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Columbidae
Temporal range: Early Miocene – recent[1]
The pink-necked green pigeon, a frugivorous species
The speckled pigeon, a granivorous species
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Clade: Columbimorphae
Order: Columbiformes
Latham, 1790
Family: Columbidae
Illiger, 1811
Type genus
Columba
Linnaeus, 1758
Subfamilies

See text

      Geographic range of the family

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",[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 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.[4] 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,[4] and 13 are extinct,[5] with the most famous examples being the dodo, a large, flightless, island bird, and the passenger pigeon, that once flocked in the billions.

Etymology

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Pigeon is a French word that derives from the Latin pīpiō, for a 'peeping' chick,[6] while dove is an ultimately Germanic word, possibly referring to the bird's diving flight.[7] The English dialectal word culver appears to derive from Latin columba.[6] A group of doves has sometimes been called a "dule", taken from the French word deuil ('mourning').[8][page needed]

Origin and evolution

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Columbiformes is one of the most diverse non-passerine clades of neoavians, and its origins are in the Cretaceous[9] and the result of a rapid diversification at the end of the K-Pg boundary.[10] Whole genome analyses have found Columbiformes is the sister clade to the clade Pteroclimesites a clade consisting the orders Pterocliformes (sandgrouses) and Mesitornithiformes (mesites).[11][12][13] The columbiform-pteroclimesitean clade, or Columbimorphae, monophyly has been supported from several studies.[11][12][14][15][16][17][18][19]

Taxonomy and systematics

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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.[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 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.[26]

These taxonomic issues are exacerbated by columbids not being well represented in the fossil record,[27] with no truly primitive forms having been found to date.[citation needed] The genus Gerandia has been described from Early Miocene deposits in France, but while it was long believed to be a pigeon,[28] it is now considered a sandgrouse.[29] Fragmentary remains of a probably "ptilinopine" Early Miocene pigeon were found in the Bannockburn Formation of New Zealand and described as Rupephaps;[29] "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).[30] Apart from that, all other fossils belong to extant genera.[31]

List of genera

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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, 1811

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Subfamily Starnoenadinae Bonaparte, 1855

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Subfamily Claravinae (American ground doves) Todd, 1913

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

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Description

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Anatomy and physiology

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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.[32] Like some other birds, the Columbidae have no gall bladders.[33] 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.[34] In fact, however, they do have bile (as Aristotle had earlier realized), which is secreted directly into the gut.[35][page needed]

The wings of most species are large, and have eleven primary 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 on treadmills; it was observed that they did not bob their heads, as their surroundings were constant.[39]

Feathers

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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.[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 a predator avoidance mechanism,[42] 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.[43]

Granivorous species tend to have dull plumage, with a few exceptions, whereas the frugivorous species have brightly coloured plumage.[44][page needed] The genera Chalcophaps, Ptilinopus and Alectroenas include some of the most brightly coloured pigeons. Pigeons and doves may be sexually monochromatic or dichromatic.[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

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Many Columbidae are excellent fliers due to the lift provided by their large wings, which results in low wing loading.[47] They are highly maneuverable in flight[48] 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.[49] 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.

Size

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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 the crowned pigeons of New Guinea,[51] which are nearly turkey-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 largest arboreal species, the Marquesan imperial pigeon with a length of 55 cm (22 in), currently battles extinction.[55][page needed][56] The extinct, flightless dodo 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 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 22 g (0.78 oz).[44][page needed][61][62] The dwarf 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][page needed]

Distribution and habitat

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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.[50] 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.[63]

Some species have large natural ranges. The eared dove ranges across the entirety of South America from Colombia to Tierra del Fuego,[64] the Eurasian collared dove has a massive (if discontinuous) distribution from Britain across Europe, the Middle East, India, Pakistan and China,[65] and the laughing 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 the rock 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, the Himalayas and up into China and Mongolia.[67] The range of the species increased dramatically upon domestication, as the species went feral 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][page needed]

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,[69] the Caroline ground dove, restricted to two islands, Truk and Pohnpei in the Caroline Islands,[70] and the Grenada dove, which is only found on the island of Grenada in the Caribbean.[71]

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,[72] the Somali pigeon, found only in a tiny area of northern Somalia,[73] and Moreno's ground dove, endemic to the area around Salta and Tucuman in northern Argentina.[44][page needed]

Behaviour

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Feeding

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White-bellied green pigeon (Treron sieboldii) feeding on fruit

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 more arboreal, 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, and esophagi, 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][page needed][75]

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.[75] One species, the atoll fruit dove, is specialised in taking insect and reptile prey.[75] Snails, moths, and other insects are taken by white-crowned pigeons, orange fruit doves, and ruddy ground doves.[44][page needed] Flowers are also taken by some species.[4]

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.[77]

Spotted dove (Spilopelia chinensis) on its nest, with one unhatched egg and one hatchling

Reproduction

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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.[4]

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.[4]

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,[78] and leave the nest after 25–32 days.

Status and conservation

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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.[79] 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.[79][4]

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[79] even though it was once the most numerous species of bird on Earth.[citation needed] Its former numbers are difficult to estimate, but one ornithologist, 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] 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 19th century.[citation needed]

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.[82] 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.[82]

118 columbid species are at risk (34% of the total), with 48 species NT, 40 VU, 18 EN, 11 CR, and 1 EW.[4] 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.[82] 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.[83] 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,[84] 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.[85]

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.[82][86]

Domestication

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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.[87]

See also

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References

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Further reading

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Columbidae is a family of birds in the order Columbiformes, consisting of pigeons and doves, which are characterized by their stout bodies, small heads, short legs, and slender bills. This family is the sole within its order and includes approximately 350 extant distributed across all continents except the high and . These birds exhibit a wide range of sizes, from 15 to 75 cm in length and weights from 30 g to over 2 kg, with many species displaying iridescent neck feathers and producing a unique nutrient-rich "" for feeding their young. Columbidae encompasses five subfamilies, around 50 genera, and shows the greatest species diversity in the Indomalayan and Australasian realms, though many species thrive in close association with humans in urban and agricultural environments worldwide. Habitats vary from deserts and forests to high-altitude regions up to 5,000 m and oceanic islands, with some species being highly nomadic or migratory. Primarily granivorous and frugivorous, columbids consume seeds, fruits, and occasionally insects or snails, often foraging on the ground in flocks that can number from solitary pairs to massive congregations, as seen historically with the extinct passenger pigeon (Ectopistes migratorius). Reproductively, most species are monogamous, laying one or two eggs that are incubated for 11 to 30 days by both parents, with chicks fed —a protein- and fat-rich secretion from the parents' —enabling rapid growth. Behaviorally, columbids are known for strong pair bonds, elaborate displays involving cooing and bowing, and exceptional abilities, which have led to their use in homing and . Conservation challenges are significant, with approximately 70 species classified as threatened (Critically Endangered, Endangered, or Vulnerable) by the IUCN as of 2023 and 26 listed under , primarily due to habitat loss, hunting, and , underscoring the need for targeted research and protection efforts.

Etymology and Classification

Etymology

The family name Columbidae derives from the Latin columba, meaning "dove," which itself stems from the earlier form columbus denoting a male dove, and traces back to the kolumbos, referring to a "diver" in to the bird's bobbing head movements while drinking. The suffix -idae is a standard New Latin ending for taxonomic families, first applied to this group by English zoologist in 1820 to encompass pigeons and doves as a unified biological family. In ancient Roman texts, columba was commonly used to describe these birds, as seen in Pliny the Elder's (circa 77 CE), where he details the habits, longevity, and nutritional properties of pigeons (columbae) and turtle-doves, noting their eight-year lifespan and the "milk" they produce for feeding young. This term reflected early observations of their gentle demeanor and domestic utility, influencing later European . The Greek peristera, of uncertain origin but denoting both pigeons and doves, appears in classical literature and biblical translations, paralleling columba and emphasizing shared traits like cooing and nesting behaviors across . Such naming often evoked cultural symbols of peace—stemming from the dove's role in the narrative—and fertility, due to prolific breeding, as captured in ancient Mediterranean and Near Eastern traditions. Scientific , formalized by in (1758), standardized as the for many , distinguishing it from vernacular names like "pigeon" and "dove," which lack taxonomic precision and vary regionally. In English, "pigeon" originates from pijon (a young dove or peeping chick), typically applied to larger, often urban species like the rock pigeon (Columba livia), while "dove" derives from dūfe (of Germanic roots), reserved for smaller, wilder forms such as the (Zenaida macroura). Post-Linnaean refined these common names to align with emerging classifications, shifting from broad folk terms to more descriptive ones that highlight size, habitat, or behavior, though the pigeon-dove dichotomy persists as a cultural rather than biological divide in English-speaking regions.

Taxonomy and Systematics

The family Columbidae belongs to the order Columbiformes and encompasses approximately 350 across 51 genera, representing a diverse assemblage of pigeons and doves distributed worldwide. This hierarchical structure reflects a long history of taxonomic refinement, with the family recognized as monophyletic based on shared morphological and genetic traits, such as a unique crop-milk production system and robust body form. Major subfamilies within Columbidae include Columbinae, which contains typical pigeons and doves adapted to a wide range of habitats; Claravinae, comprising small ground-dwelling doves often found in open grasslands; and Raphinae, featuring fruit doves and larger imperial pigeons specialized for arboreal and island ecosystems. Minor subfamilies, such as Gourinae (crowned pigeons of ), highlight further diversification into niche adaptations, with the (Otidiphaps nobilis) now placed within Raphinae. These groupings are supported by molecular phylogenies that delineate deep evolutionary divergences. Prominent genera illustrate diagnostic traits integral to classification. The genus Columba, including the rock pigeon (C. livia), is defined by medium-to-large size (typically 30–40 cm), a strong bill with a cere, and metallic iridescent feathers on the neck, often used in mate displays. In contrast, Streptopelia, encompassing turtle doves like the Eurasian collared-dove (S. decaocto), features a slimmer build, graduated tail with white tips, and a soft, purring vocalization, distinguishing it from more robust congeners. These morphological markers, combined with genetic profiles, facilitate species delineation within subfamilies. Recent taxonomic revisions, driven by 2020s molecular studies incorporating (mtDNA) and nuclear genes, have affirmed the of core subfamilies like Columbinae and Raphinae while prompting adjustments elsewhere. For instance, analyses of multi-locus datasets have confirmed the phylogenetic integrity of Claravinae and led to the elevation of taxa like Oxypelia from within Columbina to full genus status as of 2025, reflecting finer resolution of versus divergences. These updates underscore the dynamic nature of columbid , prioritizing phylogenetic congruence over traditional morphology alone.

Evolutionary History

Origin and Evolution

The family Columbidae traces its origins to the period, with molecular phylogenetic analyses indicating that the stem lineage diverged from close relatives such as the (Pteroclidae) around 55 million years ago in the Eocene. The crown group radiation, marking the diversification of extant lineages, is estimated to have begun around 25 million years ago (estimates vary from approximately 25 to 42 million years ago across studies), rather than in the earlier or Eocene as previously suggested by some mitochondrial studies. These estimates derive from Bayesian methods calibrated with whole-mitogenome data and fossil constraints, such as the divergence between Holarctic (including ) and Indo-Pacific clades around 24.7 million years ago. Ancestral forms likely resembled basal neoavian birds with adaptations suited to forested or woodland environments, evolving from lineages within the broader . A major evolutionary event was the radiation, driven by climatic cooling, tectonic shifts, and , which facilitated the split into distinct biogeographic clades: the predominantly group and the Holarctic assemblage encompassing Eurasian and taxa. This diversification reflects a Laurasian-Gondwanan partitioning influenced by , with the clade showing strong ties to Australasian landmasses. Genomic studies from the , incorporating next-generation sequencing of museum specimens, support these timelines, revealing accelerated rates in island systems like and during the mid-. Among key adaptive innovations, the evolution of production stands out as a defining trait of Columbidae (convergently evolved in and penguins), enabling both parents to secrete a protein- and lipid-rich nutritive fluid from the esophageal crop lining to feed altricial nestlings. This secretion, regulated by , likely arose in the early to support rapid chick growth on a - and fruit-based diet, distinguishing columbids from other avian families reliant on regurgitation or predation. Biogeographically, initial diversification centered in , particularly as a hotspot with high , followed by dispersals to via the islands, through continental bridges, and the via trans-Beringian routes or overwater crossings. These patterns, corroborated by 2020 phylogenetic reconstructions using multi-locus nuclear and mitochondrial data, underscore a history of vicariance and long-distance dispersal shaping the family's .

Fossil Record and Phylogeny

The fossil record of Columbidae begins in the Late Oligocene, with the earliest known specimens consisting of a bone attributed to Primophaps schoddei, a phabine pigeon from the Riversleigh World Heritage site in northwestern , . This discovery represents the oldest definitive columbid globally, dating to approximately 25 million years ago (mya) and indicating that pigeons had already diversified into specialized fruit-eating lineages by this time. Subsequent Early deposits yield more substantial evidence of columbid diversity, such as the small-bodied Arenicolumba prattae from the Thomas Farm locality in , , known from hundreds of postcranial elements including , humeri, and sterna that reveal adaptations similar to modern ground doves. Another Early taxon, the large fruit pigeon Rupephaps taketake from New Zealand's St Bathans Fauna, further highlights early Gondwanan radiation, with its robust suggesting a diet of larger fruits akin to extant imperial pigeons. Notable extinct genera span multiple continents and epochs, providing snapshots of lost diversity. In , Miocene assemblages include fragmentary remains tentatively assigned to early Zenaida-like doves, while European Pliocene sites such as those in preserve bones of Columba species closely resembling modern rock pigeons. The Raphinae subfamily exemplifies dramatic insular and , with genera like Raphus (the dodo, extinct circa 1662 CE) and Pezophaps (, extinct circa 1761 CE) from the evolving flightlessness and large size before -induced loss through hunting and habitat alteration. Overall, Pleistocene megafaunal die-offs and subsequent of islands contributed to the of numerous columbid taxa, particularly flightless or endemic forms, with at least 13 species lost since the , representing about 4% of current recognized diversity but a higher proportion when accounting for undescribed prehistoric losses. Phylogenetic reconstructions of Columbidae rely on molecular data integrated with sparse fossil evidence, employing on mitochondrial and nuclear sequences to generate time-calibrated trees. A 2016 study using complete mitochondrial genomes from 24 species, including extinct taxa like the dodo and , estimated the crown-group radiation at approximately 24.7 mya (95% CI: 18.9–31.3 mya) during the Late , with major Indo-Pacific and Holarctic/New World clades diverging shortly thereafter. More recent analyses, such as a 2023 supermatrix approach incorporating 247 columbid species, confirm this timeline and resolve Raphinae as a derived clade sister to typical pigeons (Columbinae), while placing Columbiformes within the larger assemblage, with (Pteroclidae) as the closest based on shared morphological and genomic traits like precise bill control. These -calibrated phylogenies underscore an Australasian origin, with subsequent dispersals to other continents, though the lack of pre- columbid fossils limits precision on deeper divergences. The fossil record of Columbidae exhibits significant gaps, particularly in tropical habitats where over 80% of extant evolved and persist; acidic soils and high in rainforests hinder bone preservation, resulting in a bias toward lacustrine or temperate deposits like those at Riversleigh or Thomas Farm. This incompleteness challenges reconstructions of early diversification, as no columbid s predate the , despite molecular clocks suggesting a origin for the order.

Physical Characteristics

Morphology and Size

Columbidae, the family encompassing pigeons and doves, are characterized by a compact and rounded , featuring a small head relative to the torso, a short and stout bill, and relatively short legs adapted for perching or brief ground movements. This morphology contributes to their streamlined form, facilitating efficient flight despite their stocky build. is generally minimal within the , with differences primarily limited to slight variations in size, where males tend to be marginally larger than females in certain species. Species within Columbidae exhibit substantial variation in overall size, spanning lengths from 13 cm to 80 cm and weights from 22 g to over 4 kg. For instance, the dwarf fruit dove (Ptilinopus nainus), a representative of the smaller end of the spectrum, measures 13–15 cm in length and weighs around 49 g, though some ground doves like the plain-breasted ground dove (Columbina minuta) are lighter at 24–26 g, making it one of the lightest members of the family. At the opposite extreme, the Victoria crowned-pigeon (Goura victoria) stands as one of the largest living species, attaining lengths up to 80 cm and weights up to 3.5 kg, which underscores the family's diverse evolutionary adaptations. These size disparities reflect ecological niches, with smaller forms often suited to dense vegetation and larger ones to more exposed environments. Structural variations among Columbidae are closely tied to and , particularly in limb and morphology. Ground-dwelling , such as those in the Claravinae (American ground doves), typically possess stronger and more robust legs for enhanced terrestrial mobility and foraging on the forest floor or open ground. In contrast, arboreal like fruit doves (Ptilinopus spp.) often feature relatively longer , which provide balance and maneuverability while navigating tree canopies. Body size shows allometric scaling with preferences, where larger individuals predominate in open areas, potentially aiding predator avoidance through greater visibility and stride length during escape. Comparative morphology in Columbidae relies on standardized measurements to quantify these variations, including total body length, wing chord (a proxy for ), culmen (bill) length, and tarsus length, which collectively inform assessments of size, proportions, and adaptations across . These metrics highlight how size influences functional traits, such as flight efficiency, with larger exhibiting proportionally broader wings to sustain powered flight.

Anatomy and Physiology

Columbidae possess a specialized digestive system adapted for their granivorous diet and unique reproductive needs. The serves as a for ingested food, allowing temporary holding before further processing in the proventriculus and . This structure enables efficient bolus formation and supports rapid feeding during foraging bouts. A distinctive feature is the production of , a nutrient-rich regurgitated to feed nestlings, formed through sloughing of hyperplastic epithelial cells in the crop lining. This process is hormonally regulated, primarily by elevated levels from the , which stimulate cellular proliferation and lipid accumulation in the crop epithelium during the breeding season. The respiratory system in Columbidae is highly efficient, supporting prolonged flight through a unidirectional airflow mechanism distinct from the tidal breathing in mammals. Air sacs facilitate this by acting as bellows, with fresh air passing through the lungs in a continuous stream during both inhalation and exhalation. Pigeons and doves typically have nine air sacs—two cervical, two clavicular (fused into one interclavicular), two anterior thoracic, two posterior thoracic, and two abdominal—extending into pneumatic bones to reduce weight while enhancing oxygen delivery. This configuration, more extensive than the seven or eight in many other avian orders, contributes to sustained aerobic performance during migration and evasion. Sensory anatomy in Columbidae emphasizes visual reliance over olfaction, reflecting their diurnal and open-habitat lifestyles. Vision is predominantly , with eyes positioned laterally to provide a wide field of view exceeding 300 degrees, but the binocular overlap is limited to approximately 24-30 degrees in the frontal plane, sufficient for precise pecking during feeding. Despite this narrow stereoscopic field, they exhibit acute , tetrachromatic with sensitivity to ultraviolet light via four cone types, aiding in mate selection and detection. Olfactory capabilities are minimal, with a small relative to brain size and few functional glomeruli, rendering smell secondary to visual and tactile cues in and foraging. Skeletal features in Columbidae support agile flight and maneuverability, with adaptations varying across the family. The neck is notably flexible, comprising 12 cervical vertebrae that allow extensive head rotation and positioning for scanning the environment. The sternum features a prominent keel for attachment of flight muscles in volant species, but in flightless forms such as the dodo (Raphus cucullatus), the keel is reduced and flattened, correlating with loss of aerial locomotion and reliance on terrestrial movement. These skeletal traits, combined with lightweight pneumatic bones, optimize the balance between structural integrity and reduced mass. Physiological tolerances in Columbidae include elevated metabolic demands during breeding to support . Energy expenditure increases significantly for synthesis and , with metabolic rates rising up to 1.5-2 times basal levels due to heightened and activity, necessitating higher caloric intake from and grains. Uniquely among birds, they employ an intraoral drinking mechanism, using peristaltic tongue movements to create and draw water directly into the without tilting the head, enabling faster hydration in arid environments. This , akin to mammalian , conserves time and reduces vulnerability to predators.

Plumage and Coloration

The plumage of Columbidae is characterized by soft, dense contour feathers that provide effective insulation against environmental extremes, with the shaft typically broad, strong, and flattened to enhance aerodynamic efficiency and coverage. These feathers include specialized iridescent neck hackles, particularly prominent in males of species like the rock pigeon (Columba livia), where arises from granules and air-filled barbules that produce coherent light interference, resulting in shimmering greens, purples, and blues visible under certain angles. Coloration patterns vary widely across the family, reflecting ecological adaptations; ground-foraging species such as the (Zenaida macroura) exhibit cryptic browns and grays for against and , while fruit-dove genera like Ptilinopus display vibrant metallic greens and oranges derived from pigment-based mechanisms in the feather barbs. In fruit pigeons, non-iridescent greens stem from concentric arrays of granules within rounded barb ridges, contributing to their conspicuous tropical appearances. Molting cycles in Columbidae are typically annual and occur post-breeding, involving a complete prebasic molt that replaces body feathers, primaries, and secondaries sequentially from proximal to distal; for instance, in the rock pigeon, this process spans several months and may be incomplete in some individuals, retaining up to four outer primaries. Juvenile plumage is distinct, with chicks hatching in a sparse downy coat for initial before developing contour feathers, often retaining buffy or scaled patterns that differ from adult hues until the first complete molt. Plumage serves multiple functional roles, including for predator avoidance in terrestrial species, sexual signaling through iridescent display s during courtship, and by trapping air for warmth or facilitating heat dissipation in hot climates via fluffing. Melanin-based colors, common in many columbids, enhance UV protection and aid in heat absorption, while lighter pigments in tropical forms may reduce solar heat load. Variations in plumage include rare albinism, characterized by complete lack of melanin leading to white feathers and pink eyes, though more commonly documented are leucistic forms with partial pigment loss producing white patches while retaining normal eye color, as observed in feral rock pigeons. Additionally, many species exhibit ultraviolet (UV) reflectance in plumage, invisible to humans but critical for mate choice; for example, the iridescent neck feathers of rock pigeons show multiple UV peaks that enhance visual signaling during displays.

Locomotion and Flight

Columbidae species typically employ a style characterized by rapid, continuous wingbeats alternated with brief phases, enabling efficient travel over varied distances. This flight pattern is evident in species like the rock pigeon (Columba livia), where wingbeats occur at frequencies of approximately 5.5 per second during sustained flight. Takeoff involves a powerful burst, with rock pigeons capable of accelerating to speeds up to 97 mph (156 km/h) in trained individuals, facilitated by strong leg thrusts that contribute up to 25% of the initial vertical force. Wing morphology in Columbidae supports high maneuverability, featuring broad, rounded wings with relatively low aspect ratios in many species, which allow for quick turns and agile navigation in cluttered environments. Forest-dwelling doves, such as the common ground dove (Columbina passerina), exhibit even lower aspect ratios (around 1.59), optimizing for explosive takeoffs from the ground. In contrast, open-country pigeons like the rock pigeon possess higher aspect ratios with more pointed wings, enhancing efficiency for longer, straighter flights. On the ground, Columbidae display a distinctive waddling , often accompanied by head bobbing to stabilize vision during forward movement, with short bursts of running when alarmed. This locomotion is supported by short legs and a compact body, limiting sustained terrestrial travel but allowing quick evasion. Perching is facilitated by reversible outer toes and strong flexor tendons, enabling secure grips on branches or wires for resting and observation. Habitat adaptations in Columbidae include variations in flight capability, with some island endemics showing reduced size and flight prowess due to insular environments with fewer predators and limited dispersal needs; for instance, in the (Caloenas nicobarica) lineage, extinct relatives like the dodo (Raphus cucullatus) evolved near-flightlessness. The energetics of flight in Columbidae benefit from low , which minimizes power requirements for sustained aerial activity and enables long-distance migrations. Species in the genus Streptopelia, such as the (S. turtur), can cover total migration distances exceeding 5,000 km between and sub-Saharan Africa, with individual non-stop legs up to 700 km, supported by efficient muscle power outputs of 60–150 W/kg during cruising.

Distribution and Habitat

Geographic Range

The family Columbidae, comprising pigeons and doves, exhibits a near-cosmopolitan distribution, with approximately 353 occurring across all continents except , as of 2024. Recent taxonomic revisions have increased the recognized number from around 344 to 353 species. These birds are absent primarily from polar regions, high elevations above 5,000 meters, the driest parts of the Desert, and remote oceanic islands lacking suitable colonization pathways. Centers of species diversity are concentrated in the Indomalayan and Australasian realms, where environmental conditions support the highest richness; for instance, alone hosts 93 , representing a significant portion of global columbid diversity. In contrast, temperate zones, such as parts of and , support far fewer , often limited to a handful of widespread or introduced forms. Dispersal patterns in Columbidae include both natural colonizations and human-mediated introductions. Naturally, strong flight capabilities have enabled species like fruit doves (Ptilinopus) to reach and diversify across Pacific islands, leading to multiple independent radiations in and . Human activities have further expanded ranges, notably through the global introduction of the rock pigeon (Columba livia) starting in the early 1600s, which has established feral populations on every inhabited continent. Endemism is prominent on isolated islands, with over 50 restricted to archipelagos such as those in the Pacific and Indian Oceans, where geographic isolation has driven ; examples include the imperial pigeons (Ducula) of and the . These island endemics often face heightened vulnerability due to limited dispersal options and small population sizes. Contemporary range dynamics show shifts influenced by , including poleward expansions in some species. The Eurasian collared-dove (Streptopelia decaocto), for example, has rapidly extended its range northward across since the 1980s, facilitated by warmer conditions that extend breeding seasons and enhance survival in new latitudes.

Habitat Preferences

Columbidae species exhibit a broad spectrum of habitat preferences, ranging from arid deserts to humid tropical rainforests, reflecting their adaptability across diverse environmental conditions. In arid regions, species such as the spinifex pigeon (Geophaps plumifera) favor semi-arid stony habitats, including rocky hills, gorges, dry creek beds, and spinifex-dominated grasslands in northern and , where tussock-forming grasses provide essential cover and foraging opportunities. Conversely, in tropical environments, the emerald dove (Chalcophaps indica) inhabits dense wet woodlands, rainforests, mangroves, and forest understories across and the , preferring shaded, moist areas with thick foliage for concealment. This habitat diversity underscores the family's ecological versatility, with species distributed from to elevations exceeding 5,000 meters. Microhabitat utilization within these broader environments varies significantly among Columbidae groups, influencing their niche occupancy. Ground-foraging species, such as many small doves in open grasslands and savannas, exploit low for collection, while canopy-dwellers in forested habitats navigate upper tree layers for resources. Nesting preferences further diversify microhabitat use; for instance, some select cliff ledges or rocky outcrops for security, akin to the ancestral , whereas others build in tree branches or shrubs, as seen in woodland pigeons. These choices enable coexistence through spatial partitioning, particularly in heterogeneous landscapes. Certain adaptations enhance tolerance to modified habitats, notably urban environments, where populations of the rock pigeon (Columba livia) thrive on building ledges, parks, and city streets worldwide, substituting natural cliffs with anthropogenic structures. In contrast, many tropical Columbidae species show sensitivity to habitat alterations like , which disrupts and canopy availability critical for their survival. Seasonal variations in habitat use are evident in montane forms, such as the (Patagioenas fasciata), which undertakes altitudinal migrations along the , shifting from higher elevations in breeding seasons to lower valleys during non-breeding periods to track food resources in coniferous and mixed forests. Biodiversity within Columbidae correlates strongly with humid tropical regions, where niche partitioning allows multiple to occupy layered strata, from forest floors to canopies, fostering higher compared to temperate or arid zones. This pattern is particularly pronounced in and Neotropical hotspots, where environmental complexity supports specialized microhabitats.

Behavior and Ecology

Foraging and Diet

Columbidae, the family encompassing pigeons and doves, are predominantly granivorous, with seeds and grains comprising up to 90% of the diet in many species, such as the rock pigeon (Columba livia), which primarily consumes cultivated grains like (Triticum), (Hordeum), and oats (), along with and weed seeds. This seed-based diet is supplemented by fruits in frugivorous members, such as the (Columba palumbus), which exploits a variety of tree fruits and seeds from 45 species, including exotic ornamentals like plane trees ( spp.), and occasionally by like or earthworms, particularly to meet the protein needs of juveniles. The family's dietary flexibility allows adaptation to diverse environments, from forests to urban areas, where species like the rock pigeon opportunistically scavenge human food waste, including bread and discarded grains. Foraging in Columbidae typically involves ground-based pecking, where birds walk bipedally and probe or leaf litter with rapid bill movements to extract seeds, as observed in the (Streptopelia turtur), an obligate granivore that targets cultivated cereals like (66.7% frequency of occurrence in diet). In urban settings, rock pigeons employ similar pecking while scavenging, often in flocks that enhance detection and reduce predation risk through collective vigilance. A key physiological adaptation is the , a storage organ that allows irregular feeding patterns by holding undigested food for later processing in the , enabling efficient energy management during daily bouts that span daylight hours. To aid digestion of hard seeds, individuals ingest grit—small stones or sand—picked up during ground , a behavior universal across the . Seasonal variations in diet reflect environmental availability, with granivorous species like the stock dove (Columba oenas) relying more on seeds from families such as Poaceae and Fabaceae during dry periods, while frugivores like imperial pigeons (Ducula spp.) shift to fruit-heavy intake in wet seasons for hydration from high-water-content items like berries and figs. In temperate regions, the common wood pigeon increases arboreal fruit consumption (e.g., from Ligustrum and Celtis) in autumn and winter, transitioning to ground seeds in summer, demonstrating the family's opportunistic foraging strategy. This nutritional physiology supports hydration and energy needs, particularly in arid habitats where water-rich fruits supplement seed intake.

Reproduction and Breeding

Columbidae species exhibit predominantly monogamous mating systems, where pairs typically form lasting bonds for a breeding season or longer, often reusing the same partner in subsequent years. Courtship behaviors are elaborate and species-specific, involving displays such as bowing, cooing vocalizations, tail spreading, and aerial chases to attract mates and establish pair bonds. In some species, sexual selection influences plumage coloration, with brighter or more ornate displays in males enhancing mating success. While social monogamy is the norm, genetic studies in certain species, such as the feral pigeon (Columba livia), indicate occasional extra-pair copulations, suggesting a degree of promiscuity within otherwise stable pairs. Breeding patterns vary geographically: in tropical regions, many species breed year-round due to consistent food availability, while in temperate zones, reproduction is seasonal, often aligned with spring and summer photoperiods and resource peaks. Nests are typically flimsy platforms or shallow cups constructed from twigs and stems, built collaboratively by both parents over 2-4 days in sites such as tree branches, cliffs, or ground cover; these are vigorously defended against intruders. Clutch sizes are usually 1-2 unmarked white or buff eggs, with frugivorous species tending toward single-egg clutches to match energetic demands. Incubation lasts 14-19 days on average, shared biparentally, with males often taking daytime shifts and females nighttime; both sexes develop brood patches for efficient . Parental care is intensive and biparental, with both adults producing nutrient-rich —a regurgitated secretion high in proteins, fats, and water (approximately 75-77% water content)—to feed altricial squabs for the first 5-10 days post-hatching, providing essential immunity and rapid growth. After this period, the diet shifts to regurgitated seeds and fruits, with fledging occurring at 3-6 weeks, though parents may continue provisioning for up to 40 days. Life history strategies emphasize iteroparity, with multiple clutches per year common (up to 6 or more in like the rock pigeon), enabling high reproductive output; semelparity is rare, and is reached at 6-12 months. These traits contribute to the family's adaptability, particularly in human-modified environments.

Social Structure and Communication

Columbidae exhibit a wide range of , from solitary pairs to highly gregarious flocks numbering in the thousands or more, depending on species and context. For instance, urban-adapted species like the rock dove (Columba livia) often form large communal roosts in cities, while the extinct (Ectopistes migratorius) gathered in enormous flocks estimated at up to 1.1 billion individuals during migrations, creating vast, sun-obscuring formations. Within flocks, dominance hierarchies are common, typically established through minimal and influenced by factors such as body size, age, and experience, which help regulate access to resources like and perches. These hierarchies remain stable over time in species like homing pigeons, reducing physical conflicts and promoting group cohesion without frequent challenges. , where non-parents assist in rearing young, is rare across the family. Communication in Columbidae relies on multiple modalities to convey information about danger, , and group coordination. Vocalizations include characteristic coos, whistles, and croaks, with alarm calls serving to alert flock members to predators and prompt evasive maneuvers. Visual signals encompass displays such as wing-spreading and tail fanning during threats or interactions, while tactile behaviors like gentle billing reinforce social bonds within groups. Some species engage in duets, combining vocal and visual elements for synchronized signaling. Flocking provides key advantages, including enhanced vigilance during to detect predators early and improved coordination for migration, where groups achieve higher speeds and more direct paths compared to solitary individuals. In homing pigeons, for example, flock flight reduces resting time and circling, boosting overall efficiency. The Eurasian collared-dove (Streptopelia decaocto) forms flocks that aid in locating mates and resources in open habitats. Social variations occur across the family, with many species showing territoriality during breeding seasons to defend resources, contrasted by nomadic or migratory patterns in non-breeding periods for species like the , which wandered vast distances in search of food. Sedentary urban doves, such as the rock dove, maintain year-round loose aggregations, while tropical species may form smaller, more fluid groups adapted to stable environments.

Conservation and Human Relations

Conservation Status and Threats

The family Columbidae encompasses 344 species assessed by the IUCN Red List, of which 71 (≈21%) are classified as threatened with extinction, including 7 critically endangered (CR), 22 endangered (EN), and 42 vulnerable (VU). Notable examples among the CR species include the black-faced cotinga (Conioptilon mcilhennyi), though many are island endemics. One species, the (Ectopistes migratorius), is extinct, having been driven to oblivion by overhunting and loss in , with the last individual dying in captivity in 1914. The (Zenaida graysoni) is classified as (EW), with approximately 150 individuals in captivity and reintroduction efforts planned, following historical degradation and predation on . Primary threats to Columbidae species include habitat loss and fragmentation, primarily from and , which affect the majority of threatened taxa and are implicated in nearly all cases according to older analyses of threat drivers. Hunting for and sport remains a significant pressure, particularly in and where species like the yellow-eyed pigeon (Columba eversmanni) have experienced sharp declines. On islands, such as rats, cats, and competing birds pose acute risks through predation and resource competition, exacerbating vulnerability for endemics. Additionally, 26 species are listed under appendices for regulation. Population trends indicate widespread declines across the family, driven by these anthropogenic pressures, while a few widespread species like the rock pigeon (Columba livia) continue to expand globally as invasives. Vulnerability is heightened for island endemics, where approximately 78% of threatened Columbidae occur, owing to their small ranges and isolation; additionally, the family's generally low —typically one to two chicks per and delayed maturity—impedes recovery from perturbations. Monitoring efforts have intensified since the , leveraging platforms like eBird to track abundance and distribution changes at fine scales, complemented by for migratory and to inform threat assessments and trend analyses.

Domestication and Cultural Significance

The domestication of the rock pigeon (Columba livia), the progenitor of most domestic pigeons, began around 5,000 BCE in the , particularly in , where archaeological evidence from tablets indicates early use for food and messaging purposes. Over millennia, humans selectively bred these birds, leading to diverse varieties; by the , focused breeding for racing emerged in around 1818, with birds achieving average speeds of up to 100 km/h in competitions. This selective process, highlighted in Charles Darwin's observations of phenotypic variation, transformed the rock pigeon into a model for artificial selection. Domestic pigeon breeds serve multiple utilities, including meat production, communication, and ornamentation. For meat, utility breeds like the White King and French Mondain are raised specifically for —the tender young pigeons harvested at 26-32 days old—providing a high-protein source in various cultures. Homing pigeons, derived from C. livia, excelled in long-distance messaging; during , they carried thousands of messages for Allied forces, including the U.S. military, often under hazardous conditions, saving countless lives. Ornamental breeds, such as the Fantail with its distinctive fan-shaped tail of 30-40 feathers, originated in regions like and , bred for aesthetic display in shows and gardens since ancient times. Pigeons and doves hold profound cultural symbolism worldwide, often representing , , and renewal. In , the white dove emerged as an emblem of from the biblical story of , where a dove returned with an signaling the flood's end and divine reconciliation. This imagery extends to broader traditions, with doves symbolizing due to their monogamous mating. In , pigeon flying competitions known as kabootarbazi or ishkbaazi are held seasonally from December to February, particularly in cities like Hyderabad, where enthusiasts release birds to compete in endurance flights, blending with . In modern times, escaped domestic pigeons have established widespread feral populations, thriving in urban environments globally and occasionally hybridizing with wild congeners. The pigeon racing industry generates significant economic value worldwide, driven by high-stakes competitions, bird sales, and breeding in regions like China and Belgium. However, ethical concerns persist, including welfare issues in racing such as high mortality from exhaustion, predation, and stress—up to 20-30% loss per race in some cases—and the culling of underperforming birds. Management of invasive feral populations often involves humane controls like fertility inhibitors to balance urban coexistence without lethal methods.

References

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