Columbidae is a birdfamily consisting ofdoves andpigeons. It is the only family in theorderColumbiformes. These are stout-bodied birds with small heads, relatively short necks and slender bills that in some species feature fleshyceres. Theyfeed largely on plant matter, feeding on seeds (granivory), fruit (frugivory), and foliage (folivory).
Incolloquial English, the smaller species tend to be called "doves", and the larger ones "pigeons",[2] although the distinction is not consistent,[2] and there is no scientific separation between them.[3] Historically, the common names for these birds involve a great deal of variation. The bird most commonly referred to as "pigeon" is thedomestic pigeon, descendant of the wildrock dove, which is a commoninhabitant of cities as theferal pigeon.
Pigeon is a French word that derives from theLatinpīpiō, for a'peeping' chick,[6] whiledove is an ultimatelyGermanic word, possibly referring to the bird's diving flight.[7] The English dialectal wordculver appears to derive from Latincolumba.[6] A group of doves has sometimes been called a "dule", taken from the French worddeuil ('mourning').[8]
The name 'Columbidae' for the family was first used by the English zoologistWilliam Elford Leach in a guide to the contents of theBritish Museum published in 1819.[20][21] However, Illiger in 1811 established an older name for the family group ("Columbini") and would actually be the proper authority for Columbidae.[22]
The interrelationships of columbids (between subfamilies) and the ergotaxonomy of them has been debated, with many different interpretations of how they should be classified. As many as five to six families, along with many subfamilies and tribes, have been used in the past including the family Raphidae for the dodo and the Rodrigues solitaire.[23][24][25] A 2024 paper on the systematics and nomenclature of the dodo and the solitaire from Young and colleagues also provided an overview of columbid family-group nomina. They recommended recognizing three subfamilies: Columbinae (New World doves and quail-doves, and columbin doves), Claravinae (American ground-doves), and Raphinae (Old World doves and pigeons including the dodo and solitaire).[22] A 2025 paper on the molecular phylogenetic placement of theCuban endemic blue-headed quail-dove from Oswald and colleagues found the species to be asister group to Columbinae, as opposed to being a true columbine or a raphine as previous authors have suggested in the past. These authors recommended that the blue-headed quail-dove should be placed in fourth monotypic subfamily, Starnoenadinae.[26]
These taxonomic issues are exacerbated by columbids not being wellrepresented in thefossil record,[27] with no trulyprimitive forms having been found to date.[citation needed] The genusGerandia has been described fromEarly Miocene deposits in France, but while it was long believed to be a pigeon,[28] it is now considered asandgrouse.[29] Fragmentary remains of a probably "ptilinopine" Early Miocene pigeon were found in the Bannockburn Formation of New Zealand and described asRupephaps;[29]"Columbina" prattae from roughly contemporary deposits ofFlorida is nowadays tentatively separated inArenicolumba, but its distinction fromColumbina/Scardafella and related genera needs to be more firmly established (e.g. bycladistic analysis).[30] Apart from that, all other fossils belong to extant genera.[31]
Overall, theanatomy of Columbidae is characterized by short legs, short bills with a fleshycere, and small heads on large, compact bodies.[32] Like some other birds, the Columbidae have nogall bladders.[33] Some medieval naturalists concluded they have nobile (gall), which in the medieval theory of thefour humours explained the allegedly sweet disposition of doves.[34] In fact, however, they do have bile (asAristotle had earlier realized), which is secreted directly into thegut.[35]
The wings of most species are large, and have elevenprimary feathers;[36] pigeons have strong wing muscles (wing muscles comprise 31–44% of their body weight[37]) and are among the strongest fliers of all birds.[36]
In a series of experiments in 1975 by Dr.Mark B. Friedman, using doves, their characteristic head bobbing was shown to be due to their natural desire to keep their vision constant.[38] It was shown yet again in a 1978 experiment by Dr.Barrie J. Frost, in which pigeons were placed ontreadmills; it was observed that they did not bob their heads, as their surroundings were constant.[39]
Columbidae have unique bodyfeathers, with the shaft being generally broad, strong, and flattened, tapering to a fine point, abruptly.[36] In general, the aftershaft is absent; however, small ones on some tail and wing feathers may be present.[40] Body feathers have very dense, fluffy bases, are attached loosely into the skin, and drop out easily.[41] Possibly serving as apredator avoidance mechanism,[42] large numbers of feathers fall out in the attacker's mouth if the bird is snatched, facilitating the bird's escape. Theplumage of the family is variable.[43]
Granivorous species tend to have dull plumage, with a few exceptions, whereas thefrugivorous species have brightly coloured plumage.[44] The generaChalcophaps,Ptilinopus andAlectroenas include some of the most brightly coloured pigeons. Pigeons and doves may be sexually monochromatic ordichromatic.[45] In addition to bright colours, some pigeon species may have crests or other ornamentation.[46]
Snow pigeons flying at an altitude of 12,000 feet (3,700 m) above sea level
Flight
Many Columbidae are excellent fliers due to the lift provided by their large wings, which results in lowwing loading;[47] They are highly maneuverable in flight[48] and have a lowaspect ratio due to the width of their wings, allowing for quick flight launches and ability to escape from predators, but at a high energy cost.[49] A few species are long-distancemigrants, with some populations of theEuropean turtle dove migrating in excess of 5,000 km between northern Europe in summer and tropical Africa in winter, and theOriental turtle dove nearly as far in eastern Asia between eastern Siberia and southern China.
Size
Pigeons and doves exhibit considerable variation in size, ranging in length from 15 to 75 centimetres (5.9 to 29.5 in), and in weight from 30 g (0.066 lb) to above 2,000 g (4.4 lb).[50] The largest extant species are thecrowned pigeons ofNew Guinea,[51] which are nearlyturkey-sized, with lengths of 66–79 cm (2.17–2.59 ft) and weights ranging 1.8–4 kg (4.0–8.8 lb).[52][53][54] One of the largestarboreal species, theMarquesan imperial pigeon with a length of 55 cm (22 in), currently battles extinction.[55][56] The extinct, flightlessdodo is the largest columbid to have ever existed, with a height of about 62.6–75 cm (24.6–29.5 in), and a range of suggested weights from 10.2–27.8 kilograms (22–61 lb), although the higher estimates are thought to be based on overweight birds.[57][58][59][60]
The least massive columbids belong to species in the genusColumbina; the common ground dove (Columbina passerina) and the plain-breasted ground dove (Columbina minuta) which are about the same size as ahouse sparrow, weighing a little above 22 g (0.78 oz).[44][61][62] Thedwarf fruit dove, which may measure as little as 13 cm (5.1 in) long, has a marginally smaller total length than any other species from this family.[44]
Diversity of Pigeons and Doves
The Nicobar pigeon (Caloenas nicobarica) is often stated to be the dodo's closest living relative.
Some species have large natural ranges. Theeared dove ranges across the entirety of South America from Colombia toTierra del Fuego,[64] theEurasian collared dove has a massive (if discontinuous) distribution fromBritain across Europe, the Middle East, India, Pakistan and China,[65] and thelaughing dove across most of sub-Saharan Africa, as well as India, Pakistan, and the Middle East.[66]
When including human-mediated introductions, the largest range of any species is that of therock dove, also known as the common pigeon.[67] This species had a large natural distribution from Britain and Ireland to northern Africa, across Europe,Arabia,Central Asia, India, theHimalayas and up into China and Mongolia.[67] The range of the species increased dramatically upon domestication, as the species wentferal in cities around the world.[67] The common pigeon is currently resident across most of North America, and has established itself in cities and urban areas in South America, sub-Saharan Africa,Southeast Asia, Japan, Australia, and New Zealand.[67] A 2020 study found that the east coast of the United States includes two pigeon genetic megacities, in New York and Boston, and observes that the birds do not mix together.[68]
As well as the rock dove, several other species of pigeon have become established outside of their natural range after escaping captivity, and other species have increased their natural ranges due to habitat changes caused by human activity.[44]
Seeds and fruit form the major component of the diets of pigeons and doves,[50][74] and the family can be loosely divided between seed-eating (granivorous) species, and fruit-and-mast-eating (frugivorous) species, though many species consume both.[75]
The granivorous species typically feed on seed found on the ground, whereas the frugivorous species are morearboreal, tending to feed in trees.[75] The morphological adaptations used to distinguish between the two groups include granivores tending to having thick walls in their gizzards,intestines, andesophagi, with the frugivores evolved with thin walls,[50] and the fruit-eating species have short intestines, as opposed to the seed eaters having longer intestines.[76] Frugivores are capable of clinging to branches and even hang upside down to reach fruit.[44][75]
Urban feral pigeons, descendants of domesticrock doves (Columbia livia),reside in urban environments, disturbing their natural feeding habits. They depend on human activities and interactions to obtain food, causing them to forage forspilled food or food provided by humans.[77]
Spotted dove (Spilopelia chinensis) on its nest, with one unhatched egg and one hatchling
Reproduction
Doves and pigeons build relatively flimsynests, often using sticks, other vegetable matter, and other debris, which may be placed on trees, onrocky ledges, or on the ground, depending on species. The female may either build the nest, with material gathered by the male, or the male builds the nest by himself. A few speciesnest colonially, others nest in aggregation.[4]
Most lay aclutch of one or (usually) two white eggs at a time which take 11-30 days to hatch (larger species have longer incubation times). Bothparents care for the young; unlike most birds, both sexes of doves and pigeons produce "crop milk" to feed their young. This fluid is secreted by a sloughing ofepithelial cells from the lining of thecrop.[4]
Unfledged baby doves and pigeons are called squabs and are generally able to fly by five weeks old. Thesefledglings, with their immature squeaking voices, are called squeakers once they areweaned,[78] and leave the nest after 25–32 days.
Status and conservation
While many species of pigeons and doves have benefited from human activities and have increased their ranges, many other species have declined in numbers and some have becomethreatened or even succumbed to extinction.[79] Among the ten species to have become extinct since 1600 (the conventional date for estimatingmodern extinctions) are two of the most famous extinct species, the dodo and the passenger pigeon.[79][4]
Thepassenger pigeon was exceptional for a number of reasons. In modern times, it is the only pigeon species that was not an island species to have become extinct[79] even though it was once the most numerous species of bird on Earth.[citation needed] Its former numbers are difficult to estimate, but oneornithologist,Alexander Wilson, estimated one flock he observed contained over two billion birds.[80] The decline of the species was abrupt; in 1871, a breeding colony was estimated to contain over a hundred million birds, yet the last individual in the species was dead by 1914.[81] Althoughhabitat loss was a contributing factor, the species is thought to have been massivelyover-hunted, being used as food forslaves and, later, the poor, in the United States throughout the 19thcentury.[citation needed]
Thedodo, and its extinction, was more typical of the extinctions of pigeons in general. Like many species that colonise remoteislands with few predators, it lost much of itspredator avoidance behaviour, along with itsability to fly.[82] The arrival of people, along with a suite of other introduced species such asrats,pigs, andcats, quickly spelled the end for this species and many other island species that have become extinct.[82]
118 columbid species are at risk (34% of the total), with 48 speciesNT, 40VU, 18EN, 11CR, and 1EW.[4] Most of these are tropical and live on islands. All of the species are threatened byintroduced predators,habitat loss,hunting, or a combination of these factors.[82] In some cases, they may beextinct in the wild, as is theSocorro dove ofSocorro Island, Mexico, last seen in the wild in 1972, driven to extinction by habitat loss and introducedferal cats.[83] In some areas, a lack of knowledge means the true status of a species is unknown (DD); theNegros fruit dove has not been seen since 1953,[84] and may or may not be extinct, and thePolynesian ground dove is classified ascritically endangered, as whether it survives or not on remote islands in the far west of the Pacific Ocean is unknown.[85]
Theringneck dove is a smaller species of domestic columbid that was kept as a source of food. As a result of selection for tame individuals who would not escape their cages, they lack a survival instinct and cannot survive release.[87]
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^Pereira, S.L.et al. (2007) Mitochondrial and nuclear DNA sequences support a Cretaceous origin of Columbiformes and a dispersal-driven radiation in the Paleocene.Syst Biol. 56:656–72
^Soares, A.E.R.et al. (2016) Complete mitochondrial genomes of living and extinct pigeons revise the timing of the columbiform radiation.BMC Evolutionary Biology, 16(230).
^Braun, Edward L.; Cracraft, Joel; Houde, Peter (2019). "Resolving the Avian Tree of Life from Top to Bottom: The Promise and Potential Boundaries of the Phylogenomic Era".Avian Genomics in Ecology and Evolution. pp. 151–210.doi:10.1007/978-3-030-16477-5_6.ISBN978-3-030-16476-8.S2CID198399272.
^Leach, William Elford (1819)."Eleventh Room".Synopsis of the Contents of the British Museum (15th ed.). London: British Museum. pp. 63–68 [66]. Although the name of the author is not specified in the document, Leach was the Keeper of Zoology at the time.
^Janoo, Anwar (2005). "Discovery of isolated dodo bonesRaphus cucullatus (L.), Aves, Columbiformes from Mauritius cave shelters highlights human predation, with a comment on the status of the family Raphidae Wetmore, 1930".Annales de Paléontologie.91 (2): 167.Bibcode:2005AnPal..91..167J.doi:10.1016/j.annpal.2004.12.002.
^Olson, Storrs L. (1985). "The fossil record of birds". In Farmer, Donald S.; King, James R.; Parkes, Kenneth C. (eds.).Avian Biology, Vol. VIII. Academic Press. pp. 79–238.hdl:10088/6553.ISBN978-0-12-249408-6.The earliest dove yet known, from the early Miocene (Aquitanian) of France, was a small species named Columba calcaria by Milne-Edwards (1867–1871) from a single humerus, for which Lambrecht (1933) later created the genus Gerandia
^abWorthy, Trevor H.; Hand, Suzanne J.; Worthy, Jennifer P.; Tennyson, Alan J. D.; Scofield, R. Paul (2009)."A large fruit pigeon (Columbidae) from the Early Miocene of New Zealand".The Auk.126 (3):649–656.doi:10.1525/auk.2009.08244.S2CID86799657.Because Columba calcaria Milne-Edwards, 1867–1871, from the Lower Miocene at Saint-Gérand-le-Puy in France, is now also considered a sandgrouse, as Gerandia calcaria (Mlíkovský 2002), there is no pre-Pliocene columbid record in Europe.
^Baptista, Luis F.; Trail, Pepper W.; Horblit, H.M.; Kirwan, Guy M.; Bonan, Arnau (2020). Del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi; Christie, David; De Juana, Eduardo (eds.)."Western Crowned-Pigeon Goura cristata".birdsoftheworld.org. Cornell Lab of Ornithology.doi:10.2173/bow.wecpig1.01. Retrieved16 February 2025.
^Baptista, Luis F.; Trail, Pepper W.; Horblit, H.M.; Kirwan, Guy M.; Garcia, Ernest (2020). Del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi; Christie, David; De Juana, Eduardo (eds.)."Scheepmaker's Crowned-Pigeon Goura scheepmakeri".birdsoftheworld.org. Cornell Lab of Ornithology.doi:10.2173/bow.soucrp2.01. Retrieved16 February 2025.
^Thorsen, M., Blanvillain, C., & Sulpice, R. (2002). Reasons for decline, conservation needs, and a translocation of the critically endangered upe (Marquesas imperial pigeon, Ducula galeata), French Polynesia. Department of Conservation.
^Baptista, Luis F.; Trail, Pepper W.; Horblit, H. M.; Sharpe, Christopher J.; Boesman, Peter F. D.; Garcia, Ernest (4 March 2020). Del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi; Christie, David; De Juana, Eduardo (eds.)."Somali Pigeon (Columba oliviae)".Birds of the World.doi:10.2173/bow.sompig1.01.S2CID240954419.