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Columbidae

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Columbidae is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with small heads, relatively short necks and slender bills that in some species feature fleshy ceres. They feed largely on plant matter, feeding on seeds (granivory), fruit (frugivory), and foliage (folivory).

In colloquial English, the smaller species tend to be called "doves", and the larger ones "pigeons",<ref name=floss>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> although the distinction is not consistent,<ref name=floss /> and there is no scientific separation between them.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Historically, the common names for these birds involve a great deal of variation. The bird most commonly referred to as "pigeon" is the domestic pigeon, descendant of the wild rock dove, which is a common inhabitant of cities as the feral pigeon.

Columbidae contains 51 genera divided into 353 species.<ref name="CornellLab">Template:Cite journal</ref> The family occurs worldwide, often in close proximity to humans, but the greatest diversity is in the Indomalayan and Australasian realms. 118 species (34%) are at risk,<ref name="CornellLab" /> and 13 are extinct,<ref name="ioc">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> with the most famous examples being the dodo, a large, flightless, island bird, and the passenger pigeon, that once flocked in the billions.

EtymologyEdit

{{#invoke:Lang|lang}} is a French word that derives from the Latin {{#invoke:Lang|lang}}, for a Template:Gloss chick,<ref name="oetymd-pigeon">Template:OEtymD</ref> while dove is an ultimately Germanic word, possibly referring to the bird's diving flight.<ref>Template:OEtymD</ref> The English dialectal word Template:Wikt-lang appears to derive from Latin Template:Wikt-lang.<ref name="oetymd-pigeon" /> A group of doves has sometimes been called a "dule", taken from the French word {{#invoke:Lang|lang}} (Template:Gloss).<ref>Template:Cite book</ref>

Origin and evolutionEdit

Columbiformes is one of the most diverse non-passerine clades of neoavians, and its origins are in the Cretaceous<ref>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</ref> and the result of a rapid diversification at the end of the K-Pg boundary.<ref>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).</ref> Whole genome analyses have found Columbiformes is the sister clade to the clade Pteroclimesites a clade consisting the orders Pterocliformes (sandgrouses) and Mesitornithiformes (mesites).<ref name=Jarvis2014>Template:Cite journal</ref><ref name=Prum2015>Template:Cite journal</ref><ref>Template:Cite journal</ref> The columbiform-pteroclimesitean clade, or Columbimorphae, monophyly has been supported from several studies.<ref name=Jarvis2014/><ref name=Prum2015/><ref name="ReddyKimball2017">Template:Cite journal</ref><ref name="BraunCracraft2019">Template:Cite book</ref><ref name="Suh2016">Template:Cite journal</ref><ref name="HoudeBraun2019">Template:Cite journal</ref><ref name="Wuetal2024">Template:Cite journal</ref><ref name="Stilleretal2024">Template:Cite journal</ref>

Taxonomy and systematicsEdit

Template:See also

The name 'Columbidae' for the family was first used by the English zoologist William Elford Leach in a guide to the contents of the British Museum published in 1819.<ref>Template:Cite book Although the name of the author is not specified in the document, Leach was the Keeper of Zoology at the time.</ref><ref>Template:Cite book</ref> However, Illiger in 1811 established an older name for the family group ("Columbini") and would actually be the proper authority for Columbidae.<ref name="Raphina2024"/>

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.<ref name="Allen2009">Template:Cite book</ref><ref name=Shapiro>Template:Cite journal</ref><ref name="Chekeetal2010">Template:Cite book</ref> 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).<ref name="Raphina2024">Template:Cite journal</ref> A 2025 paper on the molecular phylogenetic placement of the Cuban endemic blue-headed quail-dove from Oswald and colleagues found the species to be a sister 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.<ref name="Oswaldetal2025">Template:Cite journal</ref>

These taxonomic issues are exacerbated by columbids not being well represented in the fossil record,<ref>Template:Cite journal</ref> with no truly primitive forms having been found to date.Template:Citation needed The genus Gerandia has been described from Early Miocene deposits in France, but while it was long believed to be a pigeon,<ref>Template:Cite book</ref> it is now considered a sandgrouse.<ref name=auk>Template:Cite journal</ref> Fragmentary remains of a probably "ptilinopine" Early Miocene pigeon were found in the Bannockburn Formation of New Zealand and described as Rupephaps;<ref name=auk/> "Columbina" prattae from roughly contemporary deposits of Florida is nowadays tentatively separated in Arenicolumba, but its distinction from Columbina/Scardafella and related genera needs to be more firmly established (e.g. by cladistic analysis).<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Apart from that, all other fossils belong to extant genera.<ref>Template:Cite book</ref>

List of generaEdit

Fossil species of uncertain placement:

  • Genus †Arenicolumba Steadman, 2008
  • Genus †Rupephaps Worthy, Hand, Worthy, Tennyson, & Scofield, 2009 (St. Bathans pigeon, Miocene of New Zealand)

Subfamily Columbinae (typical pigeons and doves) Illiger, 1811Edit

Subfamily Starnoenadinae Bonaparte, 1855Edit

Subfamily Claravinae (American ground doves) Todd, 1913Edit

Subfamily Raphinae (Old World doves and pigeons) Oudemans, 1917 (1835)Edit

DescriptionEdit

Anatomy and physiologyEdit

File:Touchdown (pigeon).jpg
A landing collared dove (Streptopelia decaocto) displays the contour and flight feathers of its wings.

Overall, the anatomy of Columbidae is characterized by short legs, short bills with a fleshy cere, and small heads on large, compact bodies.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Like some other birds, the Columbidae have no gall bladders.<ref name="Hagey">Template:Cite journal</ref> Some medieval naturalists concluded they have no bile (gall), which in the medieval theory of the four humours explained the allegedly sweet disposition of doves.<ref name="Isidore">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In fact, however, they do have bile (as Aristotle had earlier realized), which is secreted directly into the gut.<ref name="Browne">Template:Cite book</ref>

The wings of most species are large, and have eleven primary feathers;<ref name=":1">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> pigeons have strong wing muscles (wing muscles comprise 31–44% of their body weight<ref>Template:Cite book</ref>) and are among the strongest fliers of all birds.<ref name=":1" />

In a series of experiments in 1975 by Dr.Template:NbspMark B. Friedman, using doves, their characteristic head bobbing was shown to be due to their natural desire to keep their vision constant.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> It was shown yet again in a 1978 experiment by Dr.Template:NbspBarrie J. Frost, in which pigeons were placed on treadmills; it was observed that they did not bob their heads, as their surroundings were constant.<ref>Template:Cite journal</ref>

FeathersEdit

File:Pigeon Feathers - A Dissection Sampling.pdf
Pigeon feather types, excluding down

Columbidae have unique body feathers, with the shaft being generally broad, strong, and flattened, tapering to a fine point, abruptly.<ref name=":1" /> In general, the aftershaft is absent; however, small ones on some tail and wing feathers may be present.<ref>Template:Cite book</ref> Body feathers have very dense, fluffy bases, are attached loosely into the skin, and drop out easily.<ref>Template:Cite journal</ref> Possibly serving as a predator avoidance mechanism,<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> large numbers of feathers fall out in the attacker's mouth if the bird is snatched, facilitating the bird's escape. The plumage of the family is variable.<ref>Template:Cite book</ref>

Granivorous species tend to have dull plumage, with a few exceptions, whereas the frugivorous species have brightly coloured plumage.<ref name="HBW2">Template:Cite book</ref> The genera Chalcophaps, Ptilinopus and Alectroenas include some of the most brightly coloured pigeons. Pigeons and doves may be sexually monochromatic or dichromatic.<ref>Template:Cite journal</ref> In addition to bright colours, some pigeon species may have crests or other ornamentation.<ref name=":122">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

File:Snow Pigeon Flock Flying Pangolakha WLS Sikkim India 11.02.2016.jpg
Snow pigeons flying at an altitude of Template:Convert above sea level

FlightEdit

Many Columbidae are excellent fliers due to the lift provided by their large wings, which results in low wing loading.<ref>Template:Cite book</ref> They are highly maneuverable in flight<ref>Template:Cite book</ref> and have a low aspect 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.<ref>Template:Cite journal</ref> A few species are long-distance migrants, with some populations of the European turtle dove migrating in excess of 5,000 km between northern Europe in summer and tropical Africa in winter, and the Oriental turtle dove nearly as far in eastern Asia between eastern Siberia and southern China.

SizeEdit

Pigeons and doves exhibit considerable variation in size, ranging in length from Template:Convert, and in weight from Template:Convert to above Template:Convert.<ref name=":02">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The largest extant species are the crowned pigeons of New Guinea,<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> which are nearly turkey-sized, with lengths of Template:Convert and weights ranging Template:Convert.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> One of the largest arboreal species, the Marquesan imperial pigeon with a length of Template:Convert, currently battles extinction.<ref>Template:Cite book</ref><ref>Template:Cite journal</ref> The extinct, flightless dodo is the largest columbid to have ever existed, with a height of about Template:Cvt, and a range of suggested weights from Template:Convert, although the higher estimates are thought to be based on overweight birds.<ref name=Hume2017>Template:Cite book</ref><ref>Template:Cite book</ref><ref name=Angst2011>Template:Cite journal</ref><ref name=KitchenerAugust1993>Template:Cite magazineTemplate:Subscription required</ref>

The least massive columbids belong to species in the genus Columbina; the common ground dove (Columbina passerina) and the plain-breasted ground dove (Columbina minuta) which are about the same size as a house sparrow, weighing a little above Template:Convert.<ref name="HBW2"/><ref name="Uni StA pdf">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The dwarf fruit dove, which may measure as little as Template:Convert long, has a marginally smaller total length than any other species from this family.<ref name="HBW2"/>

  • Diversity of Pigeons and Doves
  • Nicobar Pigeon on the bar.jpg

    The Nicobar pigeon (Caloenas nicobarica) is often stated to be the dodo's closest living relative.

  • SNOW-PIGEON-SELA.jpg

    Snow pigeon (Columba leuconota) in Sela, Arunachal Pradesh

  • 2019-03-17 Columba oenas, Jesmond Dene 4.jpg

    The stock dove (Columba oenas) of Europe is a typical member of the Columbinae.

  • 2018-03-14 Columba palumbus eating Cotoneaster frigidus berries.jpg

    The common wood pigeon (Columba palumbus) is common throughout Europe. This one is eating Cotoneaster frigidus berries.

  • Columbina passerina.jpg

    The common ground dove (Columbina passerina) is one of the smallest species in the family.

  • Ducula galeata Nuku Hiva.jpg

    Nuku Hiva/Marquesan imperial pigeon (Ducula galeata)

  • Goura victoria LC0384.jpg

    The Victoria crowned pigeon (Goura victoria) is one of the largest extant pigeons.

  • Blue-headed quail dove (Starnoenas cyanocephala).JPG

    The blue-headed quail-dove (Starnoenas cyanocephala) of Cuba is a relictual species with no close relatives.

  • Red-eyed dove (Streptopelia semitorquata).jpg

    A red-eyed dove (Streptopelia semitorquata) on the Zambezi in Zimbabwe

Distribution and habitatEdit

Pigeons and doves are distributed everywhere on Earth, having adapted to most terrestrial habitats available on the planet, except for the driest areas of the Sahara Desert, Antarctica and its surrounding islands, and the high Arctic.<ref name=":02"/> They have colonised most of the world's oceanic islands, reaching eastern Polynesia and the Chatham Islands in the Pacific, Mauritius, the Seychelles and Réunion in the Indian Ocean, and the Azores in the Atlantic Ocean.

Columbid species may be arboreal, terrestrial, or semi-terrestrial. They inhabit savanna, grassland, shrubland, desert, temperate woodland and forest, tropical rainforests, mangrove forest, and even the barren sands and gravels of atolls.<ref name=":16">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Some species have large natural ranges. The eared dove ranges across the entirety of South America from Colombia to Tierra del Fuego,<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> the Eurasian collared dove has a massive (if discontinuous) distribution from Britain across Europe, the Middle East, India, Pakistan and China,<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> and the laughing dove across most of sub-Saharan Africa, as well as India, Pakistan, and the Middle East.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

When including human-mediated introductions, the largest range of any species is that of the rock dove, also known as the common pigeon.<ref name=":4">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> This species had a large natural distribution from Britain and Ireland to northern Africa, across Europe, Arabia, Central Asia, India, the Himalayas and up into China and Mongolia.<ref name=":4" /> The range of the species increased dramatically upon domestication, as the species went feral in cities around the world.<ref name=":4" /> 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.<ref name=":4" /> 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.<ref>Template:Cite news</ref>

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.<ref name="HBW2"/>

Other species of Columbidae have tiny, restricted distributions, usually seen on small islands, such as the whistling dove, which is endemic to the tiny Kadavu Island in Fiji,<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> the Caroline ground dove, restricted to two islands, Truk and Pohnpei in the Caroline Islands,<ref>Template:Cite book</ref> and the Grenada dove, which is only found on the island of Grenada in the Caribbean.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Some continental species also have tiny distributions, such as the black-banded fruit dove, which is restricted to a small area of the Arnhem Land of Australia,<ref>Template:Cite book</ref> the Somali pigeon, found only in a tiny area of northern Somalia,<ref>Template:Cite journal</ref> and Moreno's ground dove, endemic to the area around Salta and Tucuman in northern Argentina.<ref name="HBW2"/>

BehaviourEdit

FeedingEdit

File:Treron Sieboldii.jpg
White-bellied green pigeon (Treron sieboldii) feeding on fruit

Seeds and fruit form the major component of the diets of pigeons and doves,<ref name=":02"/><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> and the family can be loosely divided between seed-eating (granivorous) species, and fruit-and-mast-eating (frugivorous) species, though many species consume both.<ref name=":2">Template:Cite news</ref>

The granivorous species typically feed on seed found on the ground, whereas the frugivorous species are more arboreal, tending to feed in trees.<ref name=":2" /> The morphological adaptations used to distinguish between the two groups include granivores tending to having thick walls in their gizzards, intestines, and esophagi, with the frugivores evolved with thin walls,<ref name=":02"/> and the fruit-eating species have short intestines, as opposed to the seed eaters having longer intestines.<ref>Template:Cite book</ref> Frugivores are capable of clinging to branches and even hang upside down to reach fruit.<ref name="HBW2"/><ref name=":2" />

In addition to fruit and seeds, a number of other food items are taken by many species. Some, particularly the ground doves and quail-doves, eat a large number of prey items such as insects and worms.<ref name=":2" /> One species, the atoll fruit dove, is specialised in taking insect and reptile prey.<ref name=":2" /> Snails, moths, and other insects are taken by white-crowned pigeons, orange fruit doves, and ruddy ground doves.<ref name="HBW2"/> Flowers are also taken by some species.<ref name="CornellLab" />

Urban feral pigeons, descendants of domestic rock 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 for spilled food or food provided by humans.<ref>Template:Cite journal</ref>

File:Spotted Dove and its egg.jpg
Spotted dove (Spilopelia chinensis) on its nest, with one unhatched egg and one hatchling

ReproductionEdit

Doves and pigeons build relatively flimsy nests, often using sticks, other vegetable matter, and other debris, which may be placed on trees, on rocky 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 species nest colonially, others nest in aggregation.<ref name="CornellLab" />

Most lay a clutch of one or (usually) two white eggs at a time which take 11-30 days to hatch (larger species have longer incubation times). Both parents 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 of epithelial cells from the lining of the crop.<ref name="CornellLab" />

Unfledged baby doves and pigeons are called squabs and are generally able to fly by five weeks old. These fledglings, with their immature squeaking voices, are called squeakers once they are weaned,<ref name="EoB">Template:Cite book</ref> and leave the nest after 25–32 days.

Status and conservationEdit

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 become threatened or even succumbed to extinction.<ref name=":5" /> Among the ten species to have become extinct since 1600 (the conventional date for estimating modern extinctions) are two of the most famous extinct species, the dodo and the passenger pigeon.<ref name=":5">Template:Cite news</ref><ref name="CornellLab" />

The passenger 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<ref name=":5" /> even though it was once the most numerous species of bird on Earth.Template:Citation needed Its former numbers are difficult to estimate, but one ornithologist, Alexander Wilson, estimated one flock he observed contained over two billion birds.<ref>Template:Cite magazine</ref> 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.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Although habitat loss was a contributing factor, the species is thought to have been massively over-hunted, being used as food for slaves and, later, the poor, in the United States throughout the 19thTemplate:Nbspcentury.Template:Citation needed

File:Zenaida graysoni 1.jpg
The Socorro dove (Zenaida graysoni) is extinct in the wild.

The dodo, and its extinction, was more typical of the extinctions of pigeons in general. Like many species that colonise remote islands with few predators, it lost much of its predator avoidance behaviour, along with its ability to fly.<ref name=":10">Template:Cite book</ref> The arrival of people, along with a suite of other introduced species such as rats, pigs, and cats, quickly spelled the end for this species and many other island species that have become extinct.<ref name=":10" />

118 columbid species are at risk (34% of the total), with 48 species NT, 40 VU, 18 EN, 11 CR, and 1 EW.<ref name="CornellLab" /> Most of these are tropical and live on islands. All of the species are threatened by introduced predators, habitat loss, hunting, or a combination of these factors.<ref name=":10" /> In some cases, they may be extinct in the wild, as is the Socorro dove of Socorro Island, Mexico, last seen in the wild in 1972, driven to extinction by habitat loss and introduced feral cats.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In some areas, a lack of knowledge means the true status of a species is unknown (DD); the Negros fruit dove has not been seen since 1953,<ref>Template:Cite journal</ref> and may or may not be extinct, and the Polynesian ground dove is classified as critically endangered, as whether it survives or not on remote islands in the far west of the Pacific Ocean is unknown.<ref>Template:Cite journal</ref>

Various conservation techniques are employed to prevent these extinctions, including laws and regulations to control hunting pressure, the establishment of protected areas to prevent further habitat loss, the establishment of captive populations for reintroduction back into the wild (ex situ conservation), and the translocation of individuals to suitable habitats to create additional populations.<ref name=":10" /><ref>Template:Cite book</ref>

DomesticationEdit

{{#invoke:Labelled list hatnote|labelledList|Main article|Main articles|Main page|Main pages}} The domestic pigeon (Columba livia domestica) is a descendant of the rock dove (Columba livia) that underwent domestication, with studies suggesting domestication as early as 10 thousand years ago. Domestic pigeons have long been a part of human culture; doves were important symbols of the goddesses Innana, Asherah, and Aphrodite, and revered by the early Christian, Islamic and Jewish religions. Domestication of pigeons led to significant use of homing pigeons for communication, including war pigeons, such as the 32 pigeons who were awarded the Dickin Medal for "brave service" to their country, in World War II.

The ringneck 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.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

See alsoEdit

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NotesEdit

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ReferencesEdit

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Further readingEdit

  • Blechman, Andrew, Pigeons: The Fascinating Saga of the World's Most Revered and Reviled Bird (Grove Press 2007) Template:ISBN
  • Gibbs, Barnes and Cox, Pigeons and Doves (Pica Press 2001) Template:ISBN

External linksEdit

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