Editing Dodo (section)

=== Evolution ===
[[File:NICOBAR PIGEON (8551073077).jpg|thumb|The [[Nicobar pigeon]] is the closest living relative of the dodo]]

In 2002, American geneticist [[Beth Shapiro]] and colleagues analysed the DNA of the dodo for the first time. Comparison of [[mitochondria]]l [[cytochrome b|cytochrome ''b'']] and 12S [[ribosomal RNA|rRNA]] [[DNA sequence|sequences]] isolated from a [[tarsus (skeleton)|tarsal]] of the Oxford specimen and a [[femur]] of a Rodrigues solitaire confirmed their close relationship and their placement within the Columbidae. The genetic evidence was interpreted as showing the Southeast Asian [[Nicobar pigeon]] (''Caloenas nicobarica'') to be their closest living relative, followed by the [[crowned pigeon]]s (''Goura'') of [[New Guinea]], and the superficially dodo-like [[tooth-billed pigeon]] (''Didunculus strigirostris'') from [[Samoa]] (its scientific name refers to its dodo-like beak). This [[clade]] consists of generally ground-dwelling island endemic pigeons. The following [[cladogram]] shows the dodo's closest relationships within the Columbidae, based on Shapiro and colleagues, 2002:<ref>{{cite journal| doi = 10.1126/science.295.5560.1683| last1 = Shapiro| first1 = B.| last2 = Sibthorpe| first2 = D.| last3 = Rambaut| first3 = A.| last4 = Austin| first4 = J.| last5 = Wragg| first5 = G. M.| last6 = Bininda-Emonds| first6 = O. R. P.| last7 = Lee| first7 = P. L. M.| last8 = Cooper| first8 = A.| date = 2002| title = Flight of the Dodo| journal = Science| volume = 295| issue = 5560| page = 1683| pmid = 11872833| url = http://pgl.soe.ucsc.edu/dodo_Shapiro02.pdf| ref = {{sfnRef|Shapiro et al.|2002}}| access-date = 28 August 2015| archive-date = 20 November 2018| archive-url = https://web.archive.org/web/20181120150854/https://pgl.soe.ucsc.edu/dodo_Shapiro02.pdf| url-status = live}} [http://www.sciencemag.org/cgi/content/full/295/5560/1683/DC1 Supplementary information] {{Webarchive|url=https://web.archive.org/web/20100405060329/http://www.sciencemag.org/cgi/content/full/295/5560/1683/DC1 |date=5 April 2010 }}</ref><ref>{{cite news | author = BBC | date = 28 February 2002 | title = DNA yields dodo family secrets | work = [[BBC News]] | location = London | url = http://news.bbc.co.uk/2/hi/science/nature/1847431.stm | access-date = 7 September 2006 | archive-date = 20 October 2002 | archive-url = https://web.archive.org/web/20021020210432/http://news.bbc.co.uk/2/hi/science/nature/1847431.stm | url-status = live }}</ref>

{{clade |style = font-size: 100%;
|1={{clade
   |1=''[[Didunculus strigirostris]]'' (tooth-billed pigeon)
   |2={{clade
      |1=''[[Goura victoria]]'' (Victoria crowned pigeon)
      |2={{clade
         |1={{clade
            |1={{clade
               |1=''[[Caloenas nicobarica]]'' (Nicobar pigeon)
               |2={{clade
                  |1={{extinct}}''[[Pezophaps solitaria]]'' (Rodrigues solitaire)
                  |2={{extinct}}'''''Raphus cucullatus''''' ('''dodo''')
                  }}
               }}
            }}
         }}
      }}
   }}
}}

[[File:Oxford Dodo foot.jpg|thumb|right|1848 [[lithograph]] of the Oxford specimen's foot, which has been used to sample [[DNA]] for [[genetic analyses]]]]
A similar cladogram was published in 2007, inverting the placement of ''Goura'' and ''Didunculus'' and including the [[pheasant pigeon]] (''Otidiphaps nobilis'') and the [[thick-billed ground pigeon]] (''Trugon terrestris'') at the base of the clade.<ref>{{cite journal| doi = 10.1080/10635150701549672| last1 = Pereira | first1 = S. L.| last2 = Johnson | first2 = K. P.| last3 = Clayton | first3 = D. H.| last4 = Baker | first4 = A. J.| year = 2007| title = Mitochondrial and nuclear DNA sequences support a Cretaceous origin of Columbiformes and a dispersal-driven radiation in the Paleogene| journal = Systematic Biology| volume = 56| issue = 4| pages = 656–672| pmid = 17661233| ref = {{sfnRef|Pereira et al.|2007}} | doi-access = free}}</ref> The DNA used in these studies was obtained from the Oxford specimen, and since this material is degraded, and no usable DNA has been extracted from subfossil remains, these findings still need to be independently verified.<ref name=Hume2012>{{cite journal| doi = 10.1111/j.1365-2451.2012.00843.x| last = Hume | first = J. P.| author-link = Julian Pender Hume| year = 2012| title = The Dodo: From extinction to the fossil record| journal = Geology Today| volume = 28| issue = 4| pages = 147–151| bibcode = 2012GeolT..28..147H | s2cid = 83711229 }}</ref> Based on behavioural and morphological evidence, Jolyon C. Parish proposed that the dodo and Rodrigues solitaire should be placed in the subfamily [[Gourinae]] along with the ''Goura'' pigeons and others, in agreement with the genetic evidence.{{sfn|Parish|2013|pp=134–141}} In 2014, DNA of the only known specimen of the recently extinct [[spotted green pigeon]] (''Caloenas maculata'') was analysed, and it was found to be a close relative of the Nicobar pigeon, and thus also the dodo and Rodrigues solitaire.<ref name="Spotted green pigeon">{{cite journal|last1=Heupink|first1=Tim H|last2=van Grouw|first2=Hein|last3=Lambert|first3=David M|title=The mysterious Spotted Green Pigeon and its relation to the Dodo and its kindred|journal=BMC Evolutionary Biology|volume=14|issue=1|year=2014|page=136|doi=10.1186/1471-2148-14-136|pmid=25027719|pmc=4099497 |doi-access=free |bibcode=2014BMCEE..14..136H }}</ref>

The 2002 study indicated that the ancestors of the dodo and the solitaire diverged around the [[Paleogene]]-[[Neogene]] boundary, about 23.03&nbsp;million years ago. The [[Mascarene Islands]] (Mauritius, [[Réunion]], and [[Rodrigues]]), are of [[volcano|volcanic]] origin and are less than 10&nbsp;million years old. Therefore, the ancestors of both birds probably remained capable of flight for a considerable time after the separation of their [[lineage (evolution)|lineage]].<ref>{{cite book | last1 = Cheke | first1 = Anthony S. | last2 = Hume | first2 = Julian Pender | year = 2008 | title = Lost Land of the Dodo: an Ecological History of Mauritius, Réunion & Rodrigues | publisher = T. & A. D. Poyser | location = New Haven and London | isbn = 978-0-7136-6544-4 }} pp.&nbsp;70{{ndash}}71.<!--Please keep page number(s) outside the cite template so that sfns with other page numbers can use the full citations. --></ref> The Nicobar and spotted green pigeon were placed at the base of a lineage leading to the Raphinae, which indicates the [[flightless]] raphines had ancestors that were able to fly, were semi-terrestrial, and inhabited islands. This in turn supports the hypothesis that the ancestors of those birds reached the Mascarene islands by [[island hopping]] from South Asia.<ref name="Spotted green pigeon" /> The lack of [[mammal]]ian [[herbivore]]s competing for resources on these islands allowed the solitaire and the dodo to attain [[island gigantism|very large sizes]] and flightlessness.<ref>{{cite journal| doi = 10.1086/316701| last = McNab | first = B. K.| year = 1999| title = On the Comparative Ecological and Evolutionary Significance of Total and Mass-Specific Rates of Metabolism| journal = Physiological and Biochemical Zoology| volume = 72| issue = 5| pages = 642–644| jstor = 10.1086/316701| pmid = 10521332| s2cid = 28619917 }}</ref>{{sfn|Fuller|2001|pp=37–39}} Despite its divergent skull morphology and adaptations for larger size, many features of its skeleton remained similar to those of smaller, flying pigeons.<ref name=ClaessensMeijer2016>{{cite journal|last1=Claessens|first1=L. P. A. M.|last2=Meijer|first2=H. J. M. |last3=Hume|first3=J. P. |title=The Morphology of the Thirioux dodos|journal=Journal of Vertebrate Paleontology |date=2016 |volume=35 |issue=sup 1 |pages=29–187 |doi=10.1080/02724634.2015.1121723|s2cid=87957947}}</ref> Another large, flightless pigeon, the [[Viti Levu giant pigeon]] (''Natunaornis gigoura''), was described in 2001 from [[subfossil]] material from [[Fiji]]. It was only slightly smaller than the dodo and the solitaire, and it too is thought to have been related to the crowned pigeons.<ref>{{cite journal| doi = 10.1080/03014223.2001.9517673| last = Worthy | first = T. H.| year = 2001| title = A giant flightless pigeon gen. et sp. nov. and a new species of ''Ducula'' (Aves: Columbidae), from Quaternary deposits in Fiji| journal = Journal of the Royal Society of New Zealand| volume = 31| issue = 4| pages = 763–794| bibcode = 2001JRSNZ..31..763W | s2cid = 83708873 }}</ref>