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Rodrigues solitaire
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==Taxonomy== The French explorer [[François Leguat]] was the first to refer to the bird as the "solitaire" (referring to its solitary habits), but it has been suggested that he borrowed the name from a 1689 [[Tract (literature)|tract]] by his sponsor Marquis Henri Duquesne, which used the name "solitaire" in reference to the [[Réunion ibis]].<ref>{{cite journal |doi= 10.3366/anh.2004.31.1.57 |last1= Hume |first1= J. P. |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 }}</ref> The bird was first scientifically named in 1789 as a [[species]] of dodo (''Didus solitarius'', based on Leguat's description) by the German naturalist [[Johann Friedrich Gmelin]] in the thirteenth edition of ''[[Systema Naturae]]''.<ref name="NewtonNewton1869">{{cite journal |doi= 10.1098/rstl.1869.0011 |last1= Newton |first1= Alfred |last2= Newton |first2= Edward |date= 1 January 1869 |title= On the Osteology of the Solitaire or Didine Bird of the Island of Rodriguez, ''Pezophaps solitaria'' (Gmel) |journal= Philosophical Transactions of the Royal Society of London |volume= 159 |pages= 327–362 |doi-access= free }} {{free access}}</ref> In 1786, subfossil Rodrigues solitaire bones encrusted in [[stalagmite]] were discovered in a cave and sent to the French naturalist [[Georges Cuvier]] in about 1830. For unknown reasons, he stated they had recently been found on [[Mauritius]], which caused confusion, until they were compared with other bones from Rodrigues that were found to belong to the same species.<ref>{{cite journal |doi= 10.1080/08912963.2014.886203 |url= http://julianhume.co.uk/wp-content/uploads/2010/07/Hume-et-al.-Bone-collectors1.pdf |title= In the footsteps of the bone collectors: Nineteenth-century cave exploration on Rodrigues Island, Indian Ocean |journal= Historical Biology |volume= 27 |issue= 2 |pages= 1 |year= 2014 |last1= Hume |first1= J. P. |last2= Steel |first2= L. |last3= André |first3= A. A. |last4= Meunier |first4= A.|bibcode= 2015HBio...27..265H |s2cid= 128901896 }}</ref> The English naturalists [[Hugh Edwin Strickland]] and [[Alexander Gordon Melville]] suggested the common descent of the Rodrigues solitaire and the [[dodo]] in 1848. They dissected the only known dodo specimen with soft tissue, comparing it with the few Rodrigues solitaire remains then available.<ref>{{cite journal |doi= 10.1111/j.1469-7998.1862.tb08059.x |last1= Strickland |first1= H. E. |author-link= Hugh Edwin Strickland |date=August 1859 |title= XVI. On some Bones of Birds allied to the Dodo, in the Collection of the Zoological Society of London |journal= The Transactions of the Zoological Society of London |volume= 4 |issue= 6 |pages= 187–196 |url= https://zenodo.org/record/1447617 }}</ref> Strickland stated that, although not identical, these birds shared many distinguishing features in the leg bones otherwise only known in [[pigeon]]s. The fact that the Rodrigues solitaire laid only one egg, fed on fruits, was [[Monogamy in animals|monogamous]] and cared for its nestlings also supported this relationship. Strickland recognised its generic distinction and named the new [[genus]] ''Pezophaps'', from [[ancient Greek]] ''{{Transliteration|grc|pezos}}'' ({{lang|grc|πεζός}} 'pedestrian') and ''{{Transliteration|grc|phaps}}'' ({{lang|grc|φάψ}} 'pigeon').<ref name="Parish">{{cite book | last= Parish| first= Jolyon C.| year= 2013| title=The Dodo and the Solitaire: A Natural History| publisher=Indiana University Press| url= https://books.google.com/books?id=bp8wK8zCg7wC| isbn= 978-0253000996 | location=Bloomington | pages=140, 265–282, 336–340}}</ref><ref name="StricklandMelville1848p46">{{cite book| last1= Strickland| first1= H. E.| author-link= Hugh Edwin Strickland| last2= Melville| first2= A. G.| title= 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| publisher= Reeve, Benham and Reeve| location= London| year= 1848| url= https://archive.org/details/dodoitskindredor00stri| pages= [https://archive.org/details/dodoitskindredor00stri/page/n73 46]–55}}</ref> The [[sexual dimorphism|differences between the sexes]] of the bird were so large that Strickland thought they belonged to two species, naming the smaller female bird ''Pezophaps minor''.<ref name="NewtonClark1879">{{cite journal |doi= 10.1098/rstl.1879.0044 |last1= Newton |first1= Edward |last2= Clark |first2= John Willis |date= 1 January 1879 |title= On the Osteology of the Solitaire (''Pezophaps solitaria'', Gmel.) |journal= Philosophical Transactions of the Royal Society of London |volume= 168 |pages= 438–451 |bibcode= 1879RSPT..168..438N |doi-access= free }} {{free access}}</ref> [[File:Pezophaps stalagmite.jpg|thumb|left|The first [[stalagmite]]-encrusted remains of this bird known by 1848|alt=Rodrigues solitaire bones]] Additional subfossils were recovered during the 1860s, but more complete remains were found during the [[1874 transit of Venus]], since an observation station was located on the island.<ref name="NewtonClark1879"/> Many of these excavations were requested by the English ornithologists (and brothers) [[Alfred Newton|Alfred]] and [[Edward Newton]], who used them to describe the [[osteology]] of the bird in detail. Thousands of bones were excavated, and mounted skeletons were composed from the remains of several specimens.<ref>{{cite journal |doi= 10.1080/00222937808682294 |last= Owen |first= R. |year= 1878 |url= https://archive.org/details/annalsmagazineof511878lond |title= XII. On the Solitaire (''Didus solitarius'', Gm.; ''Pezophaps solitaria'', Strkl.) |journal= Journal of Natural History|series=Series 5 |volume= 1 |issue= 1 |pages= [https://archive.org/details/annalsmagazineof511878lond/page/87 87]–98 }}</ref> Study of skeletal features by the Newtons indicated that the solitaire was [[Morphology (biology)|morphologically]] intermediate between the dodo and ordinary pigeons, but differed from them in its unique [[carpal]] knob.<ref name="NewtonNewton1869"/> Some scientists believed that [[Réunion]] was home not only to a white dodo, but also to a white bird similar to the Rodrigues solitaire, both of which are now believed to be misinterpretations of old reports of the Réunion ibis.<ref>{{cite journal |doi= 10.1093/jhc/15.2.201 |last1= de Lozoya |first1= A. V. |year= 2003 |url= https://www.academia.edu/35150001 |title= An unnoticed painting of a white Dodo |journal= Journal of the History of Collections |volume= 15 |issue= 2 |pages= 201–210 }}</ref> An atypical 17th-century description of a dodo and bones found on [[Rodrigues]], now known to have belonged to the Rodrigues solitaire, led the British taxidermist [[Abraham Dee Bartlett]] to name a new species, ''Didus nazarenus''; it is now a [[junior synonym]] of this species.<ref>{{cite journal |doi= 10.1111/j.1469-7998.1865.tb02320.x |last= Newton |first= A. |date=January 1865 |url= https://biodiversitylibrary.org/page/28497132 |title= 2. On Some Recently Discovered Bones of the Largest Known Species of Dodo (''Didus Nazarenus'', Bartlett) |journal= Proceedings of the Zoological Society of London |volume= 33 |issue= 1 |pages= 199–201 }}</ref><ref>{{cite book |last= Lydekker |first= R. |year= 1891 |title= Catalogue of the Fossil Birds in the British Museum (Natural History) |publisher= [[Taylor & Francis]] |doi= 10.5962/bhl.title.8301 |oclc= 4170867 |page=[https://biodiversitylibrary.org/page/8389911 128] |url= https://www.biodiversitylibrary.org/item/34492 }}</ref> At one point it was suggested that the skeleton of this species is the best described after that of [[human]]s.<ref name="ChekeHume2008p111">{{cite book| last1= Cheke| first1= A. S.| last2= Hume| first2= J. P.| 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| pages= 111–114}}</ref> In spite of the evidence, some later scholars doubted Leguat's story, and the existence of the Rodrigues solitaire. In 1921, the American linguist Geoffroy Atkinson claimed Leguat's memoir was merely a novel, and that the man had never even existed, and in 1955, the British ecologist [[George Evelyn Hutchinson]] doubted aspects of the bird's biology mentioned by Leguat. Today, it is widely accepted that Leguat's memoirs are credible observations of the bird in life.<ref name="Fuller2002">{{cite book| last= Fuller| first= E.| author-link= Errol Fuller| year= 2002| title= Dodo – From Extinction To Icon| publisher= [[HarperCollins]]| location= London| isbn= 978-0-00-714572-0| pages= 156–164}}</ref><ref name="islandnovel">{{cite journal |last1=Atkinson |first1=G. |title=A French desert island novel of 1708 |journal=PMLA |date=1921 |volume=36 |issue=4 |pages=509–528 |jstor=457349 |language=en|doi=10.2307/457349 |s2cid=163478864 }} {{free access}}</ref><ref>{{cite journal |jstor=27826550 |last= Hutchinson |first= G. E. |year= 1954 |title= MARGINALIA |journal= American Scientist |volume= 42 |issue= 2 |pages= 300–308 }}</ref> For many years the dodo and the Rodrigues solitaire were placed in a [[family (biology)|family]] of their own, the Raphidae (formerly Dididae), because their exact relationships with other pigeons were unresolved. Each was also placed in a [[monotypic taxon|monotypic]] family (Raphidae and Pezophapidae, respectively), as it was thought that they had [[Convergent evolution|evolved their similarities independently]].<ref>{{cite journal |doi= 10.2307/4083934 |last= Storer |first= R. W. |year= 1970 |title= Independent Evolution of the Dodo and the Solitaire |journal= The Auk |volume= 87 |issue= 2 |url=http://sora.unm.edu/node/21993 |pages= 369–370 |jstor= 4083934 }}</ref> [[Osteological]] and [[DNA analysis]] has since led to the dissolution of the family Raphidae, and the dodo and solitaire are now placed in their own subfamily, Raphinae, within the family Columbidae.<ref name="Janoo2005">{{cite journal |doi= 10.1016/j.annpal.2004.12.002 |last= Janoo |first= A. |date=April–June 2005 |url= https://www.academia.edu/3378817 |title= Discovery of Isolated Dodo Bones [''Raphus cucullatus'' (L.), Aves, Columbiformes] from Mauritius Cave Shelters Highlights Human Predation, with a Comment on the Status of the Family Raphidae Wetmore, 1930 |journal= Annales de Paléontologie |volume= 91 |issue= 2 |pages= 167–180 |bibcode= 2005AnPal..91..167J }}</ref> ===Evolution=== In 2002, American geneticist [[Beth Shapiro]] and colleagues analysed the DNA of the dodo and the Rodrigues solitaire for the first time. Comparison of [[mitochondria]]l [[cytochrome b|cytochrome ''b'']] and 12S [[ribosomal RNA|rRNA]] [[DNA sequence|sequences]] isolated from the [[femur]] of a Rodrigues solitaire and the [[tarsus (skeleton)|tarsal]] of a dodo 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]]. This [[clade]] consists of generally ground-dwelling island endemic pigeons. The following [[cladogram]] shows the closest relationships of the dodo and the Rodrigues solitaire within Columbidae, based on Shapiro et al., 2002:<ref name="Shapiroetal2002">{{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 |pages= 1683 |pmid= 11872833 |url= http://pgl.soe.ucsc.edu/dodo_Shapiro02.pdf }} [http://www.sciencemag.org/cgi/content/full/295/5560/1683/DC1 Supplementary information]</ref><ref>{{cite web| work= [[BBC News]]| date= 28 February 2002| title= DNA yields dodo family secrets| url= http://news.bbc.co.uk/2/hi/science/nature/1847431.stm| access-date= 6 December 2006 }}</ref> [[File:NICOBAR PIGEON (8551073077).jpg|thumb|A [[Nicobar pigeon]], the closest living relative of the Rodrigues solitaire and the [[dodo]] according to [[DNA]] studies]] {{clade|style=font-size:100%; |label1= |1={{clade |1=''[[Didunculus strigirostris]]'' (tooth-billed pigeon) |2={{clade |1=''[[Goura victoria]]'' (Victoria crowned pigeon) |2={{clade |1={{clade |label1= |1={{clade |1=''[[Caloenas nicobarica]]'' (Nicobar pigeon) |2={{clade |1={{extinct}}'''''Pezophaps solitaria''''' ('''Rodrigues solitaire''') |2={{extinct}}''[[Raphus cucullatus]]'' (dodo) }} }} }} }} }} }} }} 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 |doi-access= free }} {{free access}}</ref> Based on behavioural and morphological evidence, Jolyon C. Parish proposed that the dodo and Rodrigues solitaire should be placed in the [[Gourinae]] subfamily along with the [[Crowned pigeon|''Goura'']] pigeons and others, in agreement with the genetic evidence<ref name="Parish"/> 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|pages=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 Rodrigues solitaire and the dodo diverged around the [[Paleogene]]–[[Neogene]] boundary. The [[Mascarene Islands]] (Mauritius, Réunion, and Rodrigues), are of [[volcano|volcanic]] origin and are less than 10 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= A. S.| last2= Hume| first2= J. P.| 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| pages= 70–71}}</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]].<ref>{{cite journal |doi= 10.1086/316701 |last= McNab |first= B. K. |year= 1999 |url= https://www.researchgate.net/publication/12779859 |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> The dodo lost the ability to fly owing to the lack of mammalian predators on Mauritius.<ref>{{cite book| last= Fuller| first= E.| author-link= Errol Fuller| year= 2001| title= Extinct Birds| publisher= Comstock| edition= revised| location= New York| isbn= 978-0-8014-3954-4| pages= 37–39}}</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 Rodrigues solitaire and the dodo, 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 }} {{free access}}</ref>
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