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==Taxonomy== [[File:Pigeon-of-passage.jpg|thumb|left|Earliest published illustration of the species (a male), [[Mark Catesby]], 1731]] Swedish naturalist [[Carl Linnaeus]] coined the binomial name ''[[Columba (genus)|Columba]] macroura'' for both the [[mourning dove]] and the passenger pigeon in the 1758 edition of his work ''[[Systema Naturae]]'' (the starting point of [[biological nomenclature]]), wherein he appears to have considered the two identical. This composite description cited accounts of these birds in two pre-Linnean books. One of these was [[Mark Catesby]]'s description of the passenger pigeon, which was published in his 1731 to 1743 work ''Natural History of Carolina, Florida and the Bahama Islands'', which referred to this bird as ''Palumbus migratorius'', and was accompanied by the earliest published illustration of the species. Catesby's description was combined with the 1743 description of the mourning dove by [[George Edwards (naturalist)|George Edwards]], who used the name ''C. macroura'' for that bird. There is nothing to suggest Linnaeus ever saw specimens of these birds himself, and his description is thought to be fully derivative of these earlier accounts and their illustrations. In his 1766 edition of ''Systema Naturae'', Linnaeus dropped the name ''C. macroura'', and instead used the name ''C. migratoria'' for the passenger pigeon, and ''C. carolinensis'' for the mourning dove.<ref>{{cite book |last1=Catesby |first1=M. |title=Natural History of Carolina, Florida and the Bahama Islands |year=1729 |location=London |volume=1 |publisher=W. Innys and R. Manby |page=23 |url=https://archive.org/stream/naturalhistoryCC1V1Cate#page/n137/mode/2up}}</ref><ref name="Mourning Dove 1993">{{cite book |last1=Aldrich |first1=J. W. |year=1993 |page=48 |chapter=Classification and Distribution |title=Ecology and management of the Mourning Dove |editor=Baskett, T.S. |editor2=Sayre, M.W. |editor3=Tomlinson, R.E. |editor4=Mirarchi, R.E. |publisher=[[Stackpole Books]] |isbn=978-0-8117-1940-7 |location=Harrisburg, PA}}</ref><ref name="Hemming 1952">{{cite journal |last=Hemming |first=F. |title=Proposed use of the plenary powers to secure that the name ''Columba migratoria'' Linnaeus, 1766, shall be the oldest available name for the Passenger Pigeon, the type species of the genus ''Ectopistes'' Swainson, 1827 |journal=[[Bulletin of Zoological Nomenclature]] |volume=9 |pages=80β84 |year=1952 |doi=10.5962/bhl.part.10238 |doi-access=free |url=https://www.biodiversitylibrary.org/page/12158465#page/152/mode/1up}}</ref> In the same edition, Linnaeus also named ''C. canadensis'', based on ''Turtur canadensis'', as used by [[Mathurin Jacques Brisson]] in 1760. Brisson's description was later shown to have been based on a female passenger pigeon.<ref name="Oberholser 1918"/> In 1827, [[William Swainson]] moved the passenger pigeon from the genus ''Columba'' to the new [[monotypic]] genus ''Ectopistes'', due in part to the length of the wings and the wedge shape of the tail.<ref>{{cite journal |last=Swainson |first=W. |title=Mr. Swainson on several new groups in Ornithology |journal=[[The Zoological Journal]] |volume=3 |page=362 |year=1827 |url=https://archive.org/stream/zoologicaljourna03lond#page/362/mode/2up}}</ref> In 1906 [[Outram Bangs]] suggested that because Linnaeus had wholly copied Catesby's text when coining ''C. macroura'', this name should apply to the passenger pigeon, as ''E. macroura''.<ref>{{cite journal |last=Bangs |first=O. |title=The names of the passenger pigeon and the mourning dove |journal=[[Proceedings of the Biological Society of Washington]] |volume=19 |pages=43{{ndash}}44 |year=1906 |url=https://www.biodiversitylibrary.org/page/2348105#page/67/mode/1up}}</ref> In 1918 [[Harry C. Oberholser]] suggested that ''C. canadensis'' should take precedence over ''C. migratoria'' (as ''E. canadensis''), as it appeared on an earlier page in Linnaeus' book.<ref name="Oberholser 1918">{{cite journal |last=Oberholser |first=H. C. |title=The scientific name of the Passenger Pigeon |journal=Science |volume=48 |issue=1244 |page=445 |year=1918 |bibcode=1918Sci....48..445O |pmid=17752099 |doi=10.1126/science.48.1244.445 |s2cid=28529631 |url=https://archive.org/stream/jstor-1641492/1641492#page/n1/mode/2up}}</ref> In 1952 [[Francis Hemming]] proposed that the [[International Commission on Zoological Nomenclature]] (ICZN) secure the specific name ''macroura'' for the mourning dove, and the name ''migratorius'' for the passenger pigeon, since this was the intended use by the authors on whose work Linnaeus had based his description.<ref name="Hemming 1952"/> This was accepted by the ICZN, which used its [[plenary powers]] to designate the species for the respective names in 1955.<ref>{{cite book |last1=Schenk |first1=E. T. |last2=McMasters |first2=J. H. |year=1956 |page=89 |title=Procedure in Taxonomy |edition=Third |publisher=[[Stanford University Press]] |isbn=978-0-8047-3867-5 |location=Stanford, California}}</ref> ===Evolution=== [[File:Ectopistes migratorius (passenger pigeon).jpg|thumb|Mounted male passenger pigeon, [[Field Museum of Natural History]]]] [[File:Patagioenas fasciata -San Luis Obispo, California, USA-8 (1).jpg|thumb|[[Band-tailed pigeon]], a species in the related genus ''[[Patagioenas ]]'']] [[File:Zenaida macroura -California-8-2c.jpg|thumb|The physically similar [[mourning dove]] is not closely related.]] The passenger pigeon was a member of the pigeon and dove family, [[Columbidae]]. The oldest known fossil of the genus is an isolated humerus (USNM 430960) known from the Lee Creek Mine in North Carolina in sediments belonging to the [[Yorktown Formation]], dating to the [[Zanclean]] stage of the [[Pliocene]], between 5.3 and 3.6 million years ago.<ref>{{cite journal |last1=Ray |first1=Clayton E. |last2=Bohaska |first2=David J. |title=Miocene and Pliocene birds from the Lee Creek Mine, North Carolina in Geology and Paleontology of the Lee Creek Mine, North Carolina, III |date=2001 |journal=Smithsonian Contributions to Paleobiology |volume=90 |language=en |issue=90 |pages=299β300 |doi=10.5479/si.00810266.90.1 |hdl=10088/2006 |url=https://repository.si.edu/handle/10088/2006}}</ref> Its closest living relatives were long thought to be the [[Zenaida doves|''Zenaida'' doves]], based on morphological grounds, particularly the physically similar mourning dove (now ''Z. macroura'').<ref name="Blockstein 4"/><ref>{{cite conference |last=Miller |first=W. J. |title=The Biology and Natural History of the Mourning Dove |book-title=Should Doves be Hunted in Iowa? |publisher=Ames [[Audubon Society]] |date=January 16, 1969 |location=Ames, IA |url=http://www.ringneckdove.com/Wilmer's%20WebPage/mourning__doves.htm |access-date=April 23, 2013 |url-status=dead |archive-url=https://web.archive.org/web/20120920051011/http://www.ringneckdove.com/Wilmer%27s%20WebPage/mourning__doves.htm |archive-date=September 20, 2012}}</ref> It was even suggested that the mourning dove belonged to the genus ''Ectopistes'' and was listed as ''E. carolinensis'' by some authors, including [[Thomas Mayo Brewer]].<ref name="Brewer 717">{{cite book |last=Brewer |first=T. M. |author-link=Thomas Mayo Brewer |title=Wilson's American Ornithology: with Notes by Jardine; to which is Added a Synopsis of American Birds, Including those Described by Bonaparte, Audubon, Nuttall, and Richardson |publisher=Otis, Broaders, and Company |year=1840 |location=Boston |page=[https://archive.org/details/wilsonsamericano00wils/page/717 717] |url=https://archive.org/details/wilsonsamericano00wils}}</ref> The passenger pigeon was supposedly descended from ''Zenaida'' pigeons that had adapted to the woodlands on the plains of central North America.<ref name="Extinct Birds"/> The passenger pigeon differed from the species in the genus ''Zenaida'' in being larger, lacking a facial stripe, being [[sexually dimorphic]], and having [[iridescent]] neck feathers and a smaller [[clutch (eggs)|clutch]]. In a 2002 study by American geneticist [[Beth Shapiro]] et al., museum specimens of the passenger pigeon were included in an [[ancient DNA]] analysis for the first time (in a paper focusing mainly on the [[dodo]]), and it was found to be the [[sister taxon]] of the cuckoo-dove genus ''[[Macropygia]]''. The ''Zenaida'' doves were instead shown to be related to the quail-doves of the genus ''[[Geotrygon]]'' and the ''[[Leptotila]]'' doves.<ref>{{cite journal |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. |year=2002 |title=Flight of the Dodo |journal=Science |volume=295 |issue=5560 |page=1683 |doi=10.1126/science.295.5560.1683 |pmid=11872833 |ref={{sfnRef |Shapiro et al. |2002 |pmid=11872833 |doi=10.1126/science.295.5560.1683 |url=https://pgl.soe.ucsc.edu/dodo_Shapiro02.pdf}}}}</ref><ref name="Fulton"/><ref name=Johnson/> A more extensive 2010 study instead showed that the passenger pigeon was most closely related to the New World ''[[Patagioenas]]'' pigeons, including the [[band-tailed pigeon]] (''P. fasciata'') of western North America, which are related to the Southeast Asian species in the genera ''[[Turacoena]]'', ''Macropygia'', and ''[[Reinwardtoena]]''. This [[clade]] is also related to the ''Columba'' and ''[[Streptopelia]]'' doves of the Old World (collectively termed the "typical pigeons and doves"). The authors of the study suggested that the ancestors of the passenger pigeon may have colonized the New World from South East Asia by flying across the [[Pacific Ocean]], or perhaps across [[Beringia]] in the north.<ref name=Johnson>{{cite journal |last=Johnson |first=K. P. |author2=Clayton, D. H. |author3=Dumbacher, J. P. |author4=Fleischer, R. C. |title=The flight of the Passenger Pigeon: phylogenetics and biogeographic history of an extinct species |journal=Molecular Phylogenetics and Evolution |volume=57 |issue=1 |pages=455β8 |year=2010 |bibcode=2010MolPE..57..455J |pmid=20478386 |doi=10.1016/j.ympev.2010.05.010}}</ref> In a 2012 study, the [[nuclear DNA]] of the passenger pigeon was analyzed for the first time, and its relationship with the ''Patagioenas'' pigeon was confirmed. In contrast to the 2010 study, these authors suggested that their results could indicate that the ancestors of the passenger pigeon and its Old World relatives may have originated in the [[Neotropical region]] of the New World.<ref name="Fulton">{{cite journal |last=Fulton |first=T. L. |author2=Wagner, S. M. |author3=Fisher, C. |author4=Shapiro, B. |title=Nuclear DNA from the Extinct Passenger Pigeon (''Ectopistes migratorius'') Confirms a Single Origin of New World Pigeons |journal=[[Annals of Anatomy]] |volume=194 |issue=1 |pages=52β7 |year=2012 |bibcode=2012AnAAA.194...52F |pmid=21482085 |doi=10.1016/j.aanat.2011.02.017}}</ref> The [[cladogram]] below follows the 2012 DNA study showing the position of the passenger pigeon among its closest relatives:<ref name="Fulton"/> {{clade |style=font-size:100%; line-height:100% |1={{clade |1={{clade |1={{clade |1={{clade |1={{clade |1=''[[Macropygia]]'' (cuckoo-doves) |2=''[[Reinwardtoena]]''}} |2=''[[Turacoena]]''}} |2={{clade |1=''[[Columba (genus)|Columba]]'' (Old World pigeons) |2=''[[Streptopelia]]'' (turtle doves and collared doves)}}}} |2={{clade |1=''[[Patagioenas]]'' (New World pigeons) |2='''''Ectopistes''''' ('''passenger pigeon''')}}}}}}}} DNA in old museum specimens is often degraded and fragmentary, and passenger pigeon specimens have been used in various studies to discover improved methods of analyzing and assembling genomes from such material. DNA samples are often taken from the toe pads of bird skins in museums, as this can be done without causing significant damage to valuable specimens.<ref>{{cite book |chapter=Case Study: Recovery of Ancient Nuclear DNA from Toe Pads of the Extinct Passenger Pigeon |title=Ancient DNA |volume=840 |pages=29β35 |series=Methods in Molecular Biology |year=2012 |last1=Fulton |first1=T. L. |last2=Wagner |first2=S. M. |last3=Shapiro |first3=B. |isbn=978-1-61779-515-2 |pmid=22237518 |doi=10.1007/978-1-61779-516-9_4}}</ref><ref>{{cite journal |title=The De Novo Assembly of Mitochondrial Genomes of the Extinct Passenger Pigeon (''Ectopistes migratorius'') with Next Generation Sequencing |journal=[[PLOS One]] |volume=8 |issue=2 |pages=e56301 |year=2013 |last1=Hung |first1=C. M. |last2=Lin |first2=R. C. |last3=Chu |first3=J. H. |last4=Yeh |first4=C. F. |last5=Yao |first5=C. J. |last6=Li |first6=S. H. |bibcode=2013PLoSO...856301H |pmid=23437111 |doi=10.1371/journal.pone.0056301 |doi-access=free |pmc=3577829}}</ref> The passenger pigeon had no known subspecies.<ref name="Blockstein 4">{{Harvnb|Blockstein|2002|p=4}}</ref> [[Hybrid (biology)|Hybridization]] occurred between the passenger pigeon and the [[Barbary dove]] (''Streptopelia risoria'') in the aviary of [[Charles Otis Whitman]] (who owned many of the last captive birds around the turn of the 20th century, and kept them with other pigeon species) but the offspring were infertile.<ref name="Extinct Birds">{{cite book |last1=Hume |first1=J. P. |author-link=Julian Pender Hume |last2=Walters |first2=M. |title=Extinct Birds |publisher=T & AD Poyser |year=2012 |location=London |isbn=978-1-4081-5725-1 |pages=144{{ndash}}146}}</ref><ref>{{cite journal |jstor=4070695 |last=Deane |first=R. |title=The Passenger Pigeon (Ectopistes migratorius) in Confinement |journal=The Auk |volume=25 |issue=2 |pages=181β183 |year=1908 |doi=10.2307/4070695 |url=https://archive.org/stream/jstor-4070695/4070695#page/n1/mode/2up}}</ref> ===Etymology=== The genus name, ''Ectopistes'', translates as "moving about" or "wandering", while the specific name, ''migratorius'', indicates its [[bird migration|migratory]] habits.<ref name="Atkinson">{{cite book |last=Atkinson |first=G. E. |title=The Passenger Pigeon |editor=Mershon, W. B |publisher=The Outing Publishing Co |location=New York |year=1907 |page=188 |url=https://en.wikisource.org/wiki/Page:The_Passenger_Pigeon_-_Mershon.djvu/221}}</ref> The full binomial can thus be translated as "migratory wanderer". The English common name "passenger pigeon" derives from the French word ''{{lang|fr|passager}}'', which means "to pass by" in a fleeting manner.<ref name="Fuller 2014 30β47"/><ref>{{cite web |last=Harper |first=D. |title=Passenger (n.) |website=Online Etymology Dictionary |year=2012 |url=https://www.etymonline.com/word/passenger |access-date=April 23, 2013}}</ref> While the pigeon was extant, the name "passenger pigeon" was used interchangeably with "wild pigeon".<ref name="Schorger251">{{cite book |last1=Schorger |first1=A. W. |author-link=Arlie W. Schorger |title=The Passenger Pigeon: Its Natural History and Extinction |publisher=[[University of Wisconsin Press]] |year=1955 |location=Madison, WI |isbn=978-1-930665-96-5}} p. 251.<!--Please keep page number outside the cite template so that sfns with other page numbers can use the full citation. --></ref> The bird also gained some less-frequently used names, including blue pigeon, merne rouck pigeon, wandering long-tailed dove, and wood pigeon. In the 18th century, the passenger pigeon was known as ''tourte'' in [[New France]] (in modern Canada), but to the French in Europe it was known as ''tourtre''. In modern French, the bird is known as ''tourte voyageuse'' or ''pigeon migrateur'', among other names.<ref name="Schorger2523">{{Harvnb|Schorger|1955|pp=252β253}}</ref> In the Native American [[Algonquian languages]], the pigeon was called ''amimi'' by the [[Lenape language|Lenape]], ''{{lang|oj|omiimii}}'' by the [[Anishinaabe language|Ojibwe]], and ''{{lang|mia|mimia}}'' by the [[Kaskaskia]] [[Miami-Illinois language|Illinois]].<ref>{{cite web |last=Tatum |first=J. |author2=Rementer, J. |author3=the Culture Preservation Committee |title=Extinct Birds the Lenape Knew |website=Culture and History of the Delaware Tribe |publisher=[[Delaware Tribe of Indians]] |year=2010 |url=https://delawaretribe.org/blog/2013/06/27/extinct-birds-the-lenape-knew/ |access-date=October 8, 2024 |archive-url=https://web.archive.org/web/20121006163804/http://culture.delawaretribe.org/extinctbirds.htm |archive-date=October 6, 2012}}</ref><ref>{{cite web |title=Omiimii |website=Ojibwe People's Dictionary |publisher=Department of American Indian Studies, [[University of Minnesota]] |url=https://ojibwe.lib.umn.edu/main-entry/omiimii-na |access-date=March 2, 2012}}</ref><ref name="Costa">{{cite web |last1=Costa |first1=D. J. |title=The St. JΓ©rΓ΄me Dictionary of Miami-Illinois |website=Papers of the 36th Algonquian Conference |editor-first=H. C. |editor-last=Wolfart |location=Winnipeg |publisher=[[University of Manitoba]] |year=2005 |pages=107{{ndash}}133 |url=http://myaamiacenter.org/MCResources/costa_biblio/st_jerome_dictionary_costa.pdf |access-date=April 23, 2013 |url-status=dead |archive-url=https://web.archive.org/web/20160611143937/http://myaamiacenter.org/MCResources/costa_biblio/st_jerome_dictionary_costa.pdf |archive-date=June 11, 2016}}</ref> Other names in [[Indigenous peoples of the Americas|indigenous American]] languages include ''{{lang|moh|ori'te}}'' in [[Mohawk language|Mohawk]], and ''{{lang|cho|putchee nashoba}}'', or "lost dove", in [[Choctaw language|Choctaw]].<ref name="Schorger255">{{Harvnb|Schorger|1955|p=255}}</ref> The [[Seneca people]] called the pigeon ''{{lang|see|jahgowa}}'', meaning "big bread", as it was a source of food for their tribes.<ref name="Fuller 2014 72β88"/> Chief [[Simon Pokagon]] of the [[Potawatomi]] stated that his people called the pigeon ''{{lang|pot|O-me-me-wog}}'', and that the Europeans did not adopt native names for the bird, as it reminded them of their domesticated pigeons, instead calling them "wild" pigeons, as they called the native peoples "wild" men.<ref name="Fuller 2014 150β161">{{Harvnb|Fuller|2014|pp=150β161}}</ref>
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