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== Physical remains == === 17th-century specimens === {{multiple image | direction = horizontal |align = right |total_width = 350 |image1 = Dodo casts c1800.jpg |alt1 = White casts |image2 = Dodo head.jpg |alt2 = |footer = Cast of the Oxford head before dissection and the lost London foot at [[Booth Museum]], and illustration of same head }} The only extant remains of dodo specimens 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 in the [[National Museum, Prague]]. The last two were rediscovered and identified as dodo remains in the mid-19th century.{{sfn|Fuller|2002|pp=116–129}} Several stuffed dodos were also mentioned in old museum inventories, but none are known to have survived.<ref>{{cite journal| doi = 10.3366/anh.1992.19.2.145| last = Ovenell | first = R. F.| date=June 1992 | title = The Tradescant Dodo| journal = Archives of Natural History| volume = 19| issue = 2| pages = 145–152}}</ref> Apart from these remains, a dried foot which belonged to Dutch professor [[Pieter Pauw]] was mentioned by [[Carolus Clusius]] in 1605. Its provenance is unknown, and it is now lost, but it may have been collected during the Van Neck voyage.<ref name=Hume2006/> Purported stuffed dodos seen in museums around the world today have in fact been made from feathers of other birds; many by British taxidermist [[Rowland Ward]]'s company.{{sfn|Fuller|2002|pp=116–129}} {{multiple image |align = left |total_width = 350 |image1 = London dodo foot.jpg |alt1 = |image2 = London Dodo leg.jpg |alt2 = Lithograph of a dried foot |footer = Coloured engraving of the now lost London foot from 1793 (left), and 1848 lithograph of same in multiple views }} 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 was moved to the [[Ashmolean Museum]] in 1659.<ref name=Hume2006/> It has been suggested that this might be the remains of the bird that Hamon L'Estrange saw in London, the bird sent by Emanuel Altham, or a donation by Thomas Herbert. Since the remains do not show signs of having been mounted, the specimen might instead have been preserved as a [[study skin]].<ref name="Oxford Dodo 1">{{cite journal|last1=Nowak-Kemp|first1=M.|last2=Hume|first2=J. P.|title=The Oxford Dodo. Part 1: the museum history of the Tradescant Dodo: ownership, displays and audience|journal=Historical Biology|volume=29|issue=2|date=2016|pages=234–247|doi=10.1080/08912963.2016.1152471|s2cid=87191090}}</ref> In 2018, it was reported that scans of the Oxford dodo's head showed that its skin and bone contained [[Shot (pellet)|lead shot]], which was used to hunt birds in the 17th century. This indicates that the Oxford dodo was shot either before being transported to Britain, or some time after arriving. The circumstances of its killing are unknown, and the pellets are to be examined to identify where the lead was mined from.<ref name="Shot">{{cite web |title=Dodo's violent death revealed |url=https://warwick.ac.uk/newsandevents/pressreleases/dodos_violent_death/ |website=warwick.ac.uk |publisher=University of Warwick |access-date=4 October 2023 |date=2018}}</ref><ref>{{cite news|last1=Knapton|first1=Sarah|title=Who shot Lewis Carroll's dodo? Forensic scans reveal mystery death|url=https://www.telegraph.co.uk/science/2018/04/19/shot-lewis-carrolls-dodo-forensic-scans-reveal-mystery-death/ |archive-url=https://ghostarchive.org/archive/20220111/https://www.telegraph.co.uk/science/2018/04/19/shot-lewis-carrolls-dodo-forensic-scans-reveal-mystery-death/ |archive-date=11 January 2022 |url-access=subscription |url-status=live|access-date=20 April 2018|work=The Telegraph|date=20 April 2018}}{{cbignore}}</ref> Many sources state that the Ashmolean Museum burned the stuffed dodo around 1755 because of severe decay, saving only the head and leg. Statute 8 of the museum 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."<ref>{{cite journal| doi = 10.1093/jhc/13.2.125| last = MacGregor | first = A.| year = 2001| title = The Ashmolean as a museum of natural history, 1683 1860| journal = Journal of the History of Collections| volume = 13| issue = 2| pages = 125–144}}</ref> The deliberate destruction of the specimen is now believed to be a myth; it was removed from exhibition to preserve what remained of it. 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 [[tendon]]s. Very few feathers remain on the head. It is probably a female, as the foot is 11% smaller and more gracile than the London foot, yet appears to be fully grown.<ref>{{cite journal | last1 = Hume | first1 = J. P. | author-link1 = Julian Pender Hume | last2 = Datta | first2 = A. | last3 = Martill | first3 = D. M. | year = 2006 | title = Unpublished drawings of the Dodo ''Raphus cucullatus'' and notes on Dodo skin relics | journal = [[Bulletin of the British Ornithologists' Club|Bull. B.O.C.]] | volume = 126A | pages = 49–54 | issn = 0007-1595 | url = http://julianhume.co.uk/wp-content/uploads/2010/07/Hume-et-al-dodo-skin-relics.pdf | access-date = 14 September 2011 | archive-date = 5 November 2019 | archive-url = https://web.archive.org/web/20191105212809/http://julianhume.co.uk/wp-content/uploads/2010/07/Hume-et-al-dodo-skin-relics.pdf | url-status = live }}</ref> The specimen was exhibited at the Oxford museum from at least the 1860s and until 1998, where-after it was mainly kept in storage to prevent damage.<ref name="Oxford Dodo 2">{{cite journal|last1=Nowak-Kemp|first1=M.|last2=Hume|first2=J. P.|title=The Oxford Dodo. Part 2: from curiosity to icon and its role in displays, education and research|journal=Historical Biology|volume=29|issue=3|date=2016|pages=296–307|doi=10.1080/08912963.2016.1155211|s2cid=87966871}}</ref> Casts of the head can today be found in many museums worldwide.<ref name="Oxford Dodo 1"/> {{multiple image | direction = horizontal |align = right |total_width = 350 |image1 = Dodo upper beak in Prague.jpg |alt1 = Fragmentary leg and skull bones of a dodo |image2 = Prague dodo beak.jpg |alt2 = |footer = Upper jaw of a dodo in the [[National Museum of Prague]] (left) and 1855 lithograph of the specimen }} The dried London foot, first mentioned in 1665, and transferred to the British Museum in the 18th century, was displayed next to Savery's ''Edwards's Dodo'' painting until the 1840s, and it too was dissected by Strickland and Melville. It was not posed in a standing posture, which suggests that it was severed from a fresh specimen, not a mounted one. By 1896 it was mentioned as being without its [[integument]]s, and only the bones are believed to remain today, though its present whereabouts are unknown.<ref name=Hume2006/> 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]].{{sfn|Fuller|2002|p=123}} After the castle was occupied by Danish forces in 1702, the museum collection was assimilated into the Royal Danish collection. The skull was rediscovered by J. T. Reinhardt in 1840. Based on its history, it may be the oldest known surviving remains of a dodo brought to Europe in the 17th century.<ref name=Hume2006/> It is {{convert|13|mm|abbr=on}} shorter than the Oxford skull, and may have belonged to a female.<ref name=Livezey1993/> It was [[mummified]], but the skin has perished.<ref name=Hume2017/> The front part of a skull (specimen NMP P6V-004389) in the National Museum of Prague was found in 1850 among the remains of the Böhmisches Museum. Other elements supposedly belonging to this specimen have been listed in the literature, but it appears only the partial skull was ever present (a partial right limb in the museum appears to be from a Rodrigues solitaire).<ref name=Hume2006/><ref>{{cite journal | last = Jiří | first = M. | title = Extinct and nearly extinct birds in the collections of the National Museum, Prague, Czech Republic | journal = Journal of the National Museum (Prague) National History Series | volume = 181 | pages = 105–106 | year =2012 }}</ref>{{sfn|Parish|2013|p=184–188}} It may be what remains of one of the stuffed dodos known to have been at the menagerie of Emperor Rudolph II, possibly the specimen painted by Hoefnagel or Savery there.<ref name=HumeCheke2004>{{cite journal| doi = 10.3366/anh.2004.31.1.57| last1 = Hume| first1 = J. P.| author-link1 = Julian Pender Hume| last2 = Cheke| first2 = A. S.| year = 2004| title = The white dodo of Réunion Island: Unravelling a scientific and historical myth| journal = Archives of Natural History| volume = 31| issue = 1| pages = 57–79| url = http://julianhume.co.uk/wp-content/uploads/2010/07/Hume-and-Cheke-no-illustrations.pdf| access-date = 11 January 2011| archive-date = 5 November 2019| archive-url = https://web.archive.org/web/20191105212750/http://julianhume.co.uk/wp-content/uploads/2010/07/Hume-and-Cheke-no-illustrations.pdf| url-status = live}}</ref> === Subfossil specimens === {{multiple image |align = right |total_width = 350 |image1 = Owen dodo reconstruction.jpg |alt1 = |image2 = Dodo-Skeleton Natural History Museum London England.jpg |alt2 = Brown, mounted dodo skeleton |footer = [[Richard Owen]]'s 1866 reconstruction of the dodo's skeleton (left), based on bones found in the [[Mare aux Songes]] (it is too squat, following Savery's ''Edwards's Dodo'' painting), and his more upright mount at [[Natural History Museum, London]] }} Until 1860, the only known dodo remains were the four incomplete 17th-century specimens. [[Philip Burnard Ayres]] found the first subfossil bones in 1860, which were sent to Richard Owen at the British Museum, who did not publish the findings. In 1863, Owen requested the Mauritian [[Bishop]] [[Vincent Ryan (bishop)|Vincent Ryan]] to spread word that he should be informed if any dodo bones were found.<ref name=HumeCheke2009/> In 1865, George Clark, the government [[schoolmaster]] at [[Mahébourg]], finally found an abundance of subfossil dodo bones in the swamp of Mare aux Songes in Southern Mauritius, after a 30-year search inspired by Strickland and Melville's monograph.<ref name=Hume2006/> 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, feeling for bones with their feet. At first they found few bones, until they cut away herbage that covered the deepest part of the swamp, where they found many fossils.<ref>{{cite journal|last1=Clark|first1=George|title=Account of the late Discovery of Dodos' Remains in the Island of Mauritius|journal=Ibis|volume=8|issue=2|date=April 1866|pages=141–146|issn=0019-1019|doi=10.1111/j.1474-919X.1866.tb06082.x|url=https://www.biodiversitylibrary.org/part/403162 }}</ref> [[Harry Higginson|Harry Pasley Higginson]], a railway engineer from [[Yorkshire]], reports discovering the Mare aux Songes bones at the same time as Clark and there is some dispute over who found them first. Higginson sent boxes of these bones to [[World Museum|Liverpool]], [[Leeds Museums & Galleries|Leeds]] and [[York Museums Trust|York]] museums.<ref>{{Cite journal|last=Brown|first=Clare|date=November 2020|title=Harry Pasley Higginson and his role in the re-discovery of the dodo (''Raphus cucullatus'')|url=https://www.euppublishing.com/doi/abs/10.3366/anh.2020.0662|journal=Archives of Natural History|volume=47|issue=2|pages=381–391|doi=10.3366/anh.2020.0662|s2cid=229463078|via=Edinburgh University Press}}</ref><ref>{{Cite book|last=Higginson|first=Harry|title=Reminiscences of Life and Travel 1859–1872|year=1891}}</ref> The swamp yielded the remains of over 300 dodos, but very few skull and wing bones, possibly because the upper bodies were washed away or scavenged while the lower body was trapped. The situation is similar to many finds of [[moa]] remains in [[New Zealand]] marshes.<ref>{{cite journal | last = Hume | first = J. P. | author-link = Julian Pender Hume | year = 2005 | title = Contrasting taphofacies in ocean island settings: the fossil record of Mascarene vertebrates | journal = Proceedings of the International Symposium "Insular Vertebrate Evolution: The Palaeontological Approach". Monografies de la Societat d'Història Natural de les Balears | volume = 12 | pages = 129–144 }}</ref> Most dodo remains from the Mare aux Songes have a medium to dark brown colouration.<ref name=Meijer2012p177/> 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 bones in ''Memoir on the Dodo'' in October 1866, but erroneously based his reconstruction on the ''Edwards's Dodo'' painting by Savery, making it too squat and obese. In 1869 he received more bones and 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.<ref name=HumeCheke2009/><ref name="Memoir">{{cite book |last1=Owen |first1=R. |last2=Broderip |first2=W. J. |title=Memoir of the dodo (didus ineptus, Linn.) |publisher=Taylor & Francis |date=1866 |location=London |doi=10.5962/bhl.title.110122 |url=https://www.biodiversitylibrary.org/item/193035#page/13/mode/1up |page=52 |access-date=18 December 2020 |archive-date=7 November 2020 |archive-url=https://web.archive.org/web/20201107040340/https://www.biodiversitylibrary.org/item/193035#page/13/mode/1up |url-status=live }}</ref> 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.<ref>{{cite journal| doi = 10.1111/j.1469-7998.1893.tb00001.x| last1 = Newton| first1 = E.| author-link1 = Edward Newton| last2 = Gadow| first2 = H.| year = 1893| title = IX. On additional bones of the Dodo and other extinct birds of Mauritius obtained by Mr. Theodore Sauzier| journal = The Transactions of the Zoological Society of London| volume = 13| issue = 7| pages = 281–302| url = https://www.biodiversitylibrary.org/page/31083700#page/379/mode/1up| access-date = 13 January 2018| archive-date = 11 September 2017| archive-url = https://web.archive.org/web/20170911071559/http://www.biodiversitylibrary.org/page/31083700#page/379/mode/1up| url-status = live}}</ref> {{multiple image |align = left |total_width = 350 |image1 = Raphus cucullatus 1.jpg |alt1 = |caption1 = Skeleton assembled from subfossils found in 2006, [[Naturalis]] |image2 = Ma_-_Raphus_cucullatus_-_GMZ_2.jpg |caption2 = Subfossil bones rediscovered in the [[Grant Museum]] in 2011 |alt2 = }} In 2005, after a hundred years of neglect, a part of the Mare aux Songes swamp was excavated by an international team of researchers (International Dodo Research Project). To prevent [[malaria]], the British had covered the swamp with [[Stone foundation|hard core]] during their rule over Mauritius, which had to be removed. Many remains were found, including bones of at least 17 dodos in various stages of maturity (though no juveniles), and several bones obviously from the skeleton of one individual bird, which have been preserved in their natural position.<ref name=Rijsdijk2011>{{cite journal|doi=10.1177/0959683611405236|display-authors=8|last1=Rijsdijk | first1=K. F.|last2=Zinke | first2=J.|last3=de Louw | first3=P. G. B.|last4=Hume | first4=J. P.|author-link4=Julian Pender Hume|last5=Van Der Plicht | first5=H.|last6=Hooghiemstra | first6=H.|last7=Meijer | first7=H. J. M.|last8=Vonhof | first8=H. B.|last9=Porch | first9=N.|last10=Florens | first10=F. B. V.|last11=Baider | first11=C.|last12=van Geel | first12=B.|last13=Brinkkemper | first13=J.|last14=Vernimmen | first14=T.|last15=Janoo | first15=A.|date=2011|title=Mid-Holocene (4200 kyr BP) mass mortalities in Mauritius (Mascarenes): Insular vertebrates resilient to climatic extremes but vulnerable to human impact|journal=The Holocene|volume=21|issue=8|pages=1179–1194|bibcode=2011Holoc..21.1179R|s2cid=85845297|ref={{sfnRef|Rijsdijk et al.|2011}}}}</ref> 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 genus ''Cylindraspis'', and 7.1% belonged to dodos, which had been deposited within several centuries, 4,000 years ago.<ref>{{cite journal |doi=10.1016/j.quascirev.2008.09.018 |display-authors=8 |last1=Rijsdijk |first1=K. F. |last2=Hume |first2=J. P. |author-link2=Julian Pender Hume |last3=Bunnik |first3=F. |last4=Florens |first4=F. B. V. |last5=Baider |first5=C. |last6=Shapiro |first6=B. |last7=van der Plicht |first7=H. |last8=Janoo |first8=A. |last9=Griffiths |first9=O. |last10=van den Hoek Ostende |first10=L. W. |last11=Cremer |first11=H. |last12=Vernimmen |first12=T. |last13=De Louw |first13=P. |last14=Bholah |first14=A. |last15=Saumtally |first15=S. |last16=Porch |first16=N. |last17=Haile |first17=J. |last18=Buckley |first18=M. |last19=Collins |first19=M. |last20=Gittenberger |first20=E. |date=January 2009 |title=Mid-Holocene vertebrate bone Concentration-Lagerstätte on oceanic island Mauritius provides a window into the ecosystem of the dodo (''Raphus cucullatus'') |journal=Quaternary Science Reviews |volume=28 |issue=1–2 |pages=14–24 |ref={{sfnRef|Rijsdijk et al.|2009}} |bibcode=2009QSRv...28...14R |s2cid=17113275 |url=https://www.rug.nl/research/portal/files/6728502/2009QuatSciRevRijsdijk.pdf |access-date=24 September 2019 |archive-date=24 September 2019 |archive-url=https://web.archive.org/web/20190924114034/https://www.rug.nl/research/portal/files/6728502/2009QuatSciRevRijsdijk.pdf |url-status=live }}</ref> Subsequent excavations suggested that dodos and 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.<ref name=Rijsdijk2011/> Furthermore, [[cyanobacteria]] thrived in the conditions created by the excrements of animals gathered around the swamp, which died of intoxication, dehydration, trampling, and miring.<ref>{{cite journal | doi = 10.1177/0959683614567886| title = A deadly cocktail: How a drought around 4200 cal. Yr BP caused mass mortality events at the infamous 'dodo swamp' in Mauritius| journal = The Holocene| volume = 25| issue = 5| pages = 758–771 | year = 2015| last1 = De Boer | first1 = E. J.| last2 = Velez | first2 = M. I.| last3 = Rijsdijk | first3 = K. F.| last4 = De Louw | first4 = P. G.| last5 = Vernimmen | first5 = T. J.| last6 = Visser | first6 = P. M.| last7 = Tjallingii | first7 = R.| last8 = Hooghiemstra | first8 = H.| bibcode = 2015Holoc..25..758D| s2cid = 128763840}}</ref> Though many small skeletal elements were found during the recent excavations of the swamp, few were found during the 19th century, probably owing to the employment of less refined methods when collecting.<ref name=Meijer2012p177/> [[File:A Unique complete Skeleton of the Dodo, Louis Etienne Thirioux, 1904.jpg|thumb|Complete Skeleton found by Louis Etienne Thirioux in 1904, [[Natural History Museum, Port Louis]]]] Louis Étienne Thirioux, an amateur naturalist at Port Louis, also found many dodo remains around 1900 from several locations. They included the first articulated specimen, which is the first subfossil dodo skeleton found outside the Mare aux Songes, and the only remains of a juvenile specimen, a now lost [[tarsometatarsus]].<ref name=Hume2006/><ref name=Hume2017/> The former specimen was found in 1904 in a cave near [[Le Pouce]] mountain, and is the only known complete skeleton of an individual dodo. Thirioux donated the specimen to the Museum Desjardins (now [[Natural History Museum, Port Louis]]).<ref>{{cite book |last1=Gillespie |first1=Rosemary G. |last2=Clague |first2=David A. |title=Encyclopedia of Islands |date=2009 |publisher=University of California Press |location=Berkeley (US) |isbn=978-0-520-25649-1 |url=https://books.google.com/books?id=g9ZogGs_fz8C |access-date=14 March 2016 |archive-date=15 March 2023 |archive-url=https://web.archive.org/web/20230315014511/https://books.google.com/books?id=g9ZogGs_fz8C |url-status=live }} p. 231.</ref><ref>{{cite book |last1=Richards |first1=Alexandra |title=Mauritius, Rodrigues, Réunion |date=2012 |publisher=Bradt Travel Guides |location=Chalfont St. Peter, Bucks, UK |isbn=978-1-84162-410-5 |edition=8th |url=https://books.google.com/books?id=JnIb2OCzDvEC }} p. 15.</ref> Thrioux's heirs sold a second mounted composite skeleton (composed of at least two skeletons, with a mainly reconstructed skull) to the [[Durban Museum of Natural Science]] in South Africa in 1918. Together, these two skeletons represent the most completely known dodo remains, including bone elements previously unrecorded (such as knee-caps and wing bones). Though some contemporary writers noted the importance of Thrioux's specimens, they were not scientifically studied, and were largely forgotten until 2011, when sought out by a group of researchers. The mounted skeletons were [[laser scanned]], from which [[3-D computer graphics|3-D]] models were reconstructed, which became the basis of a 2016 monograph about the [[osteology]] of the dodo.<ref>{{cite journal|last1=Claessens|first1=L. P. A. M.|last2=Hume|first2=J. P. |title=Provenance and history of the Thirioux dodos|journal=Journal of Vertebrate Paleontology |date=2016 |volume=35 |issue=sup 1 |pages=21–28 |doi=10.1080/02724634.2015.1111896|s2cid=87212166}}</ref><ref>{{cite journal|last1=Claessens|first1=L. P. A. M.|last2=Meijer|first2=H. J. M. |last3=Hume|first3=J. P. |last4=Rijsdijk|first4=K. F. |title=Anatomy of the Dodo (''Raphus cucullatus'' L., 1758): An Osteological Study of the Thirioux Specimens: Preface |journal=Journal of Vertebrate Paleontology |date=2016 |volume=35 |issue=sup 1 |pages=1–2 |doi=10.1080/02724634.2015.1127721|s2cid=220413197}}</ref> In 2006, explorers discovered a complete skeleton of a dodo in a lava cave in Mauritius. This was only the second associated skeleton of an individual specimen ever found, and the only one in recent times.<ref>{{cite journal|last1=Middleton|first1=G. J.|last2=Hume|first2=J. P.|title=The discovery of a Dodo ''Raphus cucullatus'' Linn. (Aves, Columbiformes) in a highland Mauritian lava cave|journal=[[Helictite (journal)|Helictite]]|date=2016|volume=42|pages=13–20|url=https://helictite.caves.org.au/pdf1/42.Middleton-Hume.pdf}}</ref> 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 skeletons, assembled from the dissociated subfossil remains of several individuals.{{sfn|Fuller|2002|pp=123–129}} In 2011, a wooden box containing dodo bones from the [[Edwardian era]] was rediscovered at the [[Grant Museum]] at [[University College London]] during preparations for a move. They had been stored with crocodile bones until then.<ref>{{cite news | last = Kennedy | first = M. | title = Half a Dodo found in museum drawer | work = The Guardian | url = https://www.theguardian.com/science/2011/feb/21/dodo-found-grant-museum-drawer | access-date = 12 May 2012 | location = London | date = 21 February 2011 | archive-date = 4 April 2020 | archive-url = https://web.archive.org/web/20200404181557/https://www.theguardian.com/science/2011/feb/21/dodo-found-grant-museum-drawer | url-status = live }}</ref>
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