Passenger pigeon

The passenger pigeon or wild pigeon (Ectopistes migratorius) is an extinct species of pigeon that was endemic to North America. Its common name is derived from the French word passager, meaning "passing by", due to the migratory habits of the species. The scientific name also refers to its migratory characteristics. The morphologically similar mourning dove (Zenaida macroura) was long thought to be its closest relative, and the two were at times confused, but genetic analysis has shown that the genus Patagioenas is more closely related to it than the Zenaida doves.

The passenger pigeon was sexually dimorphic in size and coloration. The male was 390 to 410 mm (15.4 to 16.1 in) in length, mainly gray on the upperparts, lighter on the underparts, with iridescent bronze feathers on the neck, and black spots on the wings. The female was 380 to 400 mm (15.0 to 15.7 in), and was duller and browner than the male overall. The juvenile was similar to the female, but without iridescence. It mainly inhabited the deciduous forests of eastern North America and was also recorded elsewhere, but bred primarily around the Great Lakes. The pigeon migrated in enormous flocks, constantly searching for food, shelter, and breeding grounds, and was once the most abundant bird in North America, numbering around 3 billion, and possibly up to 5 billion. A very fast flyer, the passenger pigeon could reach a speed of 100 km/h (62 mph). The bird fed mainly on mast, and also fruits and invertebrates. It practiced communal roosting and communal breeding, and its extreme gregariousness may have been linked with searching for food and predator satiation.

Passenger pigeons were hunted by Native Americans, but hunting intensified after the arrival of Europeans, particularly in the 19th century. Pigeon meat was commercialized as cheap food, resulting in hunting on a massive scale for many decades. There were several other factors contributing to the decline and subsequent extinction of the species, including shrinking of the large breeding populations necessary for preservation of the species and widespread deforestation, which destroyed its habitat. A slow decline between about 1800 and 1870 was followed by a rapid decline between 1870 and 1890. In 1900, the last confirmed wild bird was shot in southern Ohio. The last captive birds were divided in three groups around the turn of the 20th century, some of which were photographed alive. Martha, thought to be the last passenger pigeon, died on September 1, 1914, at the Cincinnati Zoo. The eradication of the species is a notable example of anthropogenic extinction.

Swedish naturalist Carl Linnaeus coined the binomial name 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, 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. 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.

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. 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. 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. 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. This was accepted by the ICZN, which used its plenary powers to designate the species for the respective names in 1955.

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. Its closest living relatives were long thought to be the Zenaida doves, based on morphological grounds, particularly the physically similar mourning dove (now Z. macroura). 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. The passenger pigeon was supposedly descended from Zenaida pigeons that had adapted to the woodlands on the plains of central North America.

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. 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.

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.