|Adult C. l. intermedia in India|
|approximate native range introduced non-native populations|
Wild Rock Doves are pale grey with two black bars on each wing, while domestic and feral pigeons are very variable in colour and pattern. There are few visible differences between males and females. The species is generally monogamous, with two squeakers (young) per brood. Both parents care for the young for a time.
Habitats include various open and semi-open environments. Cliffs and rock ledges are used for roosting and breeding in the wild. Originally found wild in Europe, North Africa, and western Asia, feral pigeons have become established in cities around the world. The species is abundant, with an estimated population of 17 to 28 million feral and wild birds in Europe.
- 1 Taxonomy and naming
- 2 Description
- 3 Distribution and habitat
- 4 Reproduction
- 5 Predators
- 6 Parasites
- 7 Human health
- 8 Domestication
- 9 Feral pigeon
- 10 Stages of life cycle
- 11 Osmoregulation
- 12 Thermoregulation
- 13 References
- 14 External links
Taxonomy and naming
The Rock Dove was first described by Gmelin in 1789. The genus name Columba is the Latinized form of the Ancient Greek κόλυμβος (kolumbos), "a diver", from κολυμβάω (kolumbao), "dive, plunge headlong, swim". Aristophanes (Birds, 304) and others use the word κολυμβίς (kolumbis), "diver", for the name of the bird, because of its swimming motion in the air. The specific epithet is derived from the Latin livor, "bluish". Its closest relative in the Columba genus is the Hill Pigeon, followed by the other rock pigeons: the Snow, Speckled and White-collared Pigeons.
The species is also known as the Rock Pigeon or Blue Rock Dove, the former being the official name from 2004 to 2011, at which point the IOC changed their official listing to its original British name of Rock Dove. In common usage, this bird is still often simply referred to as the "pigeon". Baby pigeons are called squabs.
There are 12 subspecies recognised by Gibbs (2000); some of these may be derived from feral stock.
- C. l. livia, the nominate subspecies, occurs in western and southern Europe, northern Africa, and Asia to western Kazakhstan, the northern Caucasus, Georgia, Cyprus, Turkey, Iran, and Iraq.
- C. l. atlantis (Bannerman, 1931) of Madeira, the Azores and Cape Verde, is a very variable population with chequered upperparts obscuring the black wingbars, and is almost certainly derived from feral pigeons.
- C. l. canariensis (Bannerman, 1914) of the Canary Islands, is smaller and averages darker than the nominate subspecies.
- C. l. gymnocyclus (Gray, 1856) from Senegal and Guinea to Ghana, Benin and Nigeria is smaller and very much darker than nominate C. l. livia. It is almost blackish on the head, rump and underparts with a white back and the iridescence of the nape extending onto the head.
- C. l. targia (Geyr von Schweppenburg, 1916) breeds in the mountains of the Sahara east to Sudan. It is slightly smaller than the nominate form, with similar plumage, but the back is concolorous with the mantle instead of white.
- C. l. dakhlae (Richard Meinertzhagen, 1928) is confined to the two oases in central Egypt. It is smaller and much paler than the nominate subspecies.
- C. l. schimperi (Bonaparte, 1854) is found in the Nile Delta south to northern Sudan. It closely resembles C. l. targia, but has a distinctly paler mantle.
- C. l. palaestinae (Zedlitz, 1912) occurs from Syria to Sinai and Arabia. It is slightly larger than C. l. schimperi and has darker plumage.
- C. l. gaddi (Zarodney & Looudoni, 1906), breeds from Azerbaijan and Iran east to Uzbekistan is larger and paler than C. l. palaestinae with which it intergrades in the west. It also intergrades with the next subspecies to the east.
- C. l. neglecta (Hume, 1873), is found in the mountains of eastern Central Asia. It is similar to the nominate subspecies in size, but is darker with a stronger and more extensive iridescent sheen on the neck. It intergrades with the next race in the south.
- C. l. intermedia (Strickland, 1844) occurs in Sri Lanka and in India south of the Himalayan range of C. l. neglecta. It is similar to that subspecies, but darker with a less contrasting back.
- C. l. nigricans (Buturlin, 1908) in Mongolia and north China is variable and probably derived from feral stock.
The adult of the nominate subspecies of the Rock Dove is 29 to 37 cm (11 to 15 in) long with a 62 to 72 cm (24 to 28 in) wingspan. Weight for wild or feral Rock Doves ranges from 238–380 g (8.4–13.4 oz), though overfed domestic and semi-domestic individuals can exceed normal weights. It has a dark bluish-gray head, neck, and chest with glossy yellowish, greenish, and reddish-purple iridescence along its neck and wing feathers. The iris is orange, red or golden with a paler inner ring, and the bare skin round the eye is bluish-grey. The bill is grey-black with a conspicuous off-white cere, and the feet are purplish-red. Among standard measurements, the wing chord is typically around 22.3 cm (8.8 in), the tail is 9.5 to 11 cm (3.7 to 4.3 in), the bill is around 1.8 cm (0.71 in) and the tarsus is 2.6 to 3.5 cm (1.0 to 1.4 in).
The adult female is almost identical to the male, but the iridescence on the neck is less intense and more restricted to the rear and sides, while that on the breast is often very obscure.
The white lower back of the pure Rock Dove is its best identification character; the two black bars on its pale grey wings are also distinctive. The tail has a black band on the end and the outer web of the tail feathers are margined with white. It is strong and quick on the wing, dashing out from sea caves, flying low over the water, its lighter grey rump showing well from above.
Young birds show little lustre and are duller. Eye colour of the pigeon is generally orange but a few pigeons may have white-grey eyes. The eyelids are orange in colour and are encapsulated in a grey-white eye ring. The feet are red to pink.
When circling overhead, the white underwing of the bird becomes conspicuous. In its flight, behaviour, and voice, which is more of a dovecot coo than the phrase of the Wood Pigeon, it is a typical pigeon. Although it is a relatively strong flier, it also glides frequently, holding its wings in a very pronounced V shape as it does. Though fields are visited for grain and green food, it is often not plentiful enough as to be a viewed as pest.
Pigeons feed on the ground in flocks or individually. They roost together in buildings or on walls or statues. When drinking, most birds take small sips and tilt their heads backwards to swallow the water. Pigeons are able to dip their bills into the water and drink continuously without having to tilt their heads back. When disturbed, a pigeon in a group will take off with a noisy clapping sound.
Pigeons, especially homing or carrier breeds, are well known for their ability to find their way home from long distances. Despite these demonstrated abilities, wild Rock Doves are sedentary and rarely leave their local areas.
Distribution and habitat
The Rock Dove has a restricted natural resident range in western and southern Europe, North Africa, and into South Asia. The Rock Dove is often found in pairs in the breeding season but is usually gregarious. The species (including ferals) has a large range, with an estimated global extent of occurrence of 10,000,000 km2 (3,900,000 sq mi). It has a large global population, including an estimated 17–28 million individuals in Europe. Fossil evidence suggests the Rock Dove originated in southern Asia and skeletal remains unearthed in Israel confirm their existence there for at least three hundred thousand years. However, this species has such a long history with humans that it is impossible to tell exactly where the species' original range was. Its habitat is natural cliffs, usually on coasts. Its domesticated form, the feral pigeon, has been widely introduced elsewhere, and is common, especially in cities, over much of the world. A Rock Pigeon's lifespan is anywhere from 3–5 years in the wild to 15 years in captivity, though longer-lived specimens have been reported. The main causes of mortality in the wild are predators and persecution by humans. The species was first introduced to North America in 1606 at Port Royal, Nova Scotia.
The Rock Dove breeds at any time of the year, but peak times are spring and summer. Nesting sites are along coastal cliff faces, as well as the artificial cliff faces created by apartment buildings with accessible ledges or roof spaces.
The nest is a flimsy platform of straw and sticks, laid on a ledge, under cover, often on the window ledges of buildings. Two white eggs are laid; incubation is shared by both parents lasting from seventeen to nineteen days. The newly hatched squab (nestling) has pale yellow down and a flesh-coloured bill with a dark band. For the first few days, the baby squab is tended and fed (through regurgitation) exclusively on "crop milk" (also called "pigeon milk" or "pigeon's milk"). The pigeon milk is produced in the crops of both parents in all species of pigeons and doves. The fledging period is about 30 days.
With only its flying abilities protecting it from predation, rock pigeons are a favorite almost around the world for a wide range of raptorial birds. In fact, with feral pigeons existing in most every city in the world, they may form the majority of prey for several raptor species who live in urban areas. Peregrine Falcons and Eurasian Sparrowhawks are natural predators of pigeons that are quite adept at catching and feeding upon this species. Up to 80% of the diet of Peregrine Falcons in several cities that have breeding falcons is composed of feral pigeons. Some common predators of feral pigeons in the North America are Opossums, Raccoons, Red-tailed Hawks, Great Horned Owls, Eastern Screech Owls and Accipiters. The birds that predate pigeons in North America can range in size from American Kestrels to Golden Eagles and can even include gulls, crows, and ravens. On the ground the adults, their young and their eggs are at risk from feral and domestic cats. Doves and pigeons are considered to be game birds as many species have been hunted and used for food in many of the countries in which they are native.
|Tinaminyssus melloi, a nasal mite.||Pigeon louse fly (P. canariensis), a blood-sucking ectoparasite.|
Pigeons may harbour a diverse parasite fauna. They often host the intestinal helminths Capillaria columbae and Ascaridia columbae. Their ectoparasites include the Ischnoceran lice Columbicola columbae, Campanulotes bidentatus compar, the Amblyceran lice Bonomiella columbae, Hohorstiella lata, Colpocephalum turbinatum, the mites Tinaminyssus melloi, Dermanyssus gallinae, Dermoglyphus columbae, Falculifer rostratus, and Diplaegidia columbae. The hippoboscid fly Pseudolynchia canariensis is a typical blood-sucking ectoparasite of pigeons, found only in tropical and sub-tropical regions.
Pigeons have been falsely associated with the spread of human diseases.[verification needed] Contact with pigeon droppings poses a minor risk of contracting histoplasmosis, cryptococcosis, and psittacosis, and exposure to both droppings and feathers can produce bird fancier's lung. Pigeons are not a major concern in the spread of West Nile virus; though they can contract it, they do not appear to be able to transmit it. Pigeons are, however, at potential risk for carrying and spreading avian influenza. Although one study has shown that adult pigeons are not clinically susceptible to the most dangerous strain of avian influenza, the H5N1, other studies have presented definitive evidence of clinical signs and neurological lesions resulting from infection. Furthermore, it has been shown that pigeons are susceptible to other strains of avian influenza, such as the H7N7, from which at least one human fatality has been recorded.
Rock Doves have been domesticated for several thousand years, giving rise to the domestic pigeon (Columba livia domestica). As well as food and pets, domesticated pigeons are used as homing pigeons. They were in the past also used as carrier pigeons, and so-called war pigeons have played significant roles during wartime, with many pigeons having received bravery awards and medals for their services in saving hundreds of human lives: including, notably, the British pigeon Cher Ami who received the Croix de Guerre for her heroic actions during World War I, and the Irish Paddy and the American G.I. Joe, who both received the Dickin Medal, amongst 32 pigeons to receive this medallion, for their gallant and brave actions during World War II. There are numerous breeds of fancy pigeons of all sizes, colours and types.
Many domestic birds have escaped or been released over the years, and have given rise to the feral pigeon. These show a variety of plumages, although some have the blue barred pattern as does the pure Rock Dove. Feral pigeons are found in large numbers in cities and towns all over the world. The scarcity of the pure wild species is partly due to interbreeding with feral birds.
Stages of life cycle
Water is taken in by the Columba livia directly by drinking water or indirectly from the food they ingest. They drink water through a process called double-suction mechanism. The daily diet of the Pigeon places many physiologically challenges it must over come through osmoregulation. Protein intake for example causes an excess toxins of amine groups when it is broken down for energy. To regulate this excess and secrete these unwanted toxins the Columba livia must remove the amine groups as uric acid. Nitrogen excretion through uric acid can be considered an advantage because it doesn't require a lot of water and isn't very soluble, but producing it takes more energy because of its complex molecular composition.
The danger of desiccation is a major threat to animals living on land. Water is lost in urine and feces, but evaporation is the principal route of water loss. Water lost must be replaced by drinking and water in food. Dehydration or salt-loading decreases the filtration rate primarily by the shut down of the nephrons, which is controlled by an antidiuretic hormone, arginine vasotocin. Pigeons adjust their drinking rates and food intake in parallel and when adequate water is unavailable for excretion, food intake is limited to maintain water balance. As Columbia livia inhabit arid environments, research attributes this to their strong flying capabilities to reach the available water sources, not because of exceptional potential for water conservation. Columba livia kidneys, like mammalian kidneys, are capable of producing urine hyperosmotic to the plasma utilizing the processes of filtration, reabsorption and secretion, which will be discussed later and explained through the Starling-Landis Hypothesis. The medullary cones function as countercurrent units that achieve the production of hyperosmotic urine. Hyperosmotic urine can be understood in light of the law of diffusion and osmolarity.
Organ of osmoregulation
Unlike a number other bird species which have the salt gland as the primary osmoregulatory organ, Columba livia does not use their salt gland even though it exists. Columba livia uses the function of their kidneys to maintain homeostatic balance of ions such as sodium and potassium while preserving water quantity in the body. Filtration of the blood, reabsorption of ions and water, and secretion of uric acid are all components of the kidney's process. The kidneys of Columba livia are located in its pelvic region. Columba livia has two kidneys that are coupled, each having three partially separate lobes; the posterior lobe is the largest in size. Like mammalian kidneys, the avian kidney contains a medullary region and a cortical region. Peripherally located around the cortical region, the collecting ducts gather into cone-like ducts, medullary cones, which converge into the ureters. There are two types of nephrons in the kidney; nephrons that are located in the cortex and do not contain the loop of Henle are called loopless nephrons, the other type are called looped or mammalian nephrons. Looped nephrons contain the loop of Henle that continue down into the medulla then enter the distal tubule drain towards the ureter. Mammals generally have a more vascularized glomeruli than the nephrons in birds. The nephrons of avian species can not produce urine that is hyperosmotic to the blood, but, the loop of Henle utilizes countercurrent multiplication which allows it to become hyperosmotic in the collecting duct. This alternation of permeability between different sections of the ascending and descending loop allows for the elevation of the urine osmotic pressure 2.5 times above the blood osmotic pressure.
Specialize cell types involved in osmoregulation
The integumentary system functions in osmoregulation by acting as a barrier between the extracellular compartment and the environment to regulate water gain and loss, as well as solute flux. The permeability of the integument to water and solutes varies from animal to animal.The excretory system is responsible for regulating water and solute levels in the body fluids. Pigeons can produce hyperosmotic urine but their renal system is different from other animals. They do not produce concentrated urine to reduce water loss but produce a whitish part called urate. It is considered as uric acid solid crystals and it is less toxic than urea. The wastes move from the blood of the peritubular capillaries passes through the tubule cells and into the collecting ducts and transported as urate (uric acid). Urate is then transported to the cloaca and from there to the large intestine where uric acid particle and water and solutes in the urine can be reabsorbed and balanced. Thus this allows them to save their body water instead of excreting large volume of dilute urea. Cells of the proximal tubule have numerous microvilli and mitochondria which provide surface area and energy to the proximal tubule cells.
The blood pH is regulated by the A and B types of cells located in distal tubule and collecting duct. The A type cells are acid secreting cells that have a proton ATPase in the apical membrane and a Cl-/ HCO3- exchange system in the basolateral membrane whereas, the B type cells are base secreting cells, which secrete bicarbonate into the lumen of the tubule in exchange for chloride ions. The regulation of pH in blood determines whether bicarbonate is reabsorbed or secreted.
Transport mechanisms of osmoregulation
The filtrate contains lots of important substances. In the proximal tubules of the Columbia livia kidney, substances that are needed, such as vitamins and glucose are reabsorbed into the blood. Their kidney has a variety of ion channels involved in salt and water transport. Water is reabsorbed through aquaporins which are present in the lumen of proximal tubule, basolateral membrane, and blood vessel near proximal tubule. Water flows from the epithelial cells into the blood via osmosis. Since osmosis occurs, the osmolarity of the filtrate remains isotonic. Sodium/Potassium/ATPase transporter is located in the basolateral membrane of the epithelial cell, which is opposite of the lumen of proximal tubule, and actively pumps sodium out of the cell into the blood.
Eggshell's gas exchange and water loss
Gas exchange across eggshells results in water loss from the egg. However, the egg must retain enough water to hydrate the embryo. This results in the knowledge that changing temperatures and humidity can affect the eggshell's architecture. Behavioral adaptations in Columba livia and other birds, such as the incubation of their eggs, can help with the effects of these changing environments. It was found that eggshell architecture undergoes selection decoupled from behavioral effects, and that humidity may be a driving selective pressure. Low humidity requires enough water to keep the embryo from desiccation, and high humidity needs enough water loss to facilitate the initiation of pulmonary respiration. The water loss from the eggshell is directly linked to the growth rate of the species. The ability of the embryo to tolerate extreme water loss is due to the parental behavior in species colonizing in different environments. Studies have been done showing that wild habitats of Columba livia and other birds have a higher rate tolerance of various humidity levels, but Columba livia do prefer areas where the humidity closely matched their native breeding conditions. The pore areas of the shells allow water to diffuse in and out of the shell, preventing the possible harming of the embryo due to the high rates of water retention. If an eggshell is thinner, it can cause a decrease in pore length, and an increase in conductance and pore area. A thinner eggshell can also cause a decrease in mechanical restriction of the embryo.
The Columbia Livia is habituated within many vast environments with varying degrees of temperatures. Like all vertebrates, Columbia Livia perspires heat through evaporation of water when temperatures are high in the environment. It’s preferred niche temperature ranges between +39 - +42 degrees Celsius.
Peripheral thermoreceptors of the Columbia Liva regulate its body’s response to the cold. During low temperatures, which put the Columbia Liva’s body under stress it accommodates extreme temperatures by increasing its internal temperatures within the core and spinal cord. Along with this increase, there is also a decrease in temperature within the legs, neck and back skin.
Physiological challenges placed on organism
Columba Livia stabilize their internal body temperature independent of alteration in ambient temperature. They are also able to withstand extreme climate conditions, such as ambient temperature range of +42 to -40°C. The temperature regulation of Columba Livia is generally based on the principle of endotherms. Being endothermic they use metabolic heat to raise body temperature. Columba Livia are also homeotherms, meaning that they are thermoregulators and maintain a relatively constant body temperature. The heat exchange between animals and their surroundings occurs due to conduction, convection, radiation and evaporation. Fourier's Law of Heat Conduction describes the loss of heat experienced by animals through conduction. At low ambient temperatures the endothermic animals are able to reduce their heat loss by lowering the skin temperature and by increasing their peripheral insulation, which is discussed later.
Columba Livia does a few things to regulate its body temperature. Normally it will drink water after they have eaten, but when stressed by heat they can drink whenever needed to lower its body temperature. Another way it can regulate its heat is through Ptilomotor responses. Ptilomotor responses allow for better insulation of the body, because smooth muscle contractions make the feathers stand up straighter, which traps more air next to the skin. Columba Livia exhibits Ta (ambient temperature) selecting behavior. It will seek out its desired thermal neutral zone temperatures, in order to expend less energy heating and cooling its body.
Physiological changes to blood flow
Areas poorly or not insulated by feathers such as the beak, head, and feet have vasomotor responses. To reduce heat loss while in cold atmospheric temperatures, endothermic animals will lower the skin temperature by restricting the amount of blood that reaches it, called vasoconstriction. The sympathetic nervous system stimulates the constriction of the vascular beds at low temperatures. Vasodialation does the opposite; to increase the heat lost by convection after high muscular activity or from heat stress, Columba Livia increases its blood flow to the surface of its body. Cutaneous tissue of the beak, feet, and bends in the wings are dilated. To regulate brain temperature it uses the vascular vessels(plexus) in the eyes, in combination with vasomotion. Evaporation is usually controlled by sweat glands, however, birds use their breathing pattern to control heat dissipation. The frequency in breathing depends on body temperature, Tb; to increase respiratory evaporation the bird's breathing rate would increase. The most important thermoregulatory mechanism is called shivering thermogenesis. The skeletal muscles are used to generate heat through contractions when the surrounding air, Ta, is below its thermal neutral zone. As the temperature drops, the shivering increases to generate more heat. Non-shivering thermogenesis is used by Columba Livia, when exposed to cold to generate heat; an increase in Na+/K+-ATPase activity drives this mechanism in the liver.
A study was done by Michael E. Rashotte, et al. (1998) comparing the vigilance states and body temperature is different within in fed and fasted pigeons (Columba Livia). Fasting induces nocturnal hypothermia in pigeons. There are different sleep patterns associated with heat production in pigeons, slow wave sleep (SWS) and paradoxical sleep (PS). An increase of SWS and PS was compared to the fasting-induced nocturnal hypothermia by comparing body temperature (Tb) and vigilance states when pigeons were fed and fasted. It was found that the Tb was decreasing near the beginning of the dark phase and that the time spent in SWS and PS was elevated in the fasting pigeons due to the increase of frequency and duration. When body temperature was low in the middle of the dark phase, it showed that SWS was elevated but it did not affect the PS stage. When the body temperature was high during the last hours of dark, SWS remained elevated in fasting-induced and that PS was relatively high. Rashotte, et al. (1998) suggests that more evidence is needed to confirm these results but he suggests that pigeons may be best viewed as an animal that has a shallow hypometabolic state that fall within (or very close to) their euthermic range. It is also seen that a pigeon’s vigilance stage can be compared similarly to mammals in hibernation.
Specialize organs or anatomy involved in thermoregulation
The purpose of thermoregulation is to maintain body temperature by producing heat through physiological and metabolic reactions. Heat gain should equal to rates of heat loss. If the body temperature is unbalanced, the animal becomes either warmer or colder. Heat production in birds is associated to shivering. The large flight muscles- pectoralis as well as the leg muscles generate heat by shivering.
Columba Livia have strong wings with flexible feathers which provide enough insulation to keep their body warm and dry. The fat layers and feathers reduce the flow of heat between an animal and its environment and lower the energy cost of keeping warm. In some birds the heat loss from the legs and feet is limited in cold weather because of a countercurrent mechanism that saves heat and in hot weather it can serve as heat radiators which increase blood flow.
Thermoregulation in birds requires cooling as well as warming. At low temperature birds can tuck head and neck under their wings to reduce heat loss. The heat is lost by the pigeons as an insensible heat by evaporation of water from the respiratory system and skin when temperature gradient is less and relative humidity is low. At the relatively high temperature birds increase their respiration rate to increase their cooling by evaporation. The panting is important in birds which involves gular flutter. The pouch richly supplied with blood vessels in the floor of the mouth; the rapid movement of the upper throat tissues - fluttering the pouch increases evaporation. Pigeons can use evaporative cooling to keep body temperature close to 40°C in air temperatures as high as 60°C, as long as they have sufficient water.
Also from previous studies experiment shows that a bird is capable of evaporating enough water from the cloaca for thermoregulation and results suggests that some birds’ cloacal evaporation can be controlled and could serve as an important maneuver for thermoregulation at high ambient temperatures.
Regulation of metabolism
Columba Livia as homeothermic animals, are able to regulate heat production and external heat loss in autonomic ways, by a feedback control system. Negative feedback is the most important principle for regulation; a decrease of ambient temperature evoked by cold activates some thermoregulatory effector mechanisms, which reduce the heat loss and increase the internal heat production. The metabolic rate of resting Columba Livia at neutral ambient temperature, is reduced by a level of 5-10% during drowsiness, sleep and darkness. An increase follows every kind of muscle activity, such as flying, which increases metabolic rate by 10-12 times. Heat production throughout the day contributes to a high level of body temperature.
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- Rashotte, Michael (1998). "Vigilance states and body temperature during the circadiancycle in fed and fasted pigeons (Columba livia)". St.Petersburg, Russia.
- Rashotte, Michael (1998). "Vigilance states and body temperature during the circadian cycle in fed and fasted pigeons (Columba livia)". St.Petersburg, Russia.
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- Bird energy balance. (n.d.). Retrieved from http://people.eku.edu/ritchisong/birdmetabolism.html
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|Wikimedia Commons has media related to Columba livia.|
|Wikispecies has information related to: Columba livia|
- Ageing and sexing (PDF; 3.4 MB) by Javier Blasco-Zumeta & Gerd-Michael Heinze
- Rock Dove videos, photos, and sounds at the Internet Bird Collection
- Rock Pigeon photo gallery at VIREO (Drexel University)