Editing Dodo (section)

== Behaviour and ecology ==
[[File:Dodo by Julian P. Hume.jpg|thumb|left|upright|Modern [[life restoration]] of a slim dodo, by [[Julian P. Hume]]]]

Little is known of the behaviour of the dodo, as most contemporary descriptions are very brief. Based on weight estimates, it has been suggested the male could reach the age of 21, and the female 17.<ref name=Livezey1993/> Studies of the [[cantilever]] strength of its leg bones indicate that it could run quite fast.<ref name=KitchenerAugust1993/> The legs were robust and strong to support the bulk of the bird, and also made it agile and manoeuvrable in the dense, pre-human landscape. Though the wings were small, well-developed muscle scars on the bones show that they were not completely [[vestigial]], and may have been used for [[display behaviour]] and balance; extant pigeons also use their wings for such purposes.<ref name=ClaessensMeijer2016/> Unlike the Rodrigues solitaire, there is no evidence that the dodo used its wings in intraspecific combat. Though some dodo bones have been found with healed fractures, it had weak pectoral muscles and more reduced wings in comparison. The dodo may instead have used its large, hooked beak in territorial disputes. Since Mauritius receives more rainfall and has less seasonal variation than Rodrigues, which would have affected the availability of resources on the island, the dodo would have less reason to evolve aggressive territorial behaviour. The Rodrigues solitaire was therefore probably the more aggressive of the two.<ref>{{cite journal | last1 = Hume | first1 = J. P. | author-link1 = Julian Pender Hume| last2 = Steel | first2 = L. | doi = 10.1111/bij.12087 | title = Fight club: A unique weapon in the wing of the solitaire, ''Pezophaps solitaria'' (Aves: Columbidae), an extinct flightless bird from Rodrigues, Mascarene Islands | journal = Biological Journal of the Linnean Society | pages = 32–44| year = 2013 | volume=110| doi-access = free }}</ref> In 2016, the first 3D [[endocast]] was made from the brain of the dodo; the [[Brain-to-body mass ratio|brain-to-body-size ratio]] was similar to that of modern pigeons, indicating that dodos were probably equal in intelligence.<ref name="Endocast"/>

[[File:Gelderland1601-1603 Mauritius de Nassauw 1601.jpg|thumb|alt=Old map showing a Mauritian bay, with a D indicating where dodos were found|1601 map of a bay on Mauritius; the small D on the far right side marks where dodos were found]]
The preferred habitat of the dodo is unknown, but old descriptions suggest that it inhabited the woods on the drier coastal areas of south and west Mauritius. This view is supported by the fact that the [[Mare aux Songes]] swamp, where most dodo remains have been excavated, is close to the sea in south-eastern Mauritius.{{sfn|Fuller|2002|p=23}} Such a limited distribution across the island could well have contributed to its extinction.{{sfn|Fuller|2002|p=41}} A 1601 map from the ''Gelderland'' journal shows a small island off the coast of Mauritius where dodos were caught. Julian Hume has suggested this island was [[l'île aux Benitiers|l'île aux Bénitiers]] in [[Tamarin Bay]], on the west coast of Mauritius.{{sfn|Fuller|2002|p=54}}<ref name=Hume2003/> Subfossil bones have also been found inside caves in highland areas, indicating that it once occurred on mountains. Work at the Mare aux Songes swamp has shown that its habitat was dominated by [[tambalacoque]] and ''[[Pandanus]]'' trees and endemic palms.<ref name=Hume2017/> The near-coastal placement and wetness of the Mare aux Songes led to a high diversity of plant species, whereas the surrounding areas were drier.<ref name=Rijsdijk2016>{{cite journal|last1=Rijsdijk|first1=K. F. |last2=Hume|first2=J. P. |last3=Louw|first3=P. G. B. D. |last4=Meijer|first4=H. J. M. |last5=Janoo|first5=A. |last6=De Boer|first6=E. J. |last7=Steel|first7=L. |last8=De Vos|first8=J. |last9=Van Der Sluis|first9=L. G.|last10=Hooghiemstra|first10=H. |last11=Florens|first11=F. B. V. |last12=Baider|first12=C. |last13=Vernimmen|first13=T. J. J. |last14=Baas|first14=P. |last15=Van Heteren|first15=A. H. |last16=Rupear|first16=V. |last17=Beebeejaun|first17=G. |last18=Grihault|first18=A. |last19=Van Der Plicht|first19=J. |last20=Besselink|first20=M. |last21=Lubeek|first21=J. K. |last22=Jansen|first22=M. |last23=Kluiving|first23=S. J. |last24=Hollund|first24=H. |last25=Shapiro|first25=B. |last26=Collins|first26=M. |last27=Buckley|first27=M. |last28=Jayasena|first28=R. M. |last29=Porch|first29=N. |last30=Floore|first30=R. |last31=Bunnik|first31=F. |last32=Biedlingmaier|first32=A. |last33=Leavitt|first33=J. |last34=Monfette|first34=G. |last35=Kimelblatt|first35=A. |last36=Randall|first36=A. |last37=Floore|first37=P. |last38=Claessens|first38=L. P. A. M. |display-authors=8 |title=A review of the dodo and its ecosystem: insights from a vertebrate concentration Lagerstätte in Mauritius |journal=Journal of Vertebrate Paleontology |date=2016 |volume=35 |issue=sup 1|pages=3–20 |doi=10.1080/02724634.2015.1113803 |doi-access=free |hdl=1893/25225 |ref={{sfnRef|Rijsdijk et al.|2016}}|hdl-access=free }}</ref>

Many endemic species of Mauritius became extinct after the arrival of humans, so the [[ecosystem]] of the island is badly damaged and hard to reconstruct. Before humans arrived, Mauritius was entirely covered in forests, but very little remains of them today, because of [[deforestation]].<ref>{{cite journal| doi= 10.1017/S0030605300020457| last= Cheke | first= A. S.| year= 1987| title= The legacy of the dodo—conservation in Mauritius| journal= Oryx| volume= 21| issue= 1| pages= 29–36| s2cid= 86670941 | doi-access= free}}</ref> The surviving endemic [[fauna]] is still seriously threatened.<ref>{{cite journal| doi = 10.1017/S0030605300012643| last = Temple | first = S. A.| year = 1974| title = Wildlife in Mauritius today| journal = Oryx| volume = 12| issue = 5| pages = 584–590| s2cid = 86773685 | doi-access = free}}</ref> The dodo lived alongside other recently extinct Mauritian birds such as the flightless red rail, the [[broad-billed parrot]], the [[Mascarene grey parakeet]], the [[Mauritius blue pigeon]], the [[Mauritius scops 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. Some plants, such as ''[[Casearia tinifolia]]'' and the [[palm orchid]], have also become extinct.{{sfn|Cheke|Hume|2008|pp=49–52}}

=== 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=134–150 |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=37–38}} 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=950–963|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 = 885–886| 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 = 587–606 | 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 = 133–137| 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}}

=== Reproduction and development ===
[[File:Dodo egg replica.jpg|thumb|alt=Replica dodo egg and nest|Replica of an alleged dodo egg in a reconstructed nest, [[East London Museum]]]]

As it was flightless and terrestrial and there were no [[mammalian]] predators or other kinds of natural enemy on Mauritius, the dodo probably nested on the ground.{{sfn|Fuller|2002|pp=43–44}} The account by François Cauche from 1651 is the only description of the egg and the [[Bird call|call]]:
{{quotation |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 [[goose|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.<ref name=Strickland4to112/>}}

{{multiple image
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|image1 = Dodo bone thin sections showing ontogenetic growth series.jpg
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|caption1 = Thin sections of hindlimb bones showing stages of the [[growth series]]

|image2 = Dodo life history.jpg
|caption2 = Diagram showing life history events of a dodo based on [[histology]] and accounts
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}}
Cauche's account is problematic, since it also mentions that the bird he was describing had three toes and no tongue, unlike dodos. This led some to believe that Cauche was describing a new species of dodo ("''Didus nazarenus''"). The description was most probably mingled with that of a [[cassowary]], and Cauche's writings have other inconsistencies.{{sfn|Cheke|Hume|2008|p=27}} A mention of a "young ostrich" taken on board a ship in 1617 is the only other reference to a possible juvenile dodo.{{sfn|Cheke|Hume|2008|p=162}} An egg claimed to be that of a dodo is stored in the [[East London Museum]] in South Africa. It was donated by the South African museum official [[Marjorie Courtenay-Latimer]], whose great aunt had received it from a captain who claimed to have found it in a swamp on Mauritius. In 2010, the curator of the museum proposed using genetic studies to determine its authenticity.<ref>{{cite news | last = Laing | first = A. | date = 27 August 2010 | work = The Daily Telegraph | location = London | title = Last surviving Dodo egg could be tested for authenticity | url = https://www.telegraph.co.uk/science/science-news/7968811/Last-surviving-Dodo-egg-could-be-tested-for-authenticity.html | archive-url = https://web.archive.org/web/20100829160254/http://www.telegraph.co.uk/science/science-news/7968811/Last-surviving-Dodo-egg-could-be-tested-for-authenticity.html | url-status = dead | archive-date = 29 August 2010 }}</ref> It may instead be an aberrant [[ostrich egg]].<ref name=Fuller2002p43/>

Because of the possible single-egg clutch and the bird's large size, it has been proposed that the dodo was [[K-selected]], meaning that it produced few [[altricial]] offspring, which required parental care until they matured. Some evidence, including the large size and the fact that tropical and frugivorous birds have slower growth rates, indicates that the bird may have had a protracted development period.<ref name=Livezey1993/> The fact that no juvenile dodos have been found in the Mare aux Songes swamp may indicate that they produced little offspring, that they matured rapidly, that the breeding grounds were far away from the swamp, or that the risk of miring was seasonal.<ref name=Meijer2012p177>{{cite journal| last1 = Meijer| first1 = H. J. M.| last2 = Gill| first2 = A.| last3 = de Louw| first3 = P. G. B.| last4 = van den Hoek Ostende| first4 = L. W.| last5 = Hume| first5 = J. P.| last6 = Rijsdijk| first6 = K. F.| doi = 10.1007/s00114-012-0882-8| title = Dodo remains from an in situ context from Mare aux Songes, Mauritius| journal = [[Naturwissenschaften]]| volume = 99| issue = 3| pages = 177–184| year = 2012| pmid = 22282037| bibcode = 2012NW.....99..177M| s2cid = 514542| url = https://repository.si.edu/bitstream/handle/10088/18486/vz_meijer_naturwissenschaften_2012.pdf| type = Submitted manuscript| access-date = 6 August 2018| archive-date = 23 July 2018| archive-url = https://web.archive.org/web/20180723130845/https://repository.si.edu/bitstream/handle/10088/18486/vz_meijer_naturwissenschaften_2012.pdf| url-status = live}} pp.&nbsp;177{{ndash}}184.</ref>

A 2017 study examined the [[histology]] of [[thin-sectioned]] dodo bones, modern Mauritian birds, local ecology, and contemporary accounts, to recover information about the life history of the dodo. The study suggested that dodos bred around August, after having potentially fattened themselves, corresponding with the fat and thin cycles of many vertebrates of Mauritius. The chicks grew rapidly, reaching robust, almost adult, sizes, and sexual maturity before Austral summer or the cyclone season. Adult dodos which had just bred [[moulted]] after Austral summer, around March. The feathers of the wings and tail were replaced first, and the moulting would have completed at the end of July, in time for the next breeding season. Different stages of moulting may also account for inconsistencies in contemporary descriptions of dodo plumage.<ref>{{cite journal |last1=Angst |first1=D. |last2=Chinsamy |first2=A. |last3=Steel |first3=L. |last4=Hume |first4=J. P. |title=Bone histology sheds new light on the ecology of the dodo (''Raphus cucullatus'', Aves, Columbiformes)|journal=Scientific Reports |date=2017 |volume=7 |issue=1 |page=7993 |doi=10.1038/s41598-017-08536-3 |pmc=5570941 |issn=2045-2322 |pmid=28839147 |bibcode=2017NatSR...7.7993A|url=https://link.springer.com/content/pdf/10.1038/s41598-017-08536-3.pdf}}</ref>