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=== Diet and feeding === [[File:Roelandt Savery - 'Dodo Birds', Chalk, black and amber on cream paper.jpg|thumb|alt=Sketch of three dodos, two in the foreground, one in the distance|Savery sketch of three dodos from {{circa|lk=no|1626}}, [[Crocker Art Gallery]]]] A 1631 Dutch letter (long thought lost, but rediscovered in 2017) is the only account of the dodo's diet, and also mentions that it used its beak for defence. The document uses [[word-play]] to refer to the animals described, with dodos presumably being an [[allegory]] for wealthy mayors:<ref name=Winters2017>{{cite journal |last1=Winters|first1=R. |last2=Hume|first2=J. P. |last3=Leenstra|first3=M. |title=A famine in Surat in 1631 and Dodos on Mauritius: a long lost manuscript rediscovered |journal=Archives of Natural History |date=2017 |volume=44 |issue=1 |pages=134β150 |doi=10.3366/anh.2017.0422}}</ref> {{quotation|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.<ref name=Winters2017/>}} In addition to fallen fruits, the dodo probably subsisted on nuts, seeds, bulbs, and roots.{{sfn|Fuller|2002|p=42}} 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 probably given a wide range of food on the long sea journeys.{{sfn|Cheke|Hume|2008|pp=37β38}} 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. The Mauritian ornithologist [[France Staub]] suggested in 1996 that they mainly fed on [[palm (plant)|palm]] fruits, and he attempted to correlate the fat-cycle of the dodo with the fruiting regime of the palms.<ref name=Staub1996/> [[File:Clusius dodo.jpg|thumb|alt=Drawing of a dodo next to a large gizzard stone|left|Dodo and its [[gizzard stone]] by [[Carolus Clusius]] from 1605, copied from an illustration in the journal of [[Jacob van Neck]]]] Skeletal elements of the upper jaw appear to have been [[rhynchokinetic]] (movable in relation to each other), which must have affected its feeding behaviour. In extant birds, such as [[frugivorous]] (fruit-eating) pigeons, kinetic [[premaxillae]] help with consuming large food items. The beak also appears to have been able to withstand high force loads, which indicates a diet of hard food.<ref name=ClaessensMeijer2016/> Examination of the brain endocast found that though the brain was similar to that of other pigeons in most respects, the dodo had a comparatively large [[olfactory bulb]]. This gave the dodo a good sense of smell, which may have aided in locating fruit and small prey.<ref name="Endocast">{{cite journal|last1=Gold|first1=M. E. Leone|last2=Bourdon|first2=E.|last3=Norell|first3=M. A.|title=The first endocast of the extinct dodo (''Raphus cucullatus'') and an anatomical comparison amongst close relatives (Aves, Columbiformes)|journal=Zoological Journal of the Linnean Society|volume=177|issue=4|date=2016|pages=950β963|doi=10.1111/zoj.12388}}</ref> Several contemporary sources state that the dodo used [[Gastrolith]]s (gizzard stones) to aid digestion. The English writer Sir [[Hamon L'Estrange]] witnessed a live bird in London and described it as follows: {{quotation |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.{{sfn|Fuller|2002|p=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.<ref>{{cite journal| doi = 10.1642/0004-8038(2005)122[1003:APCBCM]2.0.CO;2| last1 = Storer| first1 = Robert W.| year = 2005| title = A possible connection between crop milk and the maximum size attainable by flightless pigeons| journal = The Auk| volume = 122| issue = 3| pages = 1003| type = Submitted manuscript| doi-access = free}}</ref> In 1973, the [[Sideroxylon grandiflorum|tambalacoque]], also known as the dodo tree, was thought to be dying out on Mauritius, to which it is [[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.<ref>{{cite journal| doi = 10.1126/science.197.4306.885| last = Temple | first = S. A.| date=August 1977 | title = Plant-Animal Mutualism: Coevolution with Dodo Leads to Near Extinction of Plant| journal = Science| volume = 197| issue = 4306| pages = 885β886| pmid = 17730171|bibcode = 1977Sci...197..885T | s2cid = 2392411 }}</ref> Temple overlooked reports from the 1940s that found that tambalacoque seeds germinated, albeit very rarely, without being [[abrasion (mechanical)|abraded]] during digestion.<ref>{{cite journal | last = Hill | first = A. W. | year = 1941 | title = The genus ''Calvaria'', with an account of the stony endocarp and germination of the seed, and description of the new species | journal = Annals of Botany | volume = 5 | issue = 4 | pages = 587β606 | doi = 10.1093/oxfordjournals.aob.a087409 }}</ref> Others have contested his hypothesis and suggested that the decline of the tree was exaggerated or seeds were also distributed by other extinct animals such as ''[[Cylindraspis]]'' [[tortoise]]s, [[fruit bat]]s, or the broad-billed parrot.<ref>{{cite web | last = Herhey | first = D. R. | year = 2004 | title = Plant Science Bulletin, Volume 50, Issue 4 | publisher = Botany.org | url = http://www.botany.org/PlantScienceBulletin/psb-2004-50-4.php#Dodo | access-date = 12 May 2012 | archive-date = 14 May 2013 | archive-url = https://web.archive.org/web/20130514022621/http://www.botany.org/PlantScienceBulletin/psb-2004-50-4.php#Dodo | url-status = live }}</ref> According to Wendy Strahm and Anthony Cheke, two experts in the [[ecology]] of the Mascarene Islands, 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.<ref>{{cite journal| doi = 10.2307/3545415| last1 = Witmer | first1 = M. C.| last2 = Cheke | first2 = A. S.| date=May 1991 | title = The Dodo and the Tambalacoque Tree: An Obligate Mutualism Reconsidered| journal = Oikos| volume = 61| issue = 1| pages = 133β137| jstor = 3545415| bibcode = 1991Oikos..61..133W }}</ref> 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 America]]n [[megafauna]] in the same way, but now rely on domesticated cattle for this service.{{sfn|Cheke|Hume|2008|p=38}}
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