Columbidae adalah satu keluarga burung yang terdiri daripada merpati dan punai (diistilahkan sebagai pigeon dan dove dalam bahasa Inggeris). Ia mempunyai badan bulat, leher pendek, paruh bengkok pendek dan pada beberapa spesies mempunyai cere berdaging. Makanan mereka terdiri daripada bijiran, buah-buahan dan tumbuhan. Keluarga burung Columbidae boleh ditemui di seluruh dunia tetapi kepelbagaian spesies boleh didapati di ekozon Indomalaya dan Australasia.
Order Columbiformes yang merangkumi merpati dan punai ini termasuk sekitar 42 genera dan 310 spesies.
Perbezaan antara "dove" (punai) dan "pigeon" (merpati) tidak konsisten dalam bahasa Inggeris dan tidak wujud dalam kebanyakan bahasa-bahasa lain. Berlawanan dalam bidang saintifik dan ornitologi, di mana "dove" cenderung merujuk pada spesies yang lebih kecil manakala "pigeon" untuk yang lebih besar; tetapi istilah ini juga tidak digunakan secara konsisten. Dari segi sejarah, nama-nama biasa bagi burung-burung ini melibatkan kesan perbezaan yang jauh. Burung ini biasanya dirujuk sebagai "pigeon" (merpati) adalah merpati domestik, di mana menjadi kebiasaan di kebanyakan bandar-bandar dengan nama merpati feral.
Pigeon (merpati) merupakan sebuah perkataan Perancis yang diterbitkan daripada perkataan Latin pipio, merujuk pada seekor anak burung "mengintai",[2] sementara dove (punai) adalah sebuah perkataan Jermanik yang merujuk pada "penerbangan junam" yang dimiliki oleh spesies burung ini.[3] The English dialectal word "culver" appears to derive from Latin columba.[2]
Merpati dan punai membina sarang nipis, sering kali menggunakan ranting kayu atau sampah sarap lain, yang mana terletak di atas pokok, belebas atau tanah bergantung pada spesies. Ia bertelur sebiji atau dua biji pada suatu masa, dan kedua-dua ibu dan bapa akan menjaga anaknya, yang mana akan meninggalkan sarang selepas 7–28 hari.[4] Tidak seperti kebanyakan burung-burung lain, kedua-dua jantina merpati dan punai akan menghasilkan "susu tembolok" untuk diberi kepada anaknya. Susu ini dihasilkan oleh satu kulit yang dipenuhi sel-sel cecair daripada lapisan tembolok. Merpati dan punai muda dipanggil "squab".
Keluarga Columbidae telah diperkenalkan oleh ahli zoologi Inggeris William Elford Leach dalam satu panduan kandungan Muzium British yang diterbitkan pada tahun 1820.[5][6] Columbidae merupakan satu-satunya keluarga hidup dalam order Columbiformes. Sandgrouse (Pteroclidae) dahulunya diletakkan di sini, sebelum dipindahkan ke order Pterocliformes berlainan berdasarkan perbezaan anatomi (contohnya ia tidak boleh minum dengan cara "menghisap" atau "mengepam");[7] mereka kini dianggap berkait rapat dengan burung pantai.[8] Kajian pilogenomik terkini menyokong pengumpulan burung merpati dan sandgrouse bersama-sama, begitu juga mesite, membentuk takson dalam Mirandornithes.[9][10][11][12]
Columbidae biasanya dibahagikan kepada lima sub-keluarga, berkemungkinan tidak berapa tepat.[13] Contohnya, punai tanah dan punai-puyuh Amerika (Geotrygon), di mana biasanya diletakkan dalam genus Columbinae, kelihatan adalah dua sub-keluarga yang berbeza.[14] The order presented here follows Baptista et al. (1997),[15] with some updates.[16][17][18]
The arrangement of genera and naming of subfamilies is in some cases provisional because analyses of different DNA sequences yield results that differ, often radically, in the placement of certain (mainly Indo-Australian) genera.[petikan diperlukan] This ambiguity, probably caused by long branch attraction, seems to confirm the first pigeons evolved in the Australasian region, and that the "Treronidae" and allied forms (crowned and pheasant pigeons, for example) represent the earliest radiation of the group.[petikan diperlukan]
The family Columbidae previously also contained the family Raphidae, consisting of the extinct Rodrigues solitaire and the dodo.[18][19][20] These species are in all likelihood part of the Indo-Australian radiation that produced the three small subfamilies mentioned above,[21] with the fruit doves and pigeons (including the Nicobar pigeon). Therefore, they are here included as a subfamily Raphinae, pending better material evidence of their exact relationships.[22]
Exacerbating these issues, columbids are not well represented in the fossil record.[23] No truly primitive forms have been found to date.[petikan diperlukan] The genus Gerandia has been described from Early Miocene deposits in France, but while it was long believed to be a pigeon,[24] it is now considered a sandgrouse.[25] Fragmentary remains of a probably "ptilinopine" Early Miocene pigeon were found in the Bannockburn Formation of New Zealand and described as Rupephaps;[25] "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).[26] Apart from that, all other fossils belong to extant genera.[27]
Taksonomi berdasarkan hasil kerja oleh John H. Boyd, III,[28] seorang professor ekonomi.[29]
Starnoenas (Blue-headed partridge-dove)
Geotrygon (punai-puyuh)
Leptotrygon (punai-puyuh belakang zaitun)
†Ectopistes (Tekukur Passenger)
Patagioenas (Merpati Amerika)
Macropygia (Tekukur Api)
Streptopelia (Tekukur Berkolar)
Columba (Merpati Dunia Lama)
Uropelia (punai tanah ekor panjang)
Gallicolumba (Punai jantung berdarah)
Ocyphaps (Tekukur jambul)
Petrophassa (Pergam Batu)
Leucosarcia (Merpati Wonga)
Geopelia (Merbuk/ketitir)
Trugon (thick-billed ground pigeon)
†Microgoura (Choiseul crested pigeon)
Otidiphaps (pheasant pigeon)
Goura (Punai mahkota)
Didunculus (merpati paruh gigi)
Caloenas (Merpati Nicobar)
†Raphus (Dodo)
†Pezophaps (Rodrigues solitaire)
Treron (Punai Hijau)
Phapitreron (Punai perang)
Chalcophaps (Punai Tanah)
Oena (Namaqua dove)
Turtur (Punai kayu Afrika)
Ducula (Pergam)
[Ptilinopus; Drepanoptila; Alectroenas] (Punai Buah)
Hemiphaga (Punai New Zealand)
Lopholaimus (Punai Topknot)
Cryptophaps (Merpati Sombre)
Gymnophaps (Pergam Gunung)
Pigeons and doves exhibit considerable variation in size, ranging in length from 15 hingga 75 sentimeter (5.9 hingga 29.5 in), and in weight from 30 g (0.066 lb) to above 2,000 g (4.4 lb).[30] The largest species is the crowned pigeon of New Guinea,[31] which is nearly turkey-sized, at a weight of 2–4 kg (4.4–8.8 lb).[32] The smallest is the New World ground dove of the genus Columbina, which is the same size as a house sparrow, weighing as little as 22 g (0.049 lb).[15] The dwarf fruit dove, which may measure as little as 13 cm (5.1 in), has a marginally smaller total length than any other species from this family.[15] One of the largest arboreal species, the Marquesan imperial pigeon, currently battles extinction.[33]
Overall, the anatomy of Columbidae is characterized by short legs, short bills with a fleshy cere, and small heads on large, compact bodies.[34] Like some other birds, the Columbidae have no gall bladders.[35] 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.[36] In fact, however, they do have bile (as Aristotle had earlier realized), which is secreted directly into the gut.[37]
The wings are large, and have eleven primary feathers;[38] pigeons have strong wing muscles (wing muscles comprise 31–44% of their body weight[39]) and are among the strongest fliers of all birds.[38]
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.[40] 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.[41]
Columbidae have unique body feathers, with the shaft being generally broad, strong, and flattened, tapering to a fine point, abruptly.[38] In general, the aftershaft is absent; however, small ones on some tail and wing feathers may be present .[42] Body feathers have very dense, fluffy bases, are attached loosely into the skin, and drop out easily.[43] Possibly serving as a predator avoidance mechanism,[44] 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.[45] Granivorous species tend to have dull plumage, with a few exceptions, whereas the frugivorous species have brightly coloured plumage.[15] The Ptilinopus (fruit doves) are some of the brightest coloured pigeons, with the three endemic species of Fiji and the Indian Ocean Alectroenas being the brightest. Pigeons and doves may be sexually monochromatic or dichromatic.[46] In addition to bright colours, pigeons may sport crests or other ornamentation.[47]
Columbidae are excellent fliers due to the lift provided by their large wings, which results in low wing loading;[48] They are highly maneuverable in flight[49] 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.[50]
Merpati dan punai boleh ditemuai hampir di seluruh dunia kecuali kawasan paling kering di padang pasir Sahara, Antartika dan kawasan pulau-pulau di sekelilinginya, dan Artik.[30] Ia juga mendiami kebanyakan pulau-pulau di lautan dunia, mencapai timur Polynesia dan kepulauan Chatham di Pasifik, Mauritius, Seychelles dan Réunion di Lautan Hindi, dan Azores di Lautan Atlantik.
The family has adapted to most of the habitats available on the planet. These species may be arboreal, terrestrial, or semi-terrestrial. Various species also inhabit savannas, grasslands, deserts, temperate woodlands and forests, mangrove forests, and even the barren sands and gravels of atolls.[51]
Some species have large natural ranges. The eared dove ranges across the entirety of South America from Colombia to Tierra del Fuego,[52] the Eurasian collared dove has a massive (if discontinuous) distribution from Britain across Europe, the Middle East, India, Pakistan and China,[53] and the laughing dove across most of sub-Saharan Africa, as well as India, Pakistan, and the Middle East.[54] Other species have tiny, restricted distributions; this is most common in island endemics. The whistling dove is endemic to the tiny Kadavu Island in Fiji,[55] the Caroline ground dove is restricted to two islands, Truk and Pohnpei in the Caroline Islands,[56] and the Grenada dove is restricted to Grenada in the Caribbean.[57] Some continental species also have tiny distributions; for example, the black-banded fruit dove is restricted to a small area of the Arnhem Land of Australia,[58] the Somali pigeon is restricted to a tiny area of northern Somalia,[59] and Moreno's ground dove is restricted to the area around Salta and Tucuman in northern Argentina.[15]
The largest range of any species is that of the rock dove.[60] 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.[60] The range of the species increased dramatically upon domestication, as the species went feral in cities around the world.[60] The species 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.[60] The species is not the only pigeon to have increased its range due to the actions of man; several other species 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.[15]
Merpati jantan boleh berpasangan dengan betina lain.[61]
Bijiran dan buah-buahan membentuk komponen sumber makanan diet utama bagi merpati dan punai.[30][62] Faktanya, keluarga ini boleh dikelaskan kepada dua iaitu pemakan bijiran atau spesoes granivorous (sub-keluarga Columbinae) dan pemakan buah-buahan atau spesies frugivorous (empat sub-keluarga yang lain).[63] The granivorous species typically feed on seed found on the ground, whereas the frugivorous species tend to feed in trees.[63] There are morphological adaptations that can be used to distinguish between the two groups: granivores tend to have thick walls in their gizzards, intestines, and esophagi whereas the frugivores tend to have thin walls.[30] In addition, fruit-eating species have short intestines whereas those that eat seeds have longer ones.[64] Frugivores are capable of clinging to branches and even hang upside down to reach fruit.[15][63]
Selain daripada buahan dan bjiran, sejumlah jenis makanan lain juga dimakan oleh banyak spesies. Beberapa spesies terutamanya punai tanah dan punai-puyuh, memakan sejumlah makanan berdaging seperti serangga dan cacing.[63] Satu spesies, atoll fruit dove memakan serangga dan reptilia.[63] Siput, rama-rama dan serangga lain juga dimakan oleh white-crowned pigeon, orange fruit dove dan ruddy ground dove.[15]
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.[65] 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.[65]
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[65] even though it was once the most numerous species of bird on Earth.[petikan diperlukan] Its former numbers are difficult to estimate, but one ornithologist, Alexander Wilson, estimated one flock he observed contained over two billion birds.[66] 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.[67] 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.[petikan diperlukan]
The dodo, and its extinction, was more typical of the extinctions of pigeons in the past. Like many species that colonise remote islands with few predators, it lost much of its predator avoidance behaviour, along with its ability to fly.[68] 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 all the other island forms that have become extinct.[68]
Around 59 species of pigeons and doves are threatened with extinction today, about 19% of all species.[69] 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.[68] 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.[70] In some areas, a lack of knowledge means the true status of a species is unknown; the Negros fruit dove has not been seen since 1953,[71] 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.[72]
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.[68][73]
The pigeon was used in both World War I and II, notably by the Australian, French, German, American, and UK forces. They were also awarded with various laurels throughout, for their service. On 2 December 1943, three pigeons – Winkie, Tyke, and White Vision – were awarded the first ever Dickin medal, serving with Britain's Royal Air Force, for rescuing an air force crew during World War II.[74] Thirty-two pigeons have been decorated with the Dickin Medal, citing them for "brave service",[75] for war contributions, including Commando, G.I. Joe,[76] Paddy, Royal Blue, and William of Orange.[77]
Cher Ami, a homing pigeon in World War I, was awarded the Croix de Guerre Medal, by France, with a palm Oak Leaf Cluster for her service in Verdun.[78] Despite having almost lost a leg and being shot in the chest, she managed to travel around twenty-five miles to deliver the message that saved 194 men of the Lost Battalion of the 77th Infantry Division in the Battle of the Argonne, in October 1918.[78][74] When Cher Ami died, she was mounted and is part of the permanent exhibit at the National Museum of American History of the Smithsonian Institution.[79]
A grand ceremony was held in Buckingham Palace to commemorate a platoon of pigeons that braved the battlefields of Normandy to deliver vital plans to Allied forces on the fringes of Germany.[80] Three of the actual birds that received the medals are on show in the London Military Museum so that well-wishers can pay their respects.[80]
The rock pigeon has been domesticated for hundreds of years.[81] It has been bred into several varieties kept by hobbyists, of which the best known is the homing pigeon or racing homer.[81] Other popular breeds are tumbling pigeons such as the Birmingham roller, and fancy varieties that are bred for certain physical characteristics such as large feathers on the feet or fan-shaped tails. Domesticated rock pigeons are also bred as carrier pigeons,[47] used for thousands of years to carry brief written messages,[82] and release doves used in ceremonies.[83] White doves are also commonly used in magic acts.[84]
In ancient Mesopotamia, doves were prominent animal symbols of Inanna-Ishtar, the goddess of love, sexuality, and war.[85][86] Doves are shown on cultic objects associated with Inanna as early as the beginning of the third millennium BC.[85] Lead dove figurines were discovered in the temple of Ishtar at Aššur, dating to the thirteenth century BC,[85] and a painted fresco from Mari, Syria, shows a giant dove emerging from a palm tree in the temple of Ishtar,[86] indicating that the goddess herself was sometimes believed to take the form of a dove.[86] In the Epic of Gilgamesh, Utnapishtim releases a dove and a raven to find land; the dove merely circles and returns.[87] Only then does Utnapishtim send forth the raven, which does not return, and Utnapishtim concludes the raven has found land.[87]
In the ancient Levant, doves were used as symbols for the Canaanite mother goddess Asherah.[85][86][88] The ancient Greek word for "dove" was peristerá,[85][86] which may be derived from the Semitic phrase peraḥ Ištar, meaning "bird of Ishtar".[85] In classical antiquity, doves were sacred to the Greek goddess Aphrodite,[89][90][85][86] who absorbed this association with doves from Inanna-Ishtar.[86] Aphrodite frequently appears with doves in ancient Greek pottery.[89] The temple of Aphrodite Pandemos on the southwest slope of the Athenian Acropolis was decorated with relief sculptures of doves with knotted fillets in their beaks[89] and votive offerings of small, white, marble doves were discovered in the temple of Aphrodite at Daphni.[89] During Aphrodite's main festival, the Aphrodisia, her altars would be purified with the blood of a sacrificed dove.[91] Aphrodite's associations with doves influenced the Roman goddesses Venus and Fortuna, causing them to become associated with doves as well.[88]
In the Hebrew Bible, doves or young pigeons are acceptable burnt offerings for those who cannot afford a more expensive animal.[92] In Genesis, Noah sends a dove out of the ark, but it came back to him because the floodwaters had not receded. Seven days later, he sent it again and it came back with an olive branch in her mouth, indicating the waters had receded enough for an olive tree to grow. "Dove" is also a term of endearment in the Song of Songs and elsewhere. In Hebrew, Jonah (יוֹנָה) means dove.[93] The "sign of Jonas" in [1] is related to the "sign of the dove".[94]
Jesus's parents sacrificed doves on his behalf after his circumcision (Luke).[94] Later, the Holy Spirit descended upon Jesus at his baptism like a dove (Matthew), and subsequently the "peace dove" became a common Christian symbol of the Holy Spirit.[94]
In Islam, doves and the pigeon family in general are respected and favoured because they are believed to have assisted the final prophet of Islam, Muhammad, in distracting his enemies outside the cave of Thaw'r, in the great Hijra.[95] A pair of pigeons had built a nest and laid eggs at once, and a spider had woven cobwebs, which in the darkness of the night made the fugitives believe that Muhammad could not be in that cave.[95]
Beberapa spesies merpati dan punai digunakan sebagai sumber makanan; bagaimanapun hampir semua spesies boleh dimakan.[96] Merpati peliharaan atau buruan telah dijadikan sumber makanan sejak zaman Timur Tengah Tua, Rome Tua, dan Eropah Zaman Pertengahan.[75] Ia merupakan sumber daging biasa bagi Yahudi, Arab, Assam dan Perancis. Berdasarkan pada Tanakh, punai merupakan kosher, dan ia satu-satunya burung yang boleh digunakan sebagai seekor korban. Burung kosher lain boleh dimakan tetapi tidak sebagai seekor korban. Merpati juga digunakan dalam masakan Asia, seperti Cina dan Indonesia.
Di Eopah, wood pigeon biasanya ditembak sebagai daging buruan,[97] sementara Pergam Batu yang asalnya dibiakkan sebagai spesies makanan, kemudian dikembangkan banyak baka-baka lain di sebabkan kualiti dagingnya.[51] Kepupusan merpati Passenger di Amerika Utara sebahagiannya diakibatkan oleh pemburuan untuk digunakan sebagai sumber makanan.[98] Mrs Beeton's Book of Household Management mengandungi resipi-resipi merpati panggang dan pai merpati, sejenis makanan popular dan tidak mahal pada zaman industri Victoria di Britain.[99]
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Columbidae adalah satu keluarga burung yang terdiri daripada merpati dan punai (diistilahkan sebagai pigeon dan dove dalam bahasa Inggeris). Ia mempunyai badan bulat, leher pendek, paruh bengkok pendek dan pada beberapa spesies mempunyai cere berdaging. Makanan mereka terdiri daripada bijiran, buah-buahan dan tumbuhan. Keluarga burung Columbidae boleh ditemui di seluruh dunia tetapi kepelbagaian spesies boleh didapati di ekozon Indomalaya dan Australasia.
Order Columbiformes yang merangkumi merpati dan punai ini termasuk sekitar 42 genera dan 310 spesies.
Perbezaan antara "dove" (punai) dan "pigeon" (merpati) tidak konsisten dalam bahasa Inggeris dan tidak wujud dalam kebanyakan bahasa-bahasa lain. Berlawanan dalam bidang saintifik dan ornitologi, di mana "dove" cenderung merujuk pada spesies yang lebih kecil manakala "pigeon" untuk yang lebih besar; tetapi istilah ini juga tidak digunakan secara konsisten. Dari segi sejarah, nama-nama biasa bagi burung-burung ini melibatkan kesan perbezaan yang jauh. Burung ini biasanya dirujuk sebagai "pigeon" (merpati) adalah merpati domestik, di mana menjadi kebiasaan di kebanyakan bandar-bandar dengan nama merpati feral.
Pigeon (merpati) merupakan sebuah perkataan Perancis yang diterbitkan daripada perkataan Latin pipio, merujuk pada seekor anak burung "mengintai", sementara dove (punai) adalah sebuah perkataan Jermanik yang merujuk pada "penerbangan junam" yang dimiliki oleh spesies burung ini. The English dialectal word "culver" appears to derive from Latin columba.
Merpati dan punai membina sarang nipis, sering kali menggunakan ranting kayu atau sampah sarap lain, yang mana terletak di atas pokok, belebas atau tanah bergantung pada spesies. Ia bertelur sebiji atau dua biji pada suatu masa, dan kedua-dua ibu dan bapa akan menjaga anaknya, yang mana akan meninggalkan sarang selepas 7–28 hari. Tidak seperti kebanyakan burung-burung lain, kedua-dua jantina merpati dan punai akan menghasilkan "susu tembolok" untuk diberi kepada anaknya. Susu ini dihasilkan oleh satu kulit yang dipenuhi sel-sel cecair daripada lapisan tembolok. Merpati dan punai muda dipanggil "squab".