovered in the 1950s shows a dodo along with native Indian birds. It depicts a slimmer, brownish bird, and is regarded by professor A. Ivanov and dodo expert Julian Hume as one of the most accurate depictions of a dodo.[25]
Not much is known of the behaviour of the dodo and most contemporary descriptions are very brief. It is mentioned that the bird was living on fruit, and nested on the ground, laying a single egg. A description by François Cauche from 1651 contains some details about the egg and call:
"The call is like that of a gosling but they are quite unpalatable to eat... They lay one egg, which is quite as large as a penny bun, against which they lay a white stone the size of a chicken's egg. They lay their egg on a nest of grass which they collect and they place the nest in the woods. If one kills the young you find a grey stone in the gizzard. We named them the birds of Nazareth.[26]"
A single account exists about the diet of the dodo, a document from 1631 rediscovered in 1887 but now lost:
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.[27]"
As Mauritius has marked dry and wet seasons, the dodo probably fattened itself on ripe fruits at the end of the wet season to live through the dry season when food was scarce; contemporary reports speak of the birds' "greedy" appetite.
Several contemporary sources state that the dodo used gizzard stones. The English historian Sir Hamon L'Estrange, who witnessed a live bird, described it as such:
"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."[28]
The tambalacoque, also known as the "dodo tree", was hypothesized by Stanley Temple to have been eaten from by dodos, and only by passing through the digestive tract of the dodo could the seeds germinate; he claimed that the tambalacocque was now nearly extinct due to the dodo's disappearance. 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 effect of feeding fruits to turkeys was unclear. Temple also overlooked reports on tambalacoque seed germination by A. W. Hill in 1941 and H. C. King in 1946, who found the seeds germinated, albeit very rarely, without abrading.[29][30][31][32]
Temple's hypothesis has been contested. Others have suggested the decline of the tree was exaggerated, or that other extinct animals may also have been distributing the seeds, such as Cylindraspis tortoises, fruit bats or the Broad-billed Parrot. Wendy Strahm and Anthony Cheke, two experts in Mascarene ecology, claim that while a rare tree, it has germinated since the demise of the Dodo and numbers a few hundred, not 13.[33]
In 2004, Botanical Society of America's Plant Science Bulletin disputed Dr. Temple's research as flawed. They published evidence as to why the dodo's extinction did not directly cause the increasing disappearance of young calvaria trees, and suggesting that tortoises would have been more likely to disperse the seeds than dodo, hence discrediting Temple's view on the dodo and the Calvaria's sole survival relationship.[34]
The earliest known descriptions of the bird were made by Dutch travelers to what is now the island of Mauritius, east of Madagascar. Only a dozen such contemporary accounts based on observation are reliable, and many seem based on earlier accounts.[35] The first accounts can be found in reports of the 1598 voyage of Admiral Jacob van Neck published in 1601, which also featured the first published illustration of the bird. According to Julian Hume, the following may be the first mention of the bird, here referred to as penguins, which was not used for sphenisciformes 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."[36]
One of the original names used for the dodo was "walghvogel" ("wallow bird" or "loathsome bird," in reference to its taste), first used in the journal of vice-admiral Wybrand van Warwijck, who visited the island with the Van Neck expedition:
"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 travel journal of the Dutch East India Company ship Gelderland (1601–1603) contains the only known sketches that are known to have been drawn from living or recently killed specimens on Mauritius, attributed to the professional artist Joris Joostensz Laerle, who also drew other now-extinct Mascarene birds.
At least three or four live birds were taken to Europe, some of which are believed to have been depicted alive, and some of which may be the source of the few non-fossil remains that are known today. Two live specimens were taken to India in the 1600s according to Peter Mundy, and one of these appears to have been depicted in an Indian painting.[37]
The existing remains of the birds taken to Europe consist of a dried head and foot bones in Oxford University Museum of Natural History, a partial skull and leg-bones in the National Museum (Prague), a skull in the Zoological Museum of Copenhagen, and a dried foot once housed in the British Museum, which is now lost. The only known soft tissue remains, the head and foot in Oxford Museum, belonged to the last known stuffed dodo, which had been kept in Oxford's Ashmolean Museum. Around 1755, however, the museum's curator or director ordered the remaining pieces discarded because of severe decay. The remaining soft tissue has since degraded further, as the head was dissected in the late 19th century, separating the skin from the skull, and the foot is in a skeletal state.[38]
Few took particular notice of the bird immediately after its extinction. By the early 19th century it seemed altogether too strange a creature, and many believed it a myth. In 1848, H. E. Strickland and A. G. Melville published a book titled 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 in which they attempted to separate Dodo myth from reality. With the discovery of the first batch of sub-fossil dodo bones in the Mauritian swamp, the Mare aux Songes, and the reports written about them by George Clarke, government schoolmaster at Mahébourg, from 1865 on, interest in the bird was rekindled.
Until recently, few associated dodo skeletons were known, most of the material consisting of isolated and scattered bones. Many of these were excavated by request of Richard Owen, who wanted to be the first to describe the post-cranial anatomy of the dodo.[39] Dublin's Natural History Museum and the Oxford University Museum of Natural History, among others, have a specimen assembled from these dissociated remains. 26 museums around the world have significant holdings of dodo material. An alleged dodo egg is on display at the East London museum in South Africa, but genetic studies are underway to determine its authencity.[40]
In October 2005, part of the Mare aux Songes, the most important site of dodo remains, was excavated by an international team of researchers. Many remains were found, including bones from birds of various stages of maturity,[41] and several bones obviously belonging to the skeleton of one individual bird and preserved in natural position.[13] These findings were made public in December 2005 in the Naturalis museum in Leiden.
In June 2007, adventurers exploring a cave in Mauritius discovered the most complete and best-preserved dodo skeleton ever.[42]
The supposed "Réunion Solitaire" or "White Dodo" of Réunion is now believed to have been 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 Reunion around 1619, mentioned that it was inhabited by "Dod-eersen", though without mentioning colouration. A white bird was first described as follows in 1625 by Mr. Tatton, the Chief Officer of Captain Castleton:
"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."
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."
Nauralists like Walter Rothschild assumed these descriptions were of the white dodo shown in the paintings, and a new species, Didus solitarius, was erectd. Rothschild suggested that the reason why the painted specimens had yellow wing-tips instead of black as in the old descriptions might have been albinism.[43] Others believed it was a species similar to the Rodrigues Solitaire, as it was referred to as "Solitaire" as well.
A record from 1735 suggests that M. de la Bourdonnaye, the Governor of Mauritius and Réunion at the time, had a live dodo sent from Réunion to France, but nothing is known of its fate.[44]
The Pieter Withoos painting, which was discovered first, appears to be based on an earlier painting by Pieter Holsteyn, of which three versions are known to have existed. According to Julian Hume and Anthony Cheke, it appears that all depictions of white dodos were based on a 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 coloring" 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.[45] It has also been suggested that the light plumage was a juvenile trait, or that it was simply artistic license.[46]
Since Reunión was not visited by Europeans until 1635, the 1611 painting could not have shown a bird from there.[47] In 1987, fossils of a recently extinct species of ibis in the genus Threskiornis were described from Réunion, and it is now believed that this bird was the source of the old sailors' descriptions, since they are white with black parts and have long, slender beaks, unlike dodos. No fossil remains of dodo-like birds have ever been found on the island, however.
Although many later writings say that the meat tasted bad, early journals say only that the meat was tough but good, though not as delectable as abundantly available pigeons.[48]
The journal of Willem Van West-Zanen from 1602, which was not published until 1648, contains some accounts of interaction with dodos:
"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."
As with many animals that have evolved in isolation from significant predators, the dodo was entirely fearless of people, and this, in combination with its flightlessness, made it easy prey for humans.[49] However, journals are full of reports regarding the bad taste and tough meat of the dodo, while other local species such as the Red Rail were praised for their taste. When humans first arrived on Mauritius, they also brought with them other animals that had not existed on the island before, including dogs, pigs, cats, rats, and Crab-eating Macaques, which plundered the dodo nests, while humans destroyed the forests where the birds made their homes;[50] the impact these animals—especially the pigs and macaques—had on the dodo population is currently considered to have been more severe than that of hunting. The 2005 expedition's finds are apparently of animals killed by a flash flood; such mass mortalities would have further jeopardized a species already in danger of becoming extinct.[51]
Although there are scattered reports of mass killings of dodos for provisioning of ships, archaeological investigations have hitherto found scant evidence of human predation on these birds. Bones of at least two dodos were found in caves at Baie du Cap, fugitive slaves and convicts used these caves for shelters in the 17th century—but due to their isolation in high, broken terrain, they were not easily accessible to dodos.[52]
There is some controversy surrounding the extinction date of the dodo. Roberts & Solow state that the last confirmed dodo sighting is the one reported by shipwrecked mariner Volkert Evertsz in 1662:
"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."
However, many other sources suggest the more conjectural date of 1681. Roberts & Solow point out that because the sighting prior to 1662 was in 1638, the dodo was likely already very rare by the 1660s, and thus a disputed report from 1674 cannot be dismissed out-of-hand.[53]
Statistical analysis of the hunting records of Isaac Johannes Lamotius give a new estimated extinction date of 1693, with a 95% confidence interval of 1688 to 1715; the last reported sighting is from the hunting records of Isaac Johannes Lamotius, who gives the year 1688, but it has been suggested that by this time the Dutch name "dodaers" had been transferred to the flightless Red Rail, which is now also extinct.[54] Considering more circumstantial evidence such as travelers' reports and the lack of good reports after 1689,[52] it is likely that the dodo became extinct before 1700; the last dodo died a little more than a century after the species' discovery in 1581.[55]
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 to symbolize a concept or object that will or has become out of date, as in the expression "dead as a dodo" or "gone the way of the dodo".[56][57] In the same year in which George Clarke started to publish his reports, the newly vindicated bird was featured as a character in Lewis Carroll's Alice's Adventures in Wonderland. With the popularity of the book, the dodo became a well-known and easily recognizable icon of extinction.[58]
The dodo is used by many environmental organizations that promote the protection of endangered species, such as the Durrell Wildlife Conservation Trust and the Jersey Zoological Park, founded by Gerald Durrell.[59]
The dodo rampant appears on the coat of arms of Mauritius.[50]
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.[60] It is unknown whether it was drawn after a live or stuffed specimen, or simply a copy of another image. It was sold for £44,450.[61]
