I have sold items to coutries such as Afghanistan
* Albania * Algeria * American Samoa (US) * Andorra * Angola *
Anguilla (GB) * Antigua and Barbuda * Argentina * Armenia * Aruba (NL) *
Australia * Austria * Azerbaijan * Bahamas * Bahrain * Bangladesh *
Barbados * Belarus * Belgium * Belize * Benin * Bermuda (GB) * Bhutan *
Bolivia * Bonaire (NL) * Bosnia and Herzegovina * Botswana * Bouvet
Island (NO) * Brazil * British Indian Ocean Territory (GB) * British
Virgin Islands (GB) * Brunei * Bulgaria * Burkina Faso * Burundi *
Cambodia * Cameroon * Canada * Cape Verde * Cayman Islands (GB) *
Central African Republic * Chad * Chile * China * Christmas Island (AU) *
Cocos Islands (AU) * Colombia * Comoros * Congo * Democratic Republic
of the Congo * Cook Islands (NZ) * Coral Sea Islands Territory (AU) *
Costa Rica * Croatia * Cuba * Curaçao (NL) * Cyprus * Czech Republic *
Denmark * Djibouti * Dominica * Dominican Republic * East Timor *
Ecuador * Egypt * El Salvador * Equatorial Guinea * Eritrea * Estonia *
Ethiopia * Falkland Islands (GB) * Faroe Islands (DK) * Fiji Islands *
Finland * France * French Guiana (FR) * French Polynesia (FR) * French
Southern Lands (FR) * Gabon * Gambia * Georgia * Germany * Ghana *
Gibraltar (GB) * Greece * Greenland (DK) * Grenada * Guadeloupe (FR) *
Guam (US) * Guatemala * Guernsey (GB) * Guinea * Guinea-Bissau * Guyana *
Haiti * Heard and McDonald Islands (AU) * Honduras * Hong Kong (CN) *
Hungary * Iceland * India * Indonesia * Iran * Iraq * Ireland * Isle of
Man (GB) * Israel * Italy * Ivory Coast * Jamaica * Jan Mayen (NO) *
Japan * Jersey (GB) * Jordan * Kazakhstan * Kenya * Kiribati * Kosovo *
Kuwait * Kyrgyzstan * Laos * Latvia * Lebanon * Lesotho * Liberia *
Libya * Liechtenstein * Lithuania * Luxembourg * Macau (CN) * Macedonia *
Madagascar * Malawi * Malaysia * Maldives * Mali * Malta * Marshall
Islands * Martinique (FR) * Mauritania * Mauritius * Mayotte (FR) *
Mexico * Micronesia * Moldova * Monaco * Mongolia * Montenegro *
Montserrat (GB) * Morocco * Mozambique * Myanmar * Namibia * Nauru *
Navassa (US) * Nepal * Netherlands * New Caledonia (FR) * New Zealand *
Nicaragua * Niger * Nigeria * Niue (NZ) * Norfolk Island (AU) * North
Korea * Northern Cyprus * Northern Mariana Islands (US) * Norway * Oman *
Pakistan * Palau * Palestinian Authority * Panama * Papua New Guinea *
Paraguay * Peru * Philippines * Pitcairn Island (GB) * Poland *
Portugal * Puerto Rico (US) * Qatar * Reunion (FR) * Romania * Russia *
Rwanda * Saba (NL) * Saint Barthelemy (FR) * Saint Helena (GB) *
Saint Kitts and Nevis * Saint Lucia * Saint Martin (FR) * Saint Pierre
and Miquelon (FR) * Saint Vincent and the Grenadines * Samoa * San
Marino * Sao Tome and Principe * Saudi Arabia * Senegal * Serbia *
Seychelles * Sierra Leone * Singapore * Sint Eustatius (NL) * Sint
Maarten (NL) * Slovakia * Slovenia * Solomon Islands * Somalia * South
Africa * South Georgia (GB) * South Korea * South Sudan * Spain * Sri
Lanka * Sudan * Suriname * Svalbard (NO) * Swaziland * Sweden *
Switzerland * Syria * Taiwan * Tajikistan * Tanzania * Thailand * Togo *
Tokelau (NZ) * Tonga * Trinidad and Tobago * Tunisia * Turkey *
Turkmenistan * Turks and Caicos Islands (GB) * Tuvalu * U.S. Minor
Pacific Islands (US) * U.S. Virgin Islands (US) * Uganda * Ukraine *
United Arab Emirates * United Kingdom * United States * Uruguay *
Uzbekistan * Vanuatu * Vatican City * Venezuela * Vietnam * Wallis and
Futuna (FR) * Yemen * Zambia * Zimbabwe and major cities such as Tokyo,
Yokohama, New York City, Sao Paulo, Seoul, Mexico City, Osaka, Kobe,
Kyoto, Manila, Mumbai, Delhi, Jakarta, Lagos, Kolkata, Cairo, Los
Angeles, Buenos Aires, Rio de Janeiro, Moscow, Shanghai, Karachi,
Paris, Istanbul, Nagoya, Beijing, Chicago, London, Shenzhen, Essen,
Düsseldorf, Tehran, Bogota, Lima, Bangkok, Johannesburg, East Rand,
Chennai, Taipei, Baghdad, Santiago, Bangalore, Hyderabad, St
Petersburg, Philadelphia, Lahore, Kinshasa, Miami, Ho Chi Minh City,
Madrid, Tianjin, Kuala Lumpur, Toronto, Milan, Shenyang, Dallas, Fort
Worth, Boston, Belo Horizonte, Khartoum, Riyadh, Singapore, Washington,
Detroit, Barcelona,, Houston, Athens, Berlin, Sydney, Atlanta,
Guadalajara, San Francisco, Oakland, Montreal, Monterey, Melbourne,
Ankara, Recife, Phoenix/Mesa, Durban, Porto Alegre, Dalian, Jeddah,
Seattle, Cape Town, San Diego, Fortaleza, Curitiba, Rome, Naples,
Minneapolis, St. Paul, Tel Aviv, Birmingham, Frankfurt, Lisbon,
Manchester, San Juan, Katowice, Tashkent, Fukuoka, Baku, Sumqayit, St.
Louis, Baltimore, Sapporo, Tampa, St. Petersburg, Taichung, Warsaw,
Denver, Cologne, Bonn, Hamburg, Dubai, Pretoria, Vancouver, Beirut,
Budapest, Cleveland, Pittsburgh, Campinas, Harare, Brasilia, Kuwait,
Munich, Portland, Brussels, Vienna, San Jose, Damman , Copenhagen,
Brisbane, Riverside, San Bernardino, Cincinnati and Accra
Rabbit
Temporal range: Late Eocene–Holocene, 53–0 Ma
PreꞒ
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S
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A small brown rabbit sat on the dirt in a forest. Its ears are small and alert and the tip of its nose, part of its chest and one of its feet are white.
European rabbit (Oryctolagus cuniculus)
Scientific classificationEdit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Lagomorpha
Family: Leporidae
Included genera
Pentalagus
Bunolagus
Nesolagus
Romerolagus
Brachylagus
Sylvilagus
Oryctolagus
Poelagus
Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). Oryctolagus cuniculus includes the European rabbit species and its descendants, the world's 305 breeds[1] of domestic rabbit. Sylvilagus includes 13 wild rabbit species, among them the seven types of cottontail. The European rabbit, which has been introduced on every continent except Antarctica, is familiar throughout the world as a wild prey animal and as a domesticated form of livestock and pet. With its widespread effect on ecologies and cultures, the rabbit is, in many areas of the world, a part of daily life—as food, clothing, a companion, and a source of artistic inspiration.
Although once considered rodents, lagomorphs like rabbits have been discovered to have diverged separately and earlier than their rodent cousins and have a number of traits rodents lack, like two extra incisors.
Terminology and etymology
A male rabbit is called a buck; a female is called a doe. An older term for an adult rabbit used until the 18th century is coney (derived ultimately from the Latin cuniculus), while rabbit once referred only to the young animals.[2] Another term for a young rabbit is bunny, though this term is often applied informally (particularly by children) to rabbits generally, especially domestic ones. More recently, the term kit or kitten has been used to refer to a young rabbit.
A group of rabbits is known as a colony or nest (or, occasionally, a warren, though this more commonly refers to where the rabbits live).[3] A group of baby rabbits produced from a single mating is referred to as a litter[4] and a group of domestic rabbits living together is sometimes called a herd.[5]
The word rabbit itself derives from the Middle English rabet, a borrowing from the Walloon robète, which was a diminutive of the French or Middle Dutch robbe.[6]
Taxonomy
See also: List of leporids
Rabbits and hares were formerly classified in the order Rodentia (rodent) until 1912, when they were moved into a new order, Lagomorpha (which also includes pikas). Below are some of the genera and species of the rabbit.
A small, round, dust-coloured rabbit with upright, close-set ears sat on the ground amidst dead branches.
Brachylagus idahoensis
Pygmy rabbit
A model of a relatively large rabbit, with a slightly longer face and shorter, rounder ears, and fur patterned black and gold.
Nesolagus netscheri
Sumatran Striped Rabbit
(Model)
A small, light-brown rabbit with upright ears sat on some grass.
Oryctolagus cuniculus
European rabbit
(Feral Tasmanian specimen)
A taxidermy of a large rabbit with dark brown fur, small, thin ears and an elongated, rodent-like face.
Pentalagus furnessi
Amami rabbit
(Taxidermy specimen)
A taxidermy of a very small light-brown rabbit mid-gallop. Its features are equally small, appearing similar to a vole.
Romerolagus diazi
Volcano rabbit
(Taxidermy specimen)
A juvenile rabbit sat on a white person's hand; its fur is a light brown ticked heavily with dark brown. It is not quite large enough to fill the person's hand completely.
Sylvilagus aquaticus
Swamp rabbit
(Juvenile)
A medium-sized rabbit with light brown fur ticked with grey, its ears large and upright. It stands on all fours on some sandy ground.
Sylvilagus audubonii
Desert cottontail
A light-brown rabbit sitting in a field, its body unusually large and squat, its limbs and head small, and its ears especially small
Sylvilagus bachmani
Brush rabbit
A taxidermy of a small rabbit, its fur a warm brown ticked with a darker brown, its ears small and set back, its face closer to a vole's than a rabbit's
Sylvilagus brasiliensis
Tapeti
(Taxidermy specimen)
A rabbit sitting upright in a field, turning to face the camera, its fur a light brown ticked with grey and dark brown, its ears upright
Sylvilagus floridanus
Eastern cottontail
Order Lagomorpha
Family Leporidae (in part)
Genus Brachylagus
Pygmy rabbit, Brachylagus idahoensis
Genus Bunolagus
Bushman rabbit, Bunolagus monticularis
Genus Lepus[a]
Genus Nesolagus
Sumatran striped rabbit, Nesolagus netscheri
Annamite striped rabbit, Nesolagus timminsi
Genus Oryctolagus
European rabbit, Oryctolagus cuniculus
Genus Pentalagus
Amami rabbit/Ryūkyū rabbit, Pentalagus furnessi
Genus Poelagus
Central African Rabbit, Poelagus marjorita
Genus Romerolagus
Volcano rabbit, Romerolagus diazi
Genus Sylvilagus
Swamp rabbit, Sylvilagus aquaticus
Desert cottontail, Sylvilagus audubonii
Brush rabbit, Sylvilagus bachmani
Forest rabbit, Sylvilagus brasiliensis
Mexican cottontail, Sylvilagus cunicularis
Dice's cottontail, Sylvilagus dicei
Eastern cottontail, Sylvilagus floridanus
Tres Marias rabbit, Sylvilagus graysoni
Omilteme cottontail, Sylvilagus insonus
San Jose brush rabbit, Sylvilagus mansuetus
Mountain cottontail, Sylvilagus nuttallii
Marsh rabbit, Sylvilagus palustris
New England cottontail, Sylvilagus transitionalis
Hare
Johann Daniel Meyer (1748)
Rabbit
Johann Daniel Meyer (1748)
Differences from hares
Main article: Hare
The term "rabbit" is typically used for all Leporidae species excluding the genus Lepus. Members of that genus are instead known as hares or jackrabbits.
Lepus species are typically precocial, born relatively mature and mobile with hair and good vision, while rabbit species are altricial, born hairless and blind, and requiring closer care. Hares live a relatively solitary life in a simple nest above the ground, while most rabbits live in social groups in burrows or warrens. Hares are generally larger than rabbits, with ears that are more elongated, and with hind legs that are larger and longer. Descendants of the European rabbit are commonly bred as livestock and kept as pets, whereas no hares have been domesticated - the breed called the Belgian hare is actually a domestic rabbit which has been selectively bred to resemble a hare.
Domestication
Main article: Domestic rabbit
Rabbits have long been domesticated. Beginning in the Middle Ages, the European rabbit has been widely kept as livestock, starting in ancient Rome. Selective breeding has generated a wide variety of rabbit breeds, of which many (since the early 19th century) are also kept as pets. Some strains of rabbit have been bred specifically as research subjects.
As livestock, rabbits are bred for their meat and fur. The earliest breeds were important sources of meat, and so became larger than wild rabbits, but domestic rabbits in modern times range in size from dwarf to giant. Rabbit fur, prized for its softness, can be found in a broad range of coat colors and patterns, as well as lengths. The Angora rabbit breed, for example, was developed for its long, silky fur, which is often hand-spun into yarn. Other domestic rabbit breeds have been developed primarily for the commercial fur trade, including the Rex, which has a short plush coat.
Biology
Evolution
Development of the rabbit heart
(wax models)
Because the rabbit's epiglottis is engaged over the soft palate except when swallowing, the rabbit is an obligate nasal breather. Rabbits have two sets of incisor teeth, one behind the other. This way they can be distinguished from rodents, with which they are often confused.[7] Carl Linnaeus originally grouped rabbits and rodents under the class Glires; later, they were separated as the scientific consensus is that many of their similarities were a result of convergent evolution. Recent DNA analysis and the discovery of a common ancestor has supported the view that they share a common lineage, so rabbits and rodents are now often grouped together in the superorder Glires.[8]
Morphology
Skeleton of the rabbit
Since speed and agility are a rabbit's main defenses against predators (including the swift fox), rabbits have large hind leg bones and well developed musculature. Though plantigrade at rest, rabbits are on their toes while running, assuming a more digitigrade posture. Rabbits use their strong claws for digging and (along with their teeth) for defense.[9] Each front foot has four toes plus a dewclaw. Each hind foot has four toes (but no dewclaw).[10]
Melanistic coloring
Oryctologus cuniculus
European rabbit (wild)
Most wild rabbits (especially compared to hares) have relatively full, egg-shaped bodies. The soft coat of the wild rabbit is agouti in coloration (or, rarely, melanistic), which aids in camouflage. The tail of the rabbit (with the exception of the cottontail species) is dark on top and white below. Cottontails have white on the top of their tails.[11]
As a result of the position of the eyes in its skull, the rabbit has a field of vision that encompasses nearly 360 degrees, with just a small blind spot at the bridge of the nose.[12]
Hind limb elements
This image comes from a specimen in the Pacific Lutheran University natural history collection. It displays all of the skeletal articulations of rabbit's hind limbs.
The anatomy of rabbits' hind limbs are structurally similar to that of other land mammals and contribute to their specialized form of locomotion. The bones of the hind limbs consist of long bones (the femur, tibia, fibula, and phalanges) as well as short bones (the tarsals). These bones are created through endochondral ossification during development. Like most land mammals, the round head of the femur articulates with the acetabulum of the os coxae. The femur articulates with the tibia, but not the fibula, which is fused to the tibia. The tibia and fibula articulate with the tarsals of the pes, commonly called the foot. The hind limbs of the rabbit are longer than the front limbs. This allows them to produce their hopping form of locomotion. Longer hind limbs are more capable of producing faster speeds. Hares, which have longer legs than cottontail rabbits, are able to move considerably faster.[13] Rabbits stay just on their toes when moving; this is called Digitigrade locomotion. The hind feet have four long toes that allow for this and are webbed to prevent them from spreading when hopping.[14] Rabbits do not have paw pads on their feet like most other animals that use digitigrade locomotion. Instead, they have coarse compressed hair that offers protection.[15]
Musculature
The rabbits hind limb (lateral view) includes muscles involved in the quadriceps and hamstrings.
Rabbits have muscled hind legs that allow for maximum force, maneuverability, and acceleration that is divided into three main parts; foot, thigh, and leg. The hind limbs of a rabbit are an exaggerated feature. They are much longer than the forelimbs, providing more force. Rabbits run on their toes to gain the optimal stride during locomotion. The force put out by the hind limbs is contributed to both the structural anatomy of the fusion tibia and fibula, and muscular features.[16] Bone formation and removal, from a cellular standpoint, is directly correlated to hind limb muscles. Action pressure from muscles creates force that is then distributed through the skeletal structures. Rabbits that generate less force, putting less stress on bones are more prone to osteoporosis due to bone rarefaction.[17] In rabbits, the more fibers in a muscle, the more resistant to fatigue. For example, hares have a greater resistance to fatigue than cottontails. The muscles of rabbit's hind limbs can be classified into four main categories: hamstrings, quadriceps, dorsiflexors, or plantar flexors. The quadriceps muscles are in charge of force production when jumping. Complementing these muscles are the hamstrings, which aid in short bursts of action. These muscles play off of one another in the same way as the plantar flexors and dorsiflexors, contributing to the generation and actions associated with force.[18]
Ears
A Holland Lop resting with one ear up and one ear down. Some rabbits can adjust their ears to hear distant sounds.
Within the order lagomorphs, the ears are utilized to detect and avoid predators. In the family Leporidae, the ears are typically longer than they are wide. For example, in black tailed jack rabbits, their long ears cover a greater surface area relative to their body size that allow them to detect predators from far away. Contrasted to cotton tailed rabbits, their ears are smaller and shorter, requiring predators to be closer to detect them before they can flee. Evolution has favored rabbits having shorter ears so the larger surface area does not cause them to lose heat in more temperate regions. The opposite can be seen in rabbits that live in hotter climates, mainly because they possess longer ears that have a larger surface area that help with dispersion of heat as well as the theory that sound does not travel well in more arid air, opposed to cooler air. Therefore, longer ears are meant to aid the organism in detecting predators sooner rather than later in warmer temperatures.[19][page needed] The rabbit is characterized by its shorter ears while hares are characterized by their longer ears.[20][page needed] Rabbits' ears are an important structure to aid thermoregulation and detect predators due to how the outer, middle, and inner ear muscles coordinate with one another. The ear muscles also aid in maintaining balance and movement when fleeing predators.[21]
Anatomy of mammalian ear
Outer ear
The auricle, also known as the pinna, is a rabbit's outer ear.[22] The rabbit's pinnae represent a fair part of the body surface area. It is theorized that the ears aid in dispersion of heat at temperatures above 30 °C with rabbits in warmer climates having longer pinnae due to this. Another theory is that the ears function as shock absorbers that could aid and stabilize rabbit's vision when fleeing predators, but this has typically only been seen in hares.[23][page needed] The rest of the outer ear has bent canals that lead to the eardrum or tympanic membrane.[24]
Middle ear
The middle ear is filled with three bones called ossicles and is separated by the outer eardrum in the back of the rabbit's skull. The three ossicles are called hammer, anvil, and stirrup and act to decrease sound before it hits the inner ear. In general, the ossicles act as a barrier to the inner ear for sound energy.[24]
Inner ear
Inner ear fluid called endolymph receives the sound energy. After receiving the energy, later within the inner ear there are two parts: the cochlea that utilizes sound waves from the ossicles and the vestibular apparatus that manages the rabbit's position in regards to movement. Within the cochlea there is a basilar membrane that contains sensory hair structures utilized to send nerve signals to the brain so it can recognize different sound frequencies. Within the vestibular apparatus the rabbit possesses three semicircular canals to help detect angular motion.[24]
Thermoregulation
Thermoregulation is the process that an organism utilizes to maintain an optimal body temperature independent of external conditions.[25] This process is carried out by the pinnae, which takes up most of the rabbit's body surface and contain a vascular network and arteriovenous shunts.[26] In a rabbit, the optimal body temperature is around 38.5–40℃.[27] If their body temperature exceeds or does not meet this optimal temperature, the rabbit must return to homeostasis. Homeostasis of body temperature is maintained by the use of their large, highly vascularized ears that are able to change the amount of blood flow that passes through the ears.
Rabbits use their large vascularized ears, which aid in thermoregulation, to keep their body temperature at an optimal level.
Constriction and dilation of blood vessels in the ears are used to control the core body temperature of a rabbit. If the core temperature exceeds its optimal temperature greatly, blood flow is constricted to limit the amount of blood going through the vessels. With this constriction, there is only a limited amount of blood that is passing through the ears where ambient heat would be able to heat the blood that is flowing through the ears and therefore, increasing the body temperature. Constriction is also used when the ambient temperature is much lower than that of the rabbit's core body temperature. When the ears are constricted it again limits blood flow through the ears to conserve the optimal body temperature of the rabbit. If the ambient temperature is either 15 degrees above or below the optimal body temperature, the blood vessels will dilate. With the blood vessels being enlarged, the blood is able to pass through the large surface area, causing it to either heat or cool down.
During hot summers, the rabbit has the capability to stretch its pinnae, which allows for greater surface area and increase heat dissipation. In cold winters, the rabbit does the opposite and folds its ears in order to decrease its surface area to the ambient air, which would decrease their body temperature.
Ventral view of dissected rabbit lungs with key structures labeled.
The jackrabbit has the largest ears within the Oryctolagus cuniculus group. Their ears contribute to 17% of their total body surface area. Their large pinna were evolved to maintain homeostasis while in the extreme temperatures of the desert.
Respiratory system
The rabbit's nasal cavity lies dorsal to the oral cavity, and the two compartments are separated by the hard and soft palate.[28] The nasal cavity itself is separated into a left and right side by a cartilage barrier, and it is covered in fine hairs that trap dust before it can enter the respiratory tract.[28][29][page needed] As the rabbit breathes, air flows in through the nostrils along the alar folds. From there, the air moves into the nasal cavity, also known as the nasopharynx, down through the trachea, through the larynx, and into the lungs.[29][page needed][30] The larynx functions as the rabbit's voice box, which enables it to produce a wide variety of sounds.[29][page needed] The trachea is a long tube embedded with cartilaginous rings that prevent the tube from collapsing as air moves in and out of the lungs. The trachea then splits into a left and right bronchus, which meet the lungs at a structure called the hilum. From there, the bronchi split into progressively more narrow and numerous branches. The bronchi branch into bronchioles, into respiratory bronchioles, and ultimately terminate at the alveolar ducts. The branching that is typically found in rabbit lungs is a clear example of monopodial branching, in which smaller branches divide out laterally from a larger central branch.[31]
The structure of the rabbit's nasal and oral cavities, necessitates breathing through the nose. This is due to the fact that the epiglottis is fixed to the backmost portion of the soft palate.[30] Within the oral cavity, a layer of tissue sits over the opening of the glottis, which blocks airflow from the oral cavity to the trachea.[28] The epiglottis functions to prevent the rabbit from aspirating on its food. Further, the presence of a soft and hard palate allow the rabbit to breathe through its nose while it feeds.[29][page needed]
Monopodial branching as seen in dissected rabbit lungs.
Rabbits lungs are divided into four lobes: the cranial, middle, caudal, and accessory lobes. The right lung is made up of all four lobes, while the left lung only has two: the cranial and caudal lobes.[31] In order to provide space for the heart, the left cranial lobe of the lungs is significantly smaller than that of the right.[28] The diaphragm is a muscular structure that lies caudal to the lungs and contracts to facilitate respiration.[28][30]
Digestion
Rabbits are herbivores that feed by grazing on grass and other leafy plants. In consequence, their diet contains large amounts of cellulose, which is hard to digest. Rabbits solve this problem via a form of hindgut fermentation. They pass two distinct types of feces: hard droppings and soft black viscous pellets, the latter of which are known as caecotrophs or "night droppings" [32] and are immediately eaten (a behaviour known as coprophagy). Rabbits reingest their own droppings (rather than chewing the cud as do cows and numerous other herbivores) to digest their food further and extract sufficient nutrients.[33]
Rabbits graze heavily and rapidly for roughly the first half-hour of a grazing period (usually in the late afternoon), followed by about half an hour of more selective feeding.[citation needed] In this time, the rabbit will also excrete many hard fecal pellets, being waste pellets that will not be reingested.[citation needed] If the environment is relatively non-threatening, the rabbit will remain outdoors for many hours, grazing at intervals.[citation needed] While out of the burrow, the rabbit will occasionally reingest its soft, partially digested pellets; this is rarely observed, since the pellets are reingested as they are produced.[citation needed]
0:54
Video of a wild European rabbit with ears twitching and a jump
Hard pellets are made up of hay-like fragments of plant cuticle and stalk, being the final waste product after redigestion of soft pellets. These are only released outside the burrow and are not reingested. Soft pellets are usually produced several hours after grazing, after the hard pellets have all been excreted.[citation needed] They are made up of micro-organisms and undigested plant cell walls.[citation needed]
Rabbits are hindgut digesters. This means that most of their digestion takes place in their large intestine and cecum. In rabbits, the cecum is about 10 times bigger than the stomach and it along with the large intestine makes up roughly 40% of the rabbit's digestive tract.[34] The unique musculature of the cecum allows the intestinal tract of the rabbit to separate fibrous material from more digestible material; the fibrous material is passed as feces, while the more nutritious material is encased in a mucous lining as a cecotrope. Cecotropes, sometimes called "night feces", are high in minerals, vitamins and proteins that are necessary to the rabbit's health. Rabbits eat these to meet their nutritional requirements; the mucous coating allows the nutrients to pass through the acidic stomach for digestion in the intestines. This process allows rabbits to extract the necessary nutrients from their food.[35]
The chewed plant material collects in the large cecum, a secondary chamber between the large and small intestine containing large quantities of symbiotic bacteria that help with the digestion of cellulose and also produce certain B vitamins. The pellets are about 56% bacteria by dry weight, largely accounting for the pellets being 24.4% protein on average. The soft feces form here and contain up to five times the vitamins of hard feces. After being excreted, they are eaten whole by the rabbit and redigested in a special part of the stomach. The pellets remain intact for up to six hours in the stomach; the bacteria within continue to digest the plant carbohydrates. This double-digestion process enables rabbits to use nutrients that they may have missed during the first passage through the gut, as well as the nutrients formed by the microbial activity and thus ensures that maximum nutrition is derived from the food they eat.[11] This process serves the same purpose in the rabbit as rumination does in cattle and sheep.[36]
Dissected image of the male rabbit reproductive system with key structures labeled.
Because rabbits cannot vomit,[37] if buildup occurs within the intestines (due often to a diet with insufficient fibre),[38] intestinal blockage can occur.[39]
Reproduction
Diagram of the male rabbit reproductive system with main components labeled.
The adult male reproductive system forms the same as most mammals with the seminiferous tubular compartment containing the Sertoli cells and an adluminal compartment that contains the Leydig cells.[40] The Leydig cells produce testosterone, which maintains libido[40] and creates secondary sex characteristics such as the genital tubercle and penis. The Sertoli cells triggers the production of Anti-Müllerian duct hormone, which absorbs the Müllerian duct. In an adult male rabbit, the sheath of the penis is cylinder-like and can be extruded as early as two months of age.[41] The scrotal sacs lay lateral to the penis and contain epididymal fat pads which protect the testes. Between 10 and 14 weeks, the testes descend and are able to retract into the pelvic cavity in order to thermoregulate.[41] Furthermore, the secondary sex characteristics, such as the testes, are complex and secrete many compounds. These compounds includes fructose, citric acid, minerals, and a uniquely high amount of catalase.[40]
Diagram of the female rabbit reproductive system with main components labeled.
The adult female reproductive tract is bipartite, which prevents an embryo from translocating between uteri.[42] The two uterine horns communicate to two cervixes and forms one vaginal canal. Along with being bipartite, the female rabbit does not go through an estrus cycle, which causes mating induced ovulation.[41]
The average female rabbit becomes sexually mature at three to eight months of age and can conceive at any time of the year for the duration of her life. Egg and sperm production can begin to decline after three years.[40] During mating, the male rabbit will mount the female rabbit from behind and insert his penis into the female and make rapid pelvic hip thrusts. The encounter lasts only 20–40 seconds and after, the male will throw himself backwards off the female.[43]
The rabbit gestation period is short and ranges from 28 to 36 days with an average period of 31 days. A longer gestation period will generally yield a smaller litter while shorter gestation periods will give birth to a larger litter. The size of a single litter can range from four to 12 kits allowing a female to deliver up to 60 new kits a year. After birth, the female can become pregnant again as early as the next day.[41]
The mortality rates of embryos are high in rabbits and can be due to infection, trauma, poor nutrition and environmental stress so a high fertility rate is necessary to counter this.[41]
Sleep
Further information: Sleep (non-human)
Rabbits may appear to be crepuscular, but their natural inclination is toward nocturnal activity.[44] In 2011, the average sleep time of a rabbit in captivity was calculated at 8.4 hours per day.[45] As with other prey animals, rabbits often sleep with their eyes open, so that sudden movements will awaken the rabbit to respond to potential danger.[46]
Diseases and immunity
See also: Category:Rabbit diseases
In addition to being at risk of disease from common pathogens such as Bordetella bronchiseptica and Escherichia coli, rabbits can contract the virulent, species-specific viruses RHD ("rabbit hemorrhagic disease", a form of calicivirus)[47] or myxomatosis. Among the parasites that infect rabbits are tapeworms (such as Taenia serialis), external parasites (including fleas and mites), coccidia species, and Toxoplasma gondii.[48][49] Domesticated rabbits with a diet lacking in high fiber sources, such as hay and grass, are susceptible to potentially lethal gastrointestinal stasis.[50] Rabbits and hares are almost never found to be infected with rabies and have not been known to transmit rabies to humans.[51]
Encephalitozoon cuniculi, an obligate intracellular parasite is also capable of infecting many mammals including rabbits.
Rabbit immunity has significantly diverged from other tetrapods in the manner it employs immunoglobulin light chains.[52][53] In one case McCartney-Francis et al., 1984 discover a unique additional disulfide bond between Cys 80 in Vκ and Cys 171 in Cκ.[52][53] They suggest that this may serve to stabilise rabbit antibodies.[52][53] Meanwhile IGKC1 shows high amino acid divergence between domesticated types and ferals derived from them.[53] This can be as high as 40%.[53]
Rabbit hemorrhagic disease is caused by strains of rabbit hemorrhagic disease virus (RHDV) including type 2 (RHDV2).[54] RHDV2 was detected for the first time in Washington state, USA in May 2022 and then in August once in Washington and twice in Oregon.[55]
Ecology
Rabbit kits one hour after birth
Rabbits are prey animals and are therefore constantly aware of their surroundings. For instance, in Mediterranean Europe, rabbits are the main prey of red foxes, badgers, and Iberian lynxes.[56] If confronted by a potential threat, a rabbit may freeze and observe then warn others in the warren with powerful thumps on the ground. Rabbits have a remarkably wide field of vision, and a good deal of it is devoted to overhead scanning.[57] The doe (mother) is aware that she gives off scent which can attract predators, so she will stay away from the nest to avoid putting the kits (babies) in danger, returning the nest only a few times a day to feed the kits.[58]
Rabbits survive predation by burrowing, hopping away in a zig-zag motion, and, if captured, delivering powerful kicks with their hind legs. Their strong teeth allow them to eat and to bite in order to escape a struggle.[59] The longest-lived rabbit on record, a domesticated European rabbit living in Tasmania, died at age 18.[60] The lifespan of wild rabbits is much shorter; the average longevity of an eastern cottontail, for instance, is less than one year.[61]
Habitat and range
Rabbit habitats include meadows, woods, forests, grasslands, deserts and wetlands.[62] Rabbits live in groups, and the best known species, the European rabbit, lives in burrows, or rabbit holes. A group of burrows is called a warren.[62]
More than half the world's rabbit population resides in North America.[62] They are also native to southwestern Europe, Southeast Asia, Sumatra, some islands of Japan, and in parts of Africa and South America. They are not naturally found in most of Eurasia, where a number of species of hares are present. Rabbits first entered South America relatively recently, as part of the Great American Interchange. Much of the continent has just one species of rabbit, the tapeti, while most of South America's southern cone is without rabbits.
The European rabbit has been introduced to many places around the world.[11]
Domestic rabbit photographed at Alligator Bay, Beauvoir, France.
Rabbits have been launched into space orbit.[63]
Environmental problems
See also: Rabbits in Australia
Impact of rabbit-proof fence, Cobar, New South Wales, 1905
Rabbits have been a source of environmental problems when introduced into the wild by humans. As a result of their appetites, and the rate at which they breed, feral rabbit depredation can be problematic for agriculture. Gassing (fumigation of warrens),[64] barriers (fences), shooting, snaring, and ferreting have been used to control rabbit populations, but the most effective measures are diseases such as myxomatosis (myxo or mixi, colloquially) and calicivirus. In Europe, where rabbits are farmed on a large scale, they are protected against myxomatosis and calicivirus with a genetically modified virus. The virus was developed in Spain, and is beneficial to rabbit farmers. If it were to make its way into wild populations in areas such as Australia, it could create a population boom, as those diseases are the most serious threats to rabbit survival. Rabbits in Australia and New Zealand are considered to be such a pest that land owners are legally obliged to control them.[65][66]
As food and clothing
Main article: Cuniculture
See also: Category:Rabbit dishes
Saint Jerome in the Desert
[Note rabbit being chased by a domesticated hound]
Taddeo Crivelli (Italian, died about 1479)
Rabbit being prepared in the kitchen
Simulation of daily life, mid-15th century
Hospices de Beaune, France
In some areas, wild rabbits and hares are hunted for their meat, a lean source of high quality protein.[67] In the wild, such hunting is accomplished with the aid of trained falcons, ferrets, or dogs, as well as with snares or other traps, and rifles. A caught rabbit may be dispatched with a sharp blow to the back of its head, a practice from which the term rabbit punch is derived.
Wild leporids comprise a small portion of global rabbit-meat consumption. Domesticated descendants of the European rabbit (Oryctolagus cuniculus) that are bred and kept as livestock (a practice called cuniculture) account for the estimated 200 million tons of rabbit meat produced annually.[68] Approximately 1.2 billion rabbits are slaughtered each year for meat worldwide.[69] In 1994, the countries with the highest consumption per capita of rabbit meat were Malta with 8.89 kg (19 lb 10 oz), Italy with 5.71 kg (12 lb 9 oz), and Cyprus with 4.37 kg (9 lb 10 oz), falling to 0.03 kg (1 oz) in Japan. The figure for the United States was 0.14 kg (5 oz) per capita. The largest producers of rabbit meat in 1994 were China, Russia, Italy, France, and Spain.[70] Rabbit meat was once a common commodity in Sydney, Australia, but declined after the myxomatosis virus was intentionally introduced to control the exploding population of feral rabbits in the area.
In the United Kingdom, fresh rabbit is sold in butcher shops and markets, and some supermarkets sell frozen rabbit meat. At farmers markets there, including the famous Borough Market in London, rabbit carcasses are sometimes displayed hanging, unbutchered (in the traditional style), next to braces of pheasant or other small game. Rabbit meat is a feature of Moroccan cuisine, where it is cooked in a tajine with "raisins and grilled almonds added a few minutes before serving".[71] In China, rabbit meat is particularly popular in Sichuan cuisine, with its stewed rabbit, spicy diced rabbit, BBQ-style rabbit, and even spicy rabbit heads, which have been compared to spicy duck neck.[68] Rabbit meat is comparatively unpopular elsewhere in the Asia-Pacific.
An extremely rare infection associated with rabbits-as-food is tularemia (also known as rabbit fever), which may be contracted from an infected rabbit.[72] Hunters are at higher risk for tularemia because of the potential for inhaling the bacteria during the skinning process.
In addition to their meat, rabbits are used for their wool, fur, and pelts, as well as their nitrogen-rich manure and their high-protein milk.[73] Production industries have developed domesticated rabbit breeds (such as the well-known Angora rabbit) to efficiently fill these needs.
In art, literature, and culture
Main article: Rabbits and hares in art
Rabbits are often used as a symbol of fertility or rebirth, and have long been associated with spring and Easter as the Easter Bunny. The species' role as a prey animal with few defenses evokes vulnerability and innocence, and in folklore and modern children's stories, rabbits often appear as sympathetic characters, able to connect easily with youth of all kinds (for example, the Velveteen Rabbit, or Thumper in Bambi).
With its reputation as a prolific breeder, the rabbit juxtaposes sexuality with innocence, as in the Playboy Bunny. The rabbit (as a swift prey animal) is also known for its speed, agility, and endurance, symbolized (for example) by the marketing icons the Energizer Bunny and the Duracell Bunny.
Folklore
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Main article: List of fictional hares and rabbits
The rabbit often appears in folklore as the trickster archetype, as he uses his cunning to outwit his enemies.
"Rabbit fools Elephant by showing the reflection of the moon".
Illustration (from 1354) of the Panchatantra
"Three rabbits" motif
Coat of arms of Corbenay, France
In Aztec mythology, a pantheon of four hundred rabbit gods known as Centzon Totochtin, led by Ometochtli or Two Rabbit, represented fertility, parties, and drunkenness.
In Central Africa, the common hare (Kalulu), is "inevitably described" as a trickster figure.[74]
In Chinese folklore, rabbits accompany Chang'e on the Moon. In the Chinese New Year, the zodiacal rabbit is one of the twelve celestial animals in the Chinese zodiac. Note that the Vietnamese zodiac includes a zodiacal cat in place of the rabbit, possibly because rabbits did not inhabit Vietnam.[citation needed] The most common explanation is that the ancient Vietnamese word for "rabbit" (mao) sounds like the Chinese word for "cat" (卯, mao).[75]
In Japanese tradition, rabbits live on the Moon where they make mochi, the popular snack of mashed sticky rice. This comes from interpreting the pattern of dark patches on the moon as a rabbit standing on tiptoes on the left pounding on an usu, a Japanese mortar.
In Jewish folklore, rabbits (shfanim שפנים) are associated with cowardice, a usage still current in contemporary Israeli spoken Hebrew (similar to the English colloquial use of "chicken" to denote cowardice).
In Korean mythology, as in Japanese, rabbits live on the moon making rice cakes ("Tteok" in Korean).
In Anishinaabe traditional beliefs, held by the Ojibwe and some other Native American peoples, Nanabozho, or Great Rabbit, is an important deity related to the creation of the world.
A Vietnamese mythological story portrays the rabbit of innocence and youthfulness. The Gods of the myth are shown to be hunting and killing rabbits to show off their power.
Buddhism, Christianity, and Judaism have associations with an ancient circular motif called the three rabbits (or "three hares"). Its meaning ranges from "peace and tranquility", to purity or the Holy Trinity, to Kabbalistic levels of the soul or to the Jewish diaspora. The tripartite symbol also appears in heraldry and even tattoos.
The rabbit as trickster is a part of American popular culture, as Br'er Rabbit (from African-American folktales and, later, Disney animation) and Bugs Bunny (the cartoon character from Warner Bros.), for example.
Anthropomorphized rabbits have appeared in film and literature, in Alice's Adventures in Wonderland (the White Rabbit and the March Hare characters), in Watership Down (including the film and television adaptations), in Rabbit Hill (by Robert Lawson), and in the Peter Rabbit stories (by Beatrix Potter). In the 1920s, Oswald the Lucky Rabbit, was a popular cartoon character.
WWII USAF pilot D. R. Emerson
"flys with a rabbit's foot talisman,
a gift from a New York girl friend"
Beatrix Potter's Peter Rabbit.
A rabbit's foot may be carried as an amulet, believed to bring protection and good luck. This belief is found in many parts of the world, with the earliest use being recorded in Europe c. 600 BC.[76]
On the Isle of Portland in Dorset, UK, the rabbit is said to be unlucky and even speaking the creature's name can cause upset among older island residents. This is thought to date back to early times in the local quarrying industry where (to save space) extracted stones that were not fit for sale were set aside in what became tall, unstable walls. The local rabbits' tendency to burrow there would weaken the walls and their collapse resulted in injuries or even death. Thus, invoking the name of the culprit became an unlucky act to be avoided. In the local culture to this day, the rabbit (when he has to be referred to) may instead be called a “long ears” or “underground mutton”, so as not to risk bringing a downfall upon oneself.[77] While it was true 50 years ago[when?] that a pub on the island could be emptied by calling out the word "rabbit", this has become more fable than fact in modern times.[citation needed]
In other parts of Britain and in North America, invoking the rabbit's name may instead bring good luck. "Rabbit rabbit rabbit" is one variant of an apotropaic or talismanic superstition that involves saying or repeating the word "rabbit" (or "rabbits" or "white rabbits" or some combination thereof) out loud upon waking on the first day of each month, because doing so will ensure good fortune for the duration of that month.
The "rabbit test" is a term, first used in 1949, for the Friedman test, an early diagnostic tool for detecting a pregnancy in humans. It is a common misconception (or perhaps an urban legend) that the test-rabbit would die if the woman was pregnant. This led to the phrase "the rabbit died" becoming a euphemism for a positive pregnancy test.
See also
iconLagomorpha portal
Animal track
Cuniculture
Dwarf rabbit
Hare games
Jackalope
List of animal names
List of rabbit breeds
Lop rabbit
Rabbits in the arts
Rabbit show jumping
References
Notes
This genus is considered a hare, not a rabbit
Citations
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Further reading
Windling, Terri. The Symbolism of Rabbits and Hares[Usurped!]
External links
Wikimedia Commons has media related to Rabbits.
Wikiquote has quotations related to Rabbit.
Wikisource has the text of the 1911 Encyclopædia Britannica article "Rabbit".
American Rabbit Breeders Association organization, which promotes all phases of rabbit keeping
House Rabbit Society an activist organization that promotes keeping rabbits indoors
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Categories:
LeporidaeHerbivorous mammalsExtant Ypresian first appearancesMammal common namesCosmopolitan mammalsParaphyletic groups
Leporidae is the family of rabbits and hares, containing over 60 species of extant mammals in all. The Latin word Leporidae means "those that resemble lepus" (hare). Together with the pikas, the Leporidae constitute the mammalian order Lagomorpha. Leporidae differ from pikas in that they have short, furry tails and elongated ears and hind legs.
The common name "rabbit" usually applies to all genera in the family except Lepus, while members of Lepus (almost half the species) usually are called hares. Like most common names, however, the distinction does not match current taxonomy completely; jackrabbits are members of Lepus, and members of the genera Pronolagus and Caprolagus sometimes are called hares.
Various countries across all continents except Antarctica and Australia have indigenous species of Leporidae. Furthermore, rabbits, most significantly the European rabbit, Oryctolagus cuniculus, also have been introduced to most of Oceania and to many other islands, where they pose serious ecological and commercial threats.
Contents
1 Characteristics
2 Reproduction
3 Evolution
4 Classification
5 Predation
6 See also
7 References
Characteristics
Leporids are small to moderately sized mammals, adapted for rapid movement. They have long hind legs, with four toes on each foot, and shorter fore legs, with five toes each. The soles of their feet are hairy, to improve grip while running, and they have strong claws on all of their toes. Leporids also have distinctive, elongated and mobile ears, and they have an excellent sense of hearing. Their eyes are large, and their night vision is good, reflecting their primarily nocturnal or crepuscular mode of living.[2]
Leporids are all roughly the same shape and fall within a small range of sizes with short tails, ranging from the 21 cm (8 in) long Tres Marias cottontail to the 76 cm (30 in) long desert hare. Female leporids are almost always larger than males, which is unusual among terrestrial mammals, in which males are usually the larger sex.[3]
Both rabbits and hares are almost exclusively herbivorous (although some Lepus species are known to eat carrion),[4][5] feeding primarily on grasses and herbs, although they also eat leaves, fruit, and seeds of various kinds. They are coprophagous, as they pass food through their digestive systems twice, first expelling it as soft green feces, called cecotropes, which they then reingest, eventually producing hard, dark fecal pellets. Like rodents, they have powerful front incisor teeth, but they also have a smaller second pair of incisors to either side of the main teeth in the upper jaw, and the structure is different from that of rodent incisors. Also like rodents, leporids lack any canine teeth, but they do have more cheek teeth than rodents do. Their jaws also contain a large diastema. The dental formula of most, though not all, leporids is: 2.0.3.31.0.2.3
They have adapted to a remarkable range of habitats, from desert to tundra, forests, mountains, and swampland. Rabbits generally dig permanent burrows for shelter, the exact form of which varies between species. In contrast, hares rarely dig shelters of any kind, and their bodies are more suited to fast running than to burrowing.[2]
The gestation period in leporids varies from around 28 to 50 days, and is generally longer in the hares. This is in part because young hares, or leverets, are born fully developed, with fur and open eyes, while rabbit kits are naked and blind at birth, having the security of the burrow to protect them.[2] Leporids can have several litters a year, which can cause their population to expand dramatically in a short time when resources are plentiful.
Reproduction
Leporids are typically polygynandrous, and have highly developed social systems. Their social hierarchies determine which males mate when the females go into estrus, which happens throughout the year. Gestation periods are variable, but in general, higher latitudes correspond to shorter gestation periods.[6] Moreover, the gestation time and litter size correspond to predation rates as well. Species nesting below ground tend to have lower predation rates and have larger litters.[7]
Evolution
Serengetilagus praecapensis skull, Naturkundemuseum, Berlin
The oldest known leporid species date from the late Eocene, by which time the family was already present in both North America and Asia. Over the course of their evolution, this group has become increasingly adapted to lives of fast running and leaping. For example, Palaeolagus, an extinct rabbit from the Oligocene of North America, had shorter hind legs than modern forms (indicating it ran rather than hopped) though it was in most other respects quite rabbit-like.[8] Two as yet unnamed fossil finds—dated ~48 Ma (from China) and ~53 Ma (India)—while primitive, display the characteristic leporid ankle, thus pushing the divergence of Ochotonidae and Leporidae yet further into the past.[9]
The cladogram is from Matthee et al., 2004, based on nuclear and mitochondrial gene analysis.[10]
Leporidae
Nesolagus (striped rabbits)
Poelagus (Bunyoro rabbit)
Pronolagus (red rock hares)
Romerolagus (volcano rabbit)
Sylvilagus (cottontails) Wild animals of North America, intimate studies of big and little creatures of the mammal kingdom (Page 511) (Sylvilagus palustris).jpg
Brachylagus (pygmy rabbit)
Caprolagus (hispid hare)
Oryctolagus (European rabbit) Lepus cuniculus - 1700-1880 - Print - Iconographia Zoologica -(white background).jpg
Bunolagus (riverine rabbit)
Pentalagus (Amami rabbit)
Lepus (hares) Lepus timidus - 1700-1880 - Print - Iconographia Zoologica -(white background).jpg
Classification
Main article: List of leporids
Family Leporidae:[1] rabbits and hares
Genus Pentalagus
Amami rabbit, Pentalagus furnessi
Genus Bunolagus
Riverine rabbit, Bunolagus monticularis
Genus Nesolagus
Sumatran striped rabbit, Nesolagus netscheri
Annamite striped rabbit, Nesolagus timminsi
Genus Romerolagus
Volcano rabbit, Romerolagus diazi
Genus Brachylagus
Pygmy rabbit, Brachylagus idahoensis
Genus Sylvilagus
Subgenus Tapeti
Swamp rabbit, Sylvilagus aquaticus
Andean tapetí, Sylvilagus andinus
Bogota tapetí, Sylvilagus apollinaris
Ecuadorian tapetí, Sylvilagus daulensis
Common tapetí, Sylvilagus brasiliensis
Fulvous tapetí, Sylvilagus fulvescens
Dice's cottontail, Sylvilagus dicei
Central American tapetí, Sylvilagus gabbi
Northern tapetí, Sylvilagus incitatus
Omilteme cottontail, Sylvilagus insonus
Nicefor's tapetí, Sylvilagus nicefori
Marsh rabbit, Sylvilagus palustris
Suriname tapetí, Sylvilagus parentum
Colombian tapetí, Sylvilagus salentus
Santa Marta tapetí, Sylvilagus sanctaemartae
Western tapetí, Sylvilagus surdaster
Coastal tapetí, Sylvilagus tapetillus
Venezuelan lowland rabbit, Sylvilagus varynaensis
Subgenus Sylvilagus
Desert cottontail, Sylvilagus audubonii
Mexican cottontail, Sylvilagus cunicularis
Eastern cottontail, Sylvilagus floridanus
Tres Marias cottontail, Sylvilagus graysoni
Robust cottontail, Sylvilagus holzneri
Mountain cottontail, Sylvilagus nuttallii
Appalachian cottontail, Sylvilagus obscurus
New England cottontail, Sylvilagus transitionalis
Subgenus Microlagus
Brush rabbit, Sylvilagus bachmani
Genus Oryctolagus
European rabbit, Oryctolagus cuniculus
Genus Poelagus
Bunyoro rabbit, Poelagus marjorita
Genus Pronolagus
Natal red rock hare, Pronolagus crassicaudatus
Jameson's red rock hare, Pronolagus randensis
Smith's red rock hare, Pronolagus rupestris
Hewitt's red rock hare, Pronolagus saundersiae
Genus Caprolagus
Hispid hare, Caprolagus hispidus
Genus Lepus
Subgenus Macrotolagus
Antelope jackrabbit, Lepus alleni
Subgenus Poecilolagus
Snowshoe hare, Lepus americanus
Subgenus Lepus
Arctic hare, Lepus arcticus
Alaskan hare, Lepus othus
Mountain hare, Lepus timidus
Subgenus Proeulagus
Black jackrabbit, Lepus insularis
Desert hare, Lepus tibetanus
Tolai hare, Lepus tolai
Subgenus Eulagos
Broom hare, Lepus castroviejoi
Yunnan hare, Lepus comus
Korean hare, Lepus coreanus
European hare, Lepus europaeus
Manchurian hare, Lepus mandshuricus
Ethiopian highland hare, Lepus starcki
Subgenus Sabanalagus
Ethiopian hare, Lepus fagani
African savanna hare, Lepus victoriae
Subgenus Indolagus
Hainan hare, Lepus hainanus
Indian hare, Lepus nigricollis
Burmese hare, Lepus peguensis
Subgenus Sinolagus
Chinese hare, Lepus sinensis
Subgenus Tarimolagus
Yarkand hare, Lepus yarkandensis
Incertae sedis
Tamaulipas jackrabbit, Lepus altamirae
Japanese hare, Lepus brachyurus
Black-tailed jackrabbit, Lepus californicus
White-sided jackrabbit, Lepus callotis
Cape hare, Lepus capensis
Corsican hare, Lepus corsicanus
Tehuantepec jackrabbit, Lepus flavigularis
Granada hare, Lepus granatensis
Abyssinian hare, Lepus habessinicus
Woolly hare, Lepus oiostolus
Scrub hare, Lepus saxatilis
White-tailed jackrabbit, Lepus townsendii
Genus †Serengetilagus
†Serengetilagus praecapensis
Genus †Aztlanolagus
†Aztlanolagus agilis
Predation
Predators of rabbits and hares include raccoons, snakes, eagles, canids, cats, mustelids, owls and hawks. Animals that eat roadkill rabbits include vultures and buzzards.
See also
iconRabbits and hares portal
Mara (mammal)
Viscacha
References
Hoffman, R.S.; Smith, A.T. (2005). "Order Lagomorpha". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 194–211. ISBN 978-0-8018-8221-0. OCLC 62265494.
Chapman, J.; Schneider, E. (1984). MacDonald, D. (ed.). The Encyclopedia of Mammals. New York: Facts on File. pp. 714–719. ISBN 978-0-87196-871-5.
Ralls, Katherine (June 1976). "Mammals in Which Females are Larger Than Males". The Quarterly Review of Biology. 51 (2): 245–276. doi:10.1086/409310. PMID 785524. S2CID 25927323.
Best, Troy L.; Henry, Travis Hill (1994). "Lepus arcticus". Mammalian Species. American Society of Mammalogists (published 2 June 1994) (457): 1–9. doi:10.2307/3504088. JSTOR 3504088. OCLC 46381503.
"Snowshoe Hare". eNature: FieldGuides. eNature.com. 2007. Archived from the original on 16 January 2009. Retrieved 23 March 2008.
Chapman, Joseph A. (1 September 1984). "Latitude and Gestation Period in New World Rabbits (Leporidae: Sylvilagus and Romerolagus)". The American Naturalist. 124 (3): 442–445. doi:10.1086/284286. JSTOR 2461471. S2CID 83584955.
Virgós, Emilio; Cabezas-Díaz, Sara; Blanco-Aguiar, José Antonio (1 August 2006). "Evolution of life history traits in Leporidae: a test of nest predation and seasonality hypotheses". Biological Journal of the Linnean Society. 88 (4): 603–610. doi:10.1111/j.1095-8312.2006.00646.x. ISSN 1095-8312.
Savage, R.J.G.; Long, M.R. (1986). Mammal Evolution: an illustrated guide. New York: Facts on File. pp. 128–129. ISBN 978-0-8160-1194-0.
Handwerk, Brian (21 March 2008). "Easter Surprise: World's Oldest Rabbit Bones Found". National Geographic News. National Geographic Society.
Matthee, Conrad A.; et al. (2004). "A Molecular Supermatrix of the Rabbits and Hares (Leporidae) Allows for the Identification of Five Intercontinental Exchanges During the Miocene". Systematic Biology. 53 (3): 433–477. doi:10.1080/10635150490445715. PMID 15503672.
A sovereign case is used for carrying either full or half gold sovereign coins and often worn on a gentleman's Albert/watch chain and kept in a waistcoat pocket. The coins would be stacked tightly together by a spring action mechanism; a case usually holds about 4 or 5 sovereigns. Some can have a number of compartments holding halves as well as full sovereigns. They can also be seen combined with vesta and stamp cases. The cases were at the height of fashion during the late Victorian and Edwardian period and can be highly decorated or completely plain.
Sovereign (British coin)
Sovereign
United Kingdom
Value £1
Mass 7.98805 g
Diameter 22.0 mm
Thickness 1.52 mm
Edge Milled (some not intended for circulation have plain edge)
Composition .917 gold, .083 copper or other metals
Gold 0.2354 troy oz
Years of minting 1817–present
Mint marks Various. Found on reverse on exergue between design and date for Saint George and the Dragon sovereigns, and under the wreath for shield back sovereigns, or below bust on obverse on earlier Australian issues.
Obverse
1959 sovereign Elizabeth II obverse.jpg
Design Reigning British monarch (Elizabeth II portrait by Mary Gillick pictured)
Reverse
1959 Elizabeth II sovereign reverse.jpg
Design Saint George and the Dragon
Designer Benedetto Pistrucci
Design date 1817
The sovereign is a British gold coin with a nominal value of one pound sterling (£1) and contains 0.2354 troy oz of pure gold. Struck since 1817, it was originally a circulating coin that was accepted in Britain and elsewhere in the world; it is now a bullion coin and is sometimes mounted in jewellery. In addition, circulation strikes and proof examples are often collected for their numismatic value. In most recent years, it has borne the design of Saint George and the Dragon on the reverse; the initials (B P) of the designer, Benedetto Pistrucci, are visible to the right of the date.
The coin was named after the English gold sovereign, which was last minted about 1603, and originated as part of the Great Recoinage of 1816. Many in Parliament believed a one-pound coin should be issued rather than the 21-shilling guinea that was struck until that time. The Master of the Mint, William Wellesley Pole had Pistrucci design the new coin; his depiction was also used for other gold coins. Originally, the coin was unpopular because the public preferred the convenience of banknotes but paper currency of value £1 was soon limited by law. With that competition gone, the sovereign became a popular circulating coin, and was used in international trade and overseas, being trusted as a coin containing a known quantity of gold.
The British government promoted the use of the sovereign as an aid to international trade, and the Royal Mint took steps to see lightweight gold coins were withdrawn from circulation. From the 1850s until 1932, the sovereign was also struck at colonial mints, initially in Australia and later in Canada, South Africa and India—they have again been struck in India for the local market since 2013, in addition to the production in Britain by the Royal Mint. The sovereigns issued in Australia initially carried a unique local design but by 1887, all new sovereigns bore Pistrucci's George and Dragon design. Strikings there were so large that by 1900, about forty per cent of the sovereigns in Britain had been minted in Australia.
With the start of the First World War in 1914, the sovereign vanished from circulation in Britain; it was replaced by paper money and did not return after the war, though issues at colonial mints continued until 1932. The coin was still used in the Middle East and demand rose in the 1950s, to which the Royal Mint eventually responded by striking new sovereigns in 1957. Since then, it has been struck both as a bullion coin and beginning in 1979 for collectors. Although the sovereign is no longer in circulation, it is still legal tender in the United Kingdom.
Background and authorisation
Gold coin showing a woman seated on a throne
Sovereign of Queen Mary I, c. 1553
There had been an English coin known as the sovereign, first authorised by Henry VII in 1489. It had a diameter of 42 millimetres (1.7 in), and weighed 15.55 grams (0.500 oz t), twice the weight of the existing gold coin, the ryal. The new coin was struck in response to a large influx of gold into Europe from West Africa in the 1480s, and Henry at first called it the double ryal, but soon changed the name to sovereign.[1] Too great in value to have any practical use in circulation, the original sovereign likely served as a presentation piece to be given to dignitaries.[2]
The English sovereign, the country's first coin to be valued at one pound,[3] was struck by the monarchs of the 16th century, the size and fineness often being altered. James I, when he came to the English throne in 1603, issued a sovereign in the year of his accession,[4] but the following year, soon after he proclaimed himself King of Great Britain, France[a] and Ireland, he issued a proclamation for a new twenty-shilling piece. About ten per cent lighter than the final sovereigns, the new coin was called the unite, symbolising that James had merged the Scottish and English crowns.[5]
In the 1660s, following the Restoration of Charles II and the mechanisation of the Royal Mint that quickly followed, a new twenty-shilling gold coin was issued. It had no special name at first but the public soon nicknamed it the guinea and this became the accepted term.[6] Coins were at the time valued by their precious metal content, and the price of gold relative to silver rose soon after the guinea's issuance. Thus, it came to trade at 21 shillings or even sixpence more. Popular in commerce, the coin's value was set by the government at 21 shillings in silver in 1717, and was subject to revision downwards, though in practice this did not occur.[7] The term sovereign, referring to a coin, fell from use—it does not appear in Samuel Johnson's dictionary, compiled in the 1750s.[8]
Old piece of paper money
A £1 note issued in 1814 by the Gloucester Old Bank
The British economy was disrupted by the Napoleonic Wars, and gold was hoarded. Among the measures taken to allow trade to continue was the issue of one-pound banknotes. The public came to like them as more convenient than the odd-value guinea. After the war, Parliament, by the Coinage Act 1816, placed Britain officially on the gold standard, with the pound to be defined as a given quantity of gold. Almost every speaker supported having a coin valued at twenty shillings, rather than continuing to use the guinea.[9] Nevertheless, the Coinage Act did not specify which coins the Mint should strike.[10] A committee of the Privy Council recommended gold coins of ten shillings, twenty shillings, two pounds and five pounds be issued, and this was accepted by George, Prince Regent on 3 August 1816.[11] The twenty-shilling piece was named a sovereign, with the resurrection of the old name possibly promoted by antiquarians with numismatic interests.[8]
Creation
A sovereign with the bust of George III on the obverse and Saint George slaying the dragon on the reverse
1817 sovereign of George III
William Wellesley Pole, elder brother of the Duke of Wellington, was appointed Master of the Mint (at that time a junior government position) in 1812, with a mandate to reform the Royal Mint. Pole had favoured retaining the guinea, due to the number extant and the amount of labour required to replace them with sovereigns.[12] Formal instruction to the Mint came with an indenture dated February 1817, directing the Royal Mint to strike gold sovereigns.[13] As one troy pound (12 troy ounces) of 22-karat gold used to be minted into 441⁄2 guineas worth 44.5*£11⁄20 = £4629⁄40, each troy pound of 22K gold was henceforth minted into 46.725 sovereigns, with each coin weighing 7.98805 g (0.256822 ozt, 123.274 grains) and containing 7.32238 g (0.235420 ozt) fine gold.[b]
The Italian sculptor Benedetto Pistrucci came to London early in 1816. His talent opened the doors of the capital's elite,[15] among them Lady Spencer, who showed Pistrucci a model in wax of Saint George and the Dragon by Nathaniel Marchant and commissioned him to reproduce it in the Greek style as part of her husband's regalia as a Knight of the Garter. Pistrucci had already been thinking of such a work, and he produced the cameo.[16] The model for the saint was an Italian waiter at Brunet's Hotel in Leicester Square, where he had stayed after coming to London.[17]
In 1816, Pole hired Pistrucci to create models for the new coinage.[18] After completing Lady Spencer's commission, by most accounts, Pistrucci suggested to Pole that an appropriate subject for the sovereign would be Saint George.[19][20] He created a head, in jasper, of King George III, to be used as model for the sovereign and the smaller silver coins. He had prepared a model in wax of Saint George and the Dragon for use on the crown; this was adapted for the sovereign. The Royal Mint's engravers were not able to successfully reproduce Pistrucci's imagery in steel, and the sculptor undertook the engraving of the dies himself.[21]
Pistrucci's George and Dragon design
The Saint George and Dragon design sketched on paper
Pistrucci's original sketch for the sovereign
Pistrucci's design for the reverse of the sovereign features Saint George on horseback. His left hand clutches the rein of the horse's bridle, and he does not wear armour, other than on his lower legs and feet, with his toes bare. Further protection is provided by the helmet, with, on early issues, a streamer or plume of hair floating behind. Also flowing behind the knight is his chlamys, or cloak; it is fastened in front by a fibula. George's right shoulder bears a balteus for suspending the gladius, the sword that he grasps in his right hand.[22] He is otherwise naked[23]—the art critic John Ruskin later considered it odd that the saint should be unclothed going into such a violent encounter.[24] The saint's horse appears to be half attacking, half shrinking from the dragon, which lies wounded by George's spear and in the throes of death.[23]
A gold coin with a man's head on one side and a crowned heraldic shield on the other
The sovereign replaced the guinea.
The original 1817 design had the saintly knight still carrying part of his broken spear. This was changed to a sword when the garter that originally surrounded the design was eliminated in 1821, and George is intended to have broken his spear earlier in the encounter with the dragon.[25] Also removed in 1821 was the plume of hair, or streamer, behind George's helmet; it was restored in 1887,[26] modified in 1893 and 1902,[22] and eliminated in 2009.[27]
The George and Dragon design is in the Neoclassical style. When Pistrucci created the coin, Neoclassicism was all the rage in London, and he may have been inspired by the Elgin Marbles, which were exhibited from 1807, and which he probably saw soon after his arrival in London. Pistrucci's sovereign was unusual for a British coin of the 19th century in not having a heraldic design, but this was consistent with Pole's desire to make the sovereign look as different from the guinea as possible.[28]
Circulation years (1817–1914)
Early years (1817–1837)
"Whereas We have thought fit to order that certain Pieces of Gold Money should be coined, which should be called 'Sovereigns or Twenty Shilling Pieces', each of which should be of the Value of Twenty Shillings, and that each Piece should be of the Weight of Five Pennyweights Three Grains 2,740⁄10,000 Troy Weight of Standard Gold ... And We have further thought fit to order that every such Piece of Gold Money, so ordered to be coined as aforesaid, shall have for the Obverse Impression the Head of His Majesty, with the Inscription 'Georgius III. D.G: Britanniar. Rex. F. D.' and the Date of the Year; and for the Reverse the Image of St. George armed sitting on Horseback encountering the Dragon with a Spear, the said Device being placed within the ennobled Garter, bearing the Motto 'Honi soit qui mal y pense', with a newly invented Graining on the Edge of the Piece."
—Proclamation of George, Prince Regent
1 July 1817[29]
When the sovereign entered circulation in late 1817, it was not initially popular, as the public preferred the convenience of the banknotes the sovereign had been intended to replace. Lack of demand meant that mintages dropped from 2,347,230 in 1818 to 3,574 the following year.[30] Another reason why few sovereigns were struck in 1819 was a proposal, eventually rejected, by economist David Ricardo to eliminate gold as a coinage metal, though making it available on demand from the Bank of England. Once this plan was abandoned in 1820, the Bank encouraged the circulation of gold sovereigns, but acceptance among the British public was slow. As difficulties over the exchange of wartime banknotes were overcome, the sovereign became more popular, and with low-value banknotes becoming scarcer, in 1826 Parliament prohibited the issuance of notes with a value of less than five pounds in England and Wales.[31]
The early sovereigns were heavily exported; in 1819, Robert Peel estimated that of the some £5,000,000 in gold struck in France since the previous year, three-quarters of the gold used had come from the new British coinage, melted down.[31] Many more sovereigns were exported to France in the 1820s as the metal alloyed with the gold included silver, which could be profitably recovered, with the gold often returned to Britain and struck again into sovereigns. Beginning in 1829, the Mint was able to eliminate the silver, but the drain on sovereigns from before then continued.[32]
George III died in January 1820, succeeded by George, Prince Regent, as George IV. Mint officials decided to continue to use the late king's head on coinage for the remainder of the year.[33] For King George IV's coinage, Pistrucci modified the George and Dragon reverse, eliminating the surrounding Garter ribbon and motto, with a reeded border substituted. Pistrucci also modified the figure of the saint, placing a sword in his hand in place of the broken lance seen previously, eliminating the streamer from his helmet, and refining the look of the cloak.[34]
The obverse design for George IV's sovereigns featured a "Laureate head" of George IV, based on the bust Pistrucci had prepared for the Coronation medal. The new version was authorised by an Order in Council of 5 May 1821. These were struck every year between 1821 and 1825, but the king was unhappy with the depiction of him and requested a new one be prepared, based on a more flattering bust by Francis Chantrey. Pistrucci refused to copy the work of another artist and was barred from further work on the coinage. Second Engraver (later Chief Engraver) William Wyon was assigned to translate Chantrey's bust into a coin design, and the new sovereign came into use during 1825. It did not bear the George and Dragon design, as the new Master of the Mint, Thomas Wallace, disliked several of the current coinage designs, and had Jean Baptiste Merlen of the Royal Mint prepare new reverse designs.[35] The new reverse for the sovereign featured the Ensigns Armorial, or royal arms of the United Kingdom, crowned, with the lions of England seen in two of the quarters, balanced by those of Scotland and the harp of Ireland. Set on the shield are the arms of Hanover,[c] again crowned, depicting the armorial bearings of Brunswick, Lüneburg and Celle. The George and Dragon design would not again appear on the sovereign until 1871.[36]
William IV's accession in 1830 upon the death of his brother George IV led to new designs for the sovereign, with the new king's depiction engraved by William Wyon based on a bust by Chantrey. Two slightly different busts were used, with what is usually called the "first bust" used for most 1831 circulating pieces (the first year of production) and some from 1832, with the "second bust" used for the prototype pattern coins that year, as well as for proof coins of 1831, some from 1832 and taking over entirely by 1833. The reverse shows another depiction by Merlen of the Ensigns Armorial, with the date accompanied by the Latin word Anno, or "in the year". These were struck every year until the year of the king's death, 1837.[37]
Victorian era
Gold coin with Queen Victoria on the obverse and the royal shield within a wreath on the other
1842 "Shield reverse" sovereign
The accession of Queen Victoria in 1837 ended the personal union between Britain and Hanover, as under the latter's Salic Law, a woman could not take the Hanoverian throne. Thus, both sides of the sovereign had to be changed.[38] Wyon designed his "Young head" portrait of the Queen, which he engraved, for the obverse, and Merlen engraved the reverse, depicting the royal arms inside a wreath, and likely played some part in designing it. The new coin was approved on 26 February 1838, and with the exception of 1840 and 1867, the "shield back" sovereign was struck at the Royal Mint in London every year from 1838 to 1874.[39] Sovereigns struck in London with the shield design between 1863 and 1874 bear small numbers under the shield, representing which coinage die was used. Records of why the numbers were used are not known to survive, with one widely printed theory that they were used to track die wear.[40] George Frederick Ansell states in his 1870 book The Royal Mint, Its Workings, Conduct, And Operations Fully And Practically Explained that "the reverse die has been made to carry, in addition to its recognised device, a small number, with a view to determine at which coining press, and on what particular day, the numbered die was used, that bad work might be traced to an individual."[41]
By 1850, some £94 million in sovereigns and half sovereigns had been struck and circulated widely, well beyond Britain's shores, a dispersion aided by the British government, who saw the sovereign's use as an auxiliary to their imperialist ambitions. Gold is a soft metal, and the hazards of circulation tended to make sovereigns lightweight over time. In 1838, when the legacy of James Smithson was converted into gold in preparation for transmission to the United States, American authorities requested recently-struck sovereigns, likely to maximise the quantity of gold when the sovereigns were melted after arrival in the United States.[42]
The weight of a newly-struck sovereign was intended to be 123.274 grains (7.98805 g). It ceased to be legal currency for £1 if found to weigh less than 1221⁄2 grains[43][d] (i.e. a deficiency of 11⁄2 pence in gold per sovereign). By the early 1840s, the Bank of England estimated that twenty per cent of the gold coins that came into its hands were lightweight. In part to boost the sovereign's reputation in trade, the Bank undertook a programme of recoinage, melting lightweight gold coins and using the gold for new, full-weight ones.[42] Between 1842 and 1845, the Bank withdrew and had recoined some £14 million in lightweight gold, about one-third the amount of that metal in circulation. This not only kept the sovereign to standard, it probably removed most of the remaining guineas still in commerce.[47] The unlucky holder of a lightweight gold coin could only turn it in as bullion, would lose at least 11⁄2 pence because of the lightness and often had to pay an equal amount to cover the Bank of England's costs.[48] There was also increased quality control within the Royal Mint; by 1866, every gold and silver coin was weighed individually.[49] The result of these efforts was that the sovereign became, in Sir John Clapham's later phrase, the "chief coin of the world".[50]
The California Gold Rush and other discoveries of the 1840s and 1850s boosted the amount of available gold and also the number of sovereigns struck, with £150 million in sovereigns and half sovereigns coined between 1850 and 1875. The wear problem continued: it was estimated that, on average, a sovereign became lightweight after fifteen years in circulation. The Coinage Act 1870 tightened standards at the Royal Mint, requiring sovereigns to be individually tested at the annual Trial of the Pyx rather than in bulk.[51] These standards resulted in a high rejection rate for newly coined sovereigns, though less than for the half sovereign, which sometimes exceeded 50 per cent.[52] When the Royal Mint was rebuilt in 1882, a decisive factor in shutting down production for renovation rather than moving to a new mint elsewhere was the Bank of England's report that there was an abnormally large stock of sovereigns and that no harm would result if they could not be coined in London for a year.[53] Advances in technology allowed sovereigns to be individually weighed by automated machines at the Bank of England by the 1890s, and efforts to keep the coin at full weight were aided by an 1889 Act of Parliament which allowed redemption of lightweight gold coin at full face value, with the loss from wear to fall upon the government.[51] The Coinage Act 1889 also authorised the Bank of England to redeem worn gold coins from before Victoria's reign, but on 22 November 1890 all gold coins from before her reign were called in by Royal Proclamation and demonetised effective 28 February 1891.[54] Owing to an ongoing programme to melt and recoin lightweight pieces, estimates of sovereigns in trade weighing less than the legal minimum had fallen to about four per cent by 1900.[51]
A metal balance with slots
Counterfeit detector. A fake will pass one test (weight or fit) and fail the other.
The sovereign was seen in fiction: in Dickens' Oliver Twist, Mrs Bumble is paid £25 in sovereigns her information. Joseph Conrad, in his novels set in Latin America, refers several times to ship's captains keeping sovereigns as a ready store of value. Although many sovereigns were melted down for recoining on reaching a foreign land (as were those for the Smithsonian) it was regarded as a circulating coin in dozens of British colonies and even in nations such as Brazil and Portugal;[55] the latter accepted it at a value of 4,500 reis.[56]
In 1871, the Deputy Master of the Mint, Charles Fremantle, restored the Pistrucci George and Dragon design to the sovereign, as part of a drive to beautify the coinage.[57] The return of Saint George was approved by the Queen, and authorised by an Order in Council dated 14 January 1871. The two designs were struck side by side in London from 1871 to 1874, and at the Australian branch mints until 1887, after which the Pistrucci design alone was used.[58] The saint returned to the rarely-struck two- and five-pound pieces in 1887, and was placed on the half sovereign in 1893.[59] Wyon's "Young head" of Queen Victoria for the sovereign's obverse was struck from 1838 until 1887, when it was replaced by the "Jubilee head" by Joseph Boehm.[60] That obverse was criticised and was replaced in 1893 by the "Old head" by Thomas Brock.[61] Victoria's death in 1901 led to a new obverse for her son and successor, Edward VII by George William de Saulles, which began production in 1902; Edward's death in 1910 necessitated a new obverse for his son, George V by Bertram Mackennal. Pistrucci's George and Dragon design continued on the reverse.[62]
Branch mint coinage
Gold coin with a crown on one side and the denomination of one pound on the other
The 1852 Adelaide Pound (on average) contains 8.75 grams of gold (0.9170 fine) and weighs 0.2580 of an ounce.[63]
The 1851 discovery of gold in Australia quickly led to calls from the local populace for the establishment of a branch of the Royal Mint in the colonies there. Authorities in Adelaide did not wait for London to act, but set up an assay office, striking what became known as the "Adelaide Pound". In 1853, an Order in Council approved the establishment of the Sydney Mint; the Melbourne Mint would follow in 1872, and the Perth Mint in 1899.[64] The act which regulated currency in New South Wales came into force on 18 July 1855 and stipulated that the gold coins were to be called sovereigns and half sovereigns. They were also to be the same weight, fineness and value as other sovereigns.[65]
Sovereign, 1855, from original gold coin, Sydney Mint, State Library of New South Wales, [http://archival.sl.nsw.gov.au/Details/archive/110493273 SAFE/DN/C 1
Early issues for Sydney, until 1870, depicted a bust of Victoria similar to those struck in Britain, but with a wreath of banksia, native to Australia, in her hair. The reverse was distinctive as well, with the name of the mint, the word AUSTRALIA and the denomination ONE SOVEREIGN on the reverse.[64] These coins were not initially legal tender outside Australia, as there were concerns about the design and about the light colour of the gold used (due to a higher percentage of silver in the alloy) but from 1866 Australian sovereigns were legal tender alongside those struck in London. Beginning in 1870, the designs were those used in London, though with a mint mark "S" or "M" (or, later, "P") denoting their origin. The mints at Melbourne and Sydney were allowed to continue striking the shield design even though it had been abandoned at the London facility, and did so until 1887 due to local popularity. The large issues of the colonial mints meant that by 1900, about forty per cent of the sovereigns circulating in Britain were from Australia.[64][66] Dies for the Australian coinage were made at London.[67]
Gold coin in a red card
2017-I sovereign in card of issue
Following the Klondike Gold Rush, the Canadian Government asked for the establishment of a Royal Mint branch in Canada. It was not until 1908 that what is now the Royal Canadian Mint, in Ottawa, opened, and it struck sovereigns with the mint mark "C" from 1908 to 1919, except 1912 and 1915, each year in small numbers.[68] Branch mints at Bombay (1918; mint mark "I") and Pretoria (1923–1932; mint mark "SA") also struck sovereigns. Melbourne and Perth stopped striking sovereigns after 1931, with Sydney having closed in 1926.[69] The 1932 sovereigns struck at Pretoria were the last to be issued intended as currency at their face value.[70]
To address the high demand for gold coins in the Indian market, which does not allow gold coins to be imported,[71] the minting of gold sovereigns in India with mint mark I has resumed since 2013. Indian/Swiss joint venture company MMTC-PAMP mints under licence in its facility close to Delhi with full quality control from the Royal Mint.[72] The coins are legal tender in the United Kingdom.[73]
Trade coin (1914–1979)
Poster depicting the gold sovereign with text urging support for the British cause in the First World War
First World War propaganda poster featuring the sovereign coin
In the late 19th century, several Chancellors of the Exchequer had questioned the wisdom of having much of Britain's stock of gold used in coinage. Lord Randolph Churchill proposed relying less on gold coinage and moving to high-value silver coins, and the short-lived double florin or four-shilling piece is a legacy of his views. Churchill's successor, George Goschen, urged issuing banknotes to replace the gold coins, saying he preferred £20 million in gold in the Bank of England to thirty million sovereigns in the hands of the public. Fears that widespread forgery of banknotes would shake confidence in the pound ended his proposal.[74]
In March 1914, John Maynard Keynes noted that the large quantities of gold arriving from South Africa were making the sovereign even more important. "The combination of the demand for sovereigns in India and Egypt with London's situation as the distributing centre of the South African gold is rapidly establishing the sovereign as the predominant gold coin of the world. Possibly it may be destined to hold in the future the same kind of international position as was held for several centuries, in the days of a silver standard, by the Mexican dollar."[75]
As Britain moved towards war in the July Crisis of 1914, many sought to convert Bank of England notes into gold, and the bank's reserves of the metal fell from £27 million on 29 July to £11 million on 1 August. Following the declaration of war against Germany on 4 August, the government circulated one-pound and ten-shilling banknotes in place of the sovereign and half sovereign.[76] Restrictions were placed on sending gold abroad, and the melting-down of coin made an offence.[77] Not all were enthusiastic about the change from gold to paper: J.J. Cullimore Allen, in his 1965 book on sovereigns, recalled meeting his first payroll after the change to banknotes, with the workers dubious about the banknotes and initially asking to be paid in gold. Allen converted five sovereigns from his own pocket into notes, and the workers made no further objection.[78] Conversion into gold was not forbidden, but the Chancellor, David Lloyd George, made it clear that such actions would be unpatriotic and would harm the war effort. Few insisted on payment in gold in the face of such appeals, and by mid-1915, the sovereign was rarely seen in London commerce. The coin was depicted on propaganda posters, which urged support for the war.[76]
Although sovereigns continued to be struck at London until the end of 1917, they were mostly held as part of the nation's gold reserves, or were paid out for war debts to the United States.[79] They were still used as currency in some foreign countries, especially in the Middle East.[80] Sovereigns continued to be struck at the Australian mints, where different economic circumstances prevailed. After the war, the sovereign did not return to commerce in Britain, with the pieces usually worth more as gold than as currency. In 1925, the Chancellor, Winston Churchill, secured the passage of the Gold Standard Act 1925, restoring Britain to that standard, but with gold to be kept in reserve rather than as a means of circulation. The effort failed—Churchill regarded it as the worst mistake of his life—but some lightweight sovereigns were melted and restruck dated 1925, and were released only later. Many of the Australian pieces struck in the postwar period were to back currency, while the South African sovereigns were mostly for export and to pay workers at the gold mines.[81][82]
By the time Edward VIII came to the throne in 1936, there was no question of issuing sovereigns for circulation, but pieces were prepared as part of the traditional proof set of coins issued in the coronation year. With a bust of King Edward by Humphrey Paget and the date 1937, these sovereigns were not authorised by royal proclamation prior to Edward VIII's abdication in December 1936, and are considered pattern coins.[83] Extremely rare, one sold in 2020 for £1,000,000, setting what was then a record (since broken) for a British coin.[84][85] Sovereigns in proof condition dated 1937 were struck for Edward's brother and successor, George VI, also designed by Paget, the only sovereigns to bear George's effigy. The 1925-dated George V sovereign was restruck in 1949, 1951 and 1952, lowering the value of the original, of which only a few had hitherto been known.[86] These were struck to meet the need for sovereigns, and to maintain the skills of the Royal Mint in striking them.[87]
The sovereign remained popular as a trade coin in the Middle East and elsewhere following the Second World War. The small strikings of 1925-dated sovereigns in the postwar period were not enough to meet the demand, which was met in part by counterfeiters in Europe and the Middle East, who often put full value of gold in the pieces. A counterfeiting prosecution was brought, to which the defence was made that the sovereign was no longer a current coin. The judge directed an acquittal although the sovereign remained legal tender under the Coinage Act 1870.[88]
Sovereigns were struck in 1953, the coronation year of Elizabeth II, bearing the portrait of her by Mary Gillick, though the gold pieces were placed only in the major museums.[89] A 1953 sovereign sold at auction in 2014 for £384,000.[73] In 1957, the Treasury decided to defend the status of the sovereign, both by continuing prosecutions and by issuing new pieces with the current date.[89] Elizabeth II sovereigns bearing Gillick's portrait were struck as bullion pieces between 1957 and 1959, and from 1962 to 1968.[90] The counterfeiting problem was minimised by the striking of about 45,000,000 sovereigns by 1968, and efforts by Treasury solicitors which resulted in the sovereign's acceptance as legal tender by the highest courts of several European nations.[91] In 1966, the Wilson government placed restrictions on the holding of gold coins to prevent hoarding against inflation, with collectors required to obtain a licence from the Bank of England. This proved ineffective, as it drove gold dealing underground, and was abandoned in 1970.[92]
The sovereign's role in popular culture continued: in the 1957 novel From Russia, with Love, Q issues James Bond with a briefcase, the handle of which contains 50 sovereigns. When held at gunpoint on the Orient Express by Red Grant, Bond uses the gold to distract Grant, leading to the villain's undoing.[93] The sovereign survived both decimalisation and the Royal Mint's move from London to Llantrisant, Wales. The last of the Gillick sovereigns had been struck in 1968; when production resumed in 1974, it was with a portrait by Arnold Machin.[94] The last coin minted at Tower Hill, in 1975, was a sovereign.[95]
Bullion and collectors coin (1979 to present)
From 1979, the sovereign was issued as a coin for the bullion market, but was also struck by the Royal Mint in proof condition for collectors, and this issuance of proof coins has continued annually. In 1985, the Machin portrait of the Queen was replaced by one by Raphael Maklouf.[96] Striking of bullion sovereigns had been suspended after 1982, and so the Maklouf portrait, struck every year but 1989 until the end of 1997, is seen on the sovereign only in proof condition.[97] In 1989, a commemorative sovereign, the first, was issued for the 500th anniversary of Henry VII's sovereign. The coin, designed by Bernard Sindall, evokes the designs of that earlier piece, showing the Queen enthroned and facing front, as Henry appeared on the old English sovereign. The reverse of the 1489 piece depicts a double Tudor rose fronted by the royal arms; a similar design with updated arms graces the reverse of the 1989 sovereign.[98]
Reverse of the 2020 sovereign
Ian Rank-Broadley designed the fourth bust of the Queen to be used on the sovereign, and this went into use in 1998 and was used until 2015. Bullion sovereigns began to be issued again in 2000, and this has continued.[99] A special reverse design was used in 2002 for the Golden Jubilee, with an adaptation of the royal arms on a shield by Timothy Noad recalling the 19th-century "shield back" sovereigns.[100] The years 2005 and 2012 (the latter, the Queen's Diamond Jubilee) saw interpretations of the George and Dragon design, the first by Noad, the later by Paul Day. In 2009, the reverse was re-engraved using tools from the reign of George III in the hope of better capturing Pistrucci's design.[101] A new portrait of the Queen by Jody Clark was introduced during 2015, and some sovereigns were issued with the new bust. The most recent special designs, in 2016 and 2017, were only for collectors. The 2016 collector's piece, for the Queen's 90th birthday, has a one-year-only portrait of her on the obverse designed by James Butler. The 2017 collector's piece returned to Pistrucci's original design of 1817 for the modern sovereign's 200th birthday, with the Garter belt and motto. A piedfort was also minted, and the bullion sovereign struck at Llantrisant, though retaining the customary design, was given a privy mark with the number 200.[102][103] For 2022, a reverse design by Noad in honour of the Queen's Platinum Jubilee, depicting his interpretation of the Royal Coat of Arms was used.[104]
In 2017, a collection of 633 gold sovereigns and 280 half sovereigns was discovered to have been hoarded inside an upright piano which had been donated to a community college in Shropshire, England.[105] The coins, which date from 1847 to 1915, were found by a technician who had been asked to tune the piano, 'stitched into seven cloth packets and a leather drawstring purse' under the piano's keyboard. Despite inquiries being made as to who could have stored the coins, no owner or claimants were found.
Collecting, other use and tax treatment
A gold coin
Sydney Mint sovereign, 1857
Many of the variant designs of the sovereign since 1989 have been intended to appeal to coin collectors, as have the other gold coins based on the sovereign, from the quarter sovereign to the five-sovereign piece. To expedite matters, the Royal Mint is authorised to sell gold sovereigns directly to the public, rather than having its output channelled through the Bank of England as was once the case.[106] As a legal tender coin, the sovereign is exempt from capital gains tax for UK residents.[107]
As well as being used as a circulating coin, the sovereign has entered fashion: some men in the 19th century placed one on their pocket watch chains (seen as a sign of integrity),[108] and others carried them in a small purse linked to the chain.[109] These customs vanished with the popularisation of the wrist watch. Women also have worn sovereigns, as bangles or ear rings.[108] In the 21st century, the wearing of a sovereign ring has been seen as a sign of chav culture.[110]
The staff carried by the Gentleman or Lady Usher of the Black Rod (known as Black Rod) as a symbol of office, and used to strike the door of the House of Commons of the United Kingdom during the State Opening of Parliament, has a sovereign inset into one of its ends.[111]
Coin auction houses deal in rare sovereigns of earlier date, as do specialist dealers.[112] As well as the 1937 Edward VIII and 1953 Elizabeth II sovereigns, rare dates in the series include the 1819,[113] and the 1863 piece with the number "827" on the obverse in place of William Wyon's initials. The 827 likely is an ingot number, used for some sort of experiment, though research has not conclusively established this.[114] Few 1879 sovereigns were struck at London, and those that remain are often well-worn.[58] Only 24,768 of the Adelaide Pound were struck; surviving specimens are rare and highly prized.[115] The sovereign itself has been the subject of commemoration; in 2005, the Perth Mint issued a gold coin with face value A$25, reproducing the reverse design of the pre-1871 Sydney Mint sovereigns.[116]
See also
iconMoney portal Numismatics portal flagUnited Kingdom portal
Crown gold
Gold Britannia coin
Krugerrand
Notes
A historic claim only. See Hubbard.
The indenture, dated 6 February 1817, directed that there be 9341⁄2 sovereigns struck from twenty troy pounds of standard gold - or 46.725 sovereigns to a troy pound.[14]
The British monarch also ruled Hanover between 1714 and 1837. See Seaby, pp. 134, 153.
Changed in 1821 from a minimum weight of 122.75 grains as experience had shown that to be two small a tolerance, and reaffirmed at the 1821 figure in 1838[44][45] and in 1843.[46]
References
Celtel & Gullbekk, p. 61.
"Tudor sovereign". The Royal Mint Museum. Retrieved 4 March 2018.
Clancy, p. 15.
Marsh 2017, pp. 3–4.
Clancy, p. 41.
Clancy, p. 45.
Clancy, p. 47.
Clancy, p. 57.
Clancy, pp. 52–55.
Seaby, pp. 116–117.
Marsh 2017, p. 7.
Clancy, p. 55.
Clancy, p. 56.
"Parliamentary Papers". Her Majesty's Stationery Office. 1866. p. 27.
ODNB.
Marsh 1996, p. 15.
Farey September 2014, p. 52.
Celtel & Gullbekk, p. 91.
Clancy, p. 58.
Rodgers, pp. 43–44.
Marsh 2017, p. 8.
Allen, p. 13.
Celtel & Gullbekk, p. 92.
Clancy, p. 63.
Marsh 2017, pp. 10–16.
Celtel & Gullbekk, p. 109.
Marsh 2017, p. 101.
Clancy, pp. 62–63.
Ruding, Rogers (1819). Supplement to the Annals of the Coinage of Britain. London: John Nichols and Son. pp. 47–48. OCLC 778858975. Archived from the original on 19 February 2018.
Rodgers, p. 44.
Clancy, pp. 64–67.
Craig, p. 304.
Marsh 2017, p. 13.
Celtel & Gullbekk, p. 98.
Clancy, pp. 66–67.
Celtel & Gullbekk, p. 99.
Marsh 2017, pp. 21–27.
Clancy, p. 69.
Marsh 2017, pp. 27–38.
Marsh 2017, pp. 31, 29.
Ansell 1870, p. 66.
Clancy, pp. 70–71.
Seyd, Ernest (1868). Bullion and Foreign Exchanges Theoretically and Practically Considered: Followed by a Defence of the Double Valuation, with Special Reference to the Proposed System of Universal Coinage. E. Wilson. p. 291. OCLC 574480898.
Ruding, Rogers (1819). Supplement to the Annals of the Coinage of Britain. London: John Nichols and Son. p. 48. OCLC 778858975. Archived from the original on 19 February 2018.
Ruding, Rogers (1840). Supplement to the Annals of the Coinage of Britain. Vol. 2 (Third ed.). London: John Hearne. pp. 128, 132. OCLC 771752141.
"By the Queen, a Proclamation". London Gazette. 10 October 1843. p. 3284.
Dyer & Gaspar, p. 484.
Craig, p. 310.
Craig, p. 322.
Dyer & Gaspar, p. 511.
Clancy, pp. 76–77.
Dyer & Gaspar, pp. 520–521.
Dyer & Gaspar, p. 525.
Hayter, p. 433.
Clancy, pp. 73, 78–79, 85.
Browne, W. A. (1899). "The Merchants' Handbook of Money, Weights and Measures, with Their British Equivalents".
Clancy, p. 73.
Marsh 2017, p. 47.
Clancy, p. 65.
Marsh 2017, pp. 47, 57.
Marsh 2017, p. 64.
Marsh 2017, pp. 69, 77.
Cuhaj, George S., ed. (2009). Standard Catalog of World Coins 1801–1900 (6 ed.). Iola, Wisconsin: Krause Publications. p. 82. ISBN 978-0-89689-940-7.
Celtel & Gullbekk, pp. 131–132.
Pamphlets issued by the New South Wales Commissioners for the World's Columbian Exposition, Chicago (1 ed.). New South Wales. Commission for the World's Columbian Exposition. 1893. p. 137.
Marsh 2017, pp. 28–29, 64.
Dyer & Gaspar, pp. 530–531.
Marsh 2017, pp. 69–72, 81.
Marsh 2017, pp. 78–87.
Rodgers, p. 46.
"Chindambaram rules out lifting ban on import of gold coins". The Hindu. 22 October 2013. Archived from the original on 1 March 2014.
"MMTC PAMP Sovereign web page". MMTC PAMP. 7 September 2014. Archived from the original on 26 December 2014. Retrieved 21 February 2018.
Rodgers, p. 47.
Clancy, p. 78.
Keynes, p. 155.
Clancy, pp. 89–91.
Josset, pp. 143–144.
Allen, p. 7.
Marsh 2017, p. 77.
Josset, p. 141.
Clancy, pp. 92–93.
Allen, p. 10.
Marsh 2017, pp. 88–89.
"'Never meant to exist': Edward VIII coin bought for record £1m". The Guardian. PA Media. 17 January 2020. Retrieved 17 January 2020.
"1937 British gold sovereign realizes $2.28M record in Heritage March 2021 sale". CoinNews.net. 29 March 2021. Retrieved 30 March 2021.
Marsh 2017, pp. 90–91.
Clancy, p. 95.
Allen, pp. 15–16.
Clancy, pp. 95–97.
Marsh 2017, p. 97.
Dyer & Gaspar, p. 598.
Seaby, p. 173.
Clancy, pp. 94–95.
Marsh 2017, pp. 94, 97–98.
Clancy, p. 99.
Celtel & Gullbekk, pp. 116–117.
Marsh 2017, pp. 98–99.
Celtel & Gullbekk, pp. 118–119.
Marsh 2017, pp. 100–101.
Clancy, p. 102.
Marsh 2017, pp. 104–105.
Marsh 2017, pp. 95–96, 106.
Clancy, pp. 102–103.
Alexander, Michael (15 November 2021). "United Kingdom: New 2022 gold sovereigns released — the first coins in the Platinum Jubilee Collection". Coin Update. Retrieved 6 March 2022.
Davies, Caroline (20 April 2017). "Mystery gold sovereign hoard found in piano declared to be treasure". The Guardian. Retrieved 2 February 2019.
Clancy, pp. 99–103.
"Gold and capital gains tax". Royal Mint. 15 November 2015. Retrieved 6 March 2018.
Allen, p. 14.
"Sovereign cases: Sampson Mordan & Co Ltd". Antiques in Oxford. Retrieved 5 October 2020.
Rumsey, Nichola; Harcourt, Diana, eds. (2014). Oxford Handbook of the Psychology of Appearance. Oxford, Oxfordshire: Oxford University Press. p. 12. ISBN 978-0-19-872322-6.
"Gentleman Usher of the Black Rod". Armchair Travel Complany. Retrieved 16 November 2020.
Marsh 2017, pp. 186–192.
Marsh 2017, p. 10.
Marsh 2017, p. 31.
Allen, pp. 56–57.
Celtel & Gullbekk, p. 133.
Bibliography
Allen, James John Cullimore (1965). Sovereigns of the British Empire. London, United Kingdom: Spink & Son, Ltd. OCLC 493287074.
Ansell, G. F. (1870). The Royal Mint: its working, conduct, and operations, fully and practically explained. London: Effingham Wilson.
Celtel, André; Gullbekk, Svein H. (2006). The Sovereign and its Golden Antecedents. Oslo, Norway: Monetarius. ISBN 978-82-996755-6-7.
Clancy, Kevin (2017) [2015]. A History of the Sovereign: Chief Coin of the World (second ed.). Llantrisant, Wales: Royal Mint Museum. ISBN 978-1-869917-00-5.
Craig, John (2010) [1953]. The Mint (paperback ed.). Cambridge, United Kingdom: Cambridge University Press. ISBN 978-0-521-17077-2.
Dyer, G.P.; Gaspar, G.P. (1992), "Reform, the New Technology and Tower Hill", in Challis, C.E. (ed.), A New History of the Royal Mint, Cambridge, United Kingdom: Cambridge University Press, pp. 398–606, ISBN 978-0-521-24026-0
Farey, Roderick (September 2014). "Benedetto Pistrucci (1782–1855), Part 1". Coin News: 51–53.
Hayter, Henry Heylyn (1891). Victorian Year-Book for 1890–91 (18th ed.). Melbourne: Sands & McDougall Ltd.
Hubbard, Arnold (14 July 2003). "How George III lost France: Or, Why Concessions Never Make Sense". Electric Review: A High Tory Online Journal of Politics, Art and Literature. Archived from the original on 5 December 2008.
Josset, Christopher Robert (1962). Money in Britain. London: Frederick Warne and Co Ltd. OCLC 923302099.
Keynes, John Maynard (March 1914). "Currency in 1912". The Economic Journal. Royal Economic Society. 24 (93): 152–157. doi:10.2307/2221837. JSTOR 2221837.
Marsh, Michael A. (1996). Benedetto Pistrucci: Principal Engraver and Chief Medallist of the Royal Mint, 1783–1855. Hardwick, Cambridgeshire: Michael A. Marsh (Publications). ISBN 978-0-9506929-2-0.
Marsh, Michael A. (2017) [1980]. The Gold Sovereign (revised ed.). Exeter, Devon: Token Publishing Ltd. ISBN 978-1-908828-36-1.
Pollard, Graham (2004). "Pistrucci, Benedetto". Oxford Dictionary of National Biography (online ed.). Oxford University Press. doi:10.1093/ref:odnb/22314. Retrieved 3 July 2017. (Subscription or UK public library membership required.)
Rodgers, Kerry (June 2017). "Britain's Gold Sovereign". Coin News: 43–47.
Seaby, Peter (1985). The Story of British Coinage. London: B. A. Seaby Ltd. ISBN 978-0-900652-74-5.
External links
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Owls are birds from the order Strigiformes (/ˈstrɪdʒəfɔːrmiːz/), which includes over 200 species of mostly solitary and nocturnal birds of prey typified by an upright stance, a large, broad head, binocular vision, binaural hearing, sharp talons, and feathers adapted for silent flight. Exceptions include the diurnal northern hawk-owl and the gregarious burrowing owl.
Owls hunt mostly small mammals, insects, and other birds, although a few species specialize in hunting fish. They are found in all regions of the Earth except the polar ice caps and some remote islands.
Owls are divided into two families: the true (or typical) owl family, Strigidae, and the barn-owl family, Tytonidae.
A group of owls is called a "parliament".[1]
Anatomy
Burrowing owl (Athene cunicularia)
Cross-eyed owl
Owls possess large, forward-facing eyes and ear-holes, a hawk-like beak, a flat face, and usually a conspicuous circle of feathers, a facial disc, around each eye. The feathers making up this disc can be adjusted to sharply focus sounds from varying distances onto the owls' asymmetrically placed ear cavities. Most birds of prey have eyes on the sides of their heads, but the stereoscopic nature of the owl's forward-facing eyes permits the greater sense of depth perception necessary for low-light hunting. Although owls have binocular vision, their large eyes are fixed in their sockets—as are those of most other birds—so they must turn their entire heads to change views. As owls are farsighted, they are unable to clearly see anything within a few centimeters of their eyes. Caught prey can be felt by owls with the use of filoplumes—hairlike feathers on the beak and feet that act as "feelers". Their far vision, particularly in low light, is exceptionally good.
Owls can rotate their heads and necks as much as 270°. Owls have 14 neck vertebrae compared to seven in humans, which makes their necks more flexible. They also have adaptations to their circulatory systems, permitting rotation without cutting off blood to the brain: the foramina in their vertebrae through which the vertebral arteries pass are about 10 times the diameter of the artery, instead of about the same size as the artery as in humans; the vertebral arteries enter the cervical vertebrae higher than in other birds, giving the vessels some slack, and the carotid arteries unite in a very large anastomosis or junction, the largest of any bird's, preventing blood supply from being cut off while they rotate their necks. Other anastomoses between the carotid and vertebral arteries support this effect.[2][3]
The smallest owl—weighing as little as 31 g (1+3⁄32 oz) and measuring some 13.5 cm (5+1⁄4 in)—is the elf owl (Micrathene whitneyi).[4] Around the same diminutive length, although slightly heavier, are the lesser known long-whiskered owlet (Xenoglaux loweryi) and Tamaulipas pygmy owl (Glaucidium sanchezi).[4] The largest owls are two similarly sized eagle owls; the Eurasian eagle-owl (Bubo bubo) and Blakiston's fish owl (Bubo blakistoni). The largest females of these species are 71 cm (28 in) long, have a 190 cm (75 in) wing span, and weigh 4.2 kg (9+1⁄4 lb).[4][5][6][7][8]
Different species of owls produce different sounds; this distribution of calls aids owls in finding mates or announcing their presence to potential competitors, and also aids ornithologists and birders in locating these birds and distinguishing species. As noted above, their facial discs help owls to funnel the sound of prey to their ears. In many species, these discs are placed asymmetrically, for better directional location.
Owl plumage is generally cryptic, although several species have facial and head markings, including face masks, ear tufts, and brightly colored irises. These markings are generally more common in species inhabiting open habitats, and are thought to be used in signaling with other owls in low-light conditions.[9]
Sexual dimorphism
Sexual dimorphism is a physical difference between males and females of a species. Females owls are typically larger than the males.[10] The degree of size dimorphism varies across multiple populations and species, and is measured through various traits, such as wing span and body mass.[10] Overall, female owls tend to be slightly larger than males, for reasons not fully agreed.
One theory suggests that selection has led males to be smaller because it allows them to be efficient foragers. The ability to obtain more food is advantageous during breeding season. In some species, female owls stay at their nest with their eggs while it is the responsibility of the male to bring back food to the nest.[11] If food is scarce, the male first feeds himself before feeding the female.[12] Small birds, which are agile, are an important source of food for owls. Male burrowing owls have been observed to have longer wing chords than females, despite being smaller than females.[12] Furthermore, owls have been observed to be roughly the same size as their prey.[12] This has also been observed in other predatory birds,[11] which suggests that owls with smaller bodies and long wing chords have been selected for because of the increased agility and speed that allows them to catch their prey.[citation needed]
Another popular theory suggests that females have not been selected to be smaller like male owls because of their sexual roles. In many species, female owls may not leave the nest. Therefore, females may have a larger mass to allow them to go for a longer period of time without starving. For example, one hypothesized sexual role is that larger females are more capable of dismembering prey and feeding it to their young, hence female owls are larger than their male counterparts.[10]
A different theory suggests that the size difference between male and females is due to sexual selection: since large females can choose their mate and may violently reject a male's sexual advances, smaller male owls that have the ability to escape unreceptive females are more likely to have been selected.[12]
If the character is stable, there can be different optimums for both sexes. Selection operates on both sexes at the same time; therefore it is necessary to explain not only why one of the sexes is relatively bigger, but also why the other sex is smaller.[13] If owls are still evolving toward smaller bodies and longer wing chords, according to V. Geodakyan's Evolutionary Theory of Sex, males should be more advanced on these characters. Males are viewed as an evolutionary vanguard of a population, and sexual dimorphism on the character, as an evolutionary “distance” between the sexes. “Phylogenetic rule of sexual dimorphism” states that if there exists a sexual dimorphism on any character, then the evolution of this trait goes from the female form toward the male one.[14]
Hunting adaptations
All owls are carnivorous birds of prey and live on diets of insects, small rodents and lagomorphs. Some owls are also specifically adapted to hunt fish. They are very adept in hunting in their respective environments. Since owls can be found in nearly all parts of the world and across a multitude of ecosystems, their hunting skills and characteristics vary slightly from species to species, though most characteristics are shared among all species.[citation needed]
Flight and feathers
Most owls share an innate ability to fly almost silently and also more slowly in comparison to other birds of prey. Most owls live a mainly nocturnal lifestyle and being able to fly without making any noise gives them a strong advantage over prey alert to the slightest sound in the night. A silent, slow flight is not as necessary for diurnal and crepuscular owls given that prey can usually see an owl approaching. Owls’ feathers are generally larger than the average birds’ feathers, have fewer radiates, longer pennulum, and achieve smooth edges with different rachis structures.[15] Serrated edges along the owl's remiges bring the flapping of the wing down to a nearly silent mechanism. The serrations are more likely reducing aerodynamic disturbances, rather than simply reducing noise.[16] The surface of the flight feathers is covered with a velvety structure that absorbs the sound of the wing moving. These unique structures reduce noise frequencies above 2 kHz,[17] making the sound level emitted drop below the typical hearing spectrum of the owl's usual prey[17][18] and also within the owl's own best hearing range.[19][20] This optimizes the owl's ability to silently fly to capture prey without the prey hearing the owl first as it flies, and to hear any noise the prey makes. It also allows the owl to monitor the sound output from its flight pattern.
A great horned owl with wet feathers, waiting out a rainstorm
The feather adaption that allows silent flight means that barn owl feathers are not waterproof. To retain the softness and silent flight, the barn owl cannot use the preen oil or powder dust that other species use for waterproofing. In wet weather, they cannot hunt and this may be disastrous during the breeding season. Barn owls are frequently found drowned in livestock drinking troughs, since they land to drink and bathe, but are unable to climb out. Owls can struggle to keep warm, because of their lack of waterproofing, so large numbers of downy feathers help them to retain body heat.[21]
Vision
Eyesight is a particular characteristic of the owl that aids in nocturnal prey capture. Owls are part of a small group of birds that live nocturnally, but do not use echolocation to guide them in flight in low-light situations. Owls are known for their disproportionally large eyes in comparison to their skulls. An apparent consequence of the evolution of an absolutely large eye in a relatively small skull is that the eye of the owl has become tubular in shape. This shape is found in other so-called nocturnal eyes, such as the eyes of strepsirrhine primates and bathypelagic fishes.[22] Since the eyes are fixed into these sclerotic tubes, they are unable to move the eyes in any direction.[23] Instead of moving their eyes, owls swivel their heads to view their surroundings. Owls' heads are capable of swiveling through an angle of roughly 270°, easily enabling them to see behind them without relocating the torso.[23] This ability keeps bodily movement at a minimum, thus reduces the amount of sound the owl makes as it waits for its prey. Owls are regarded as having the most frontally placed eyes among all avian groups, which gives them some of the largest binocular fields of vision. Owls are farsighted and cannot focus on objects within a few centimeters of their eyes.[22][24] These mechanisms are only able to function due to the large-sized retinal image.[25] Thus, the primary nocturnal function in the vision of the owl is due to its large posterior nodal distance; retinal image brightness is only maximized to the owl within secondary neural functions.[25] These attributes of the owl cause its nocturnal eyesight to be far superior to that of its average prey.[25]
Hearing
A great horned owl perched on the top of a Joshua tree at evening (twilight) in the Mojave Desert, U.S.
Owls exhibit specialized hearing functions and ear shapes that also aid in hunting. They are noted for asymmetrical ear placements on the skull in some genera. Owls can have either internal or external ears, both of which are asymmetrical. Asymmetry has not been reported to extend to the middle or internal ear of the owl. Asymmetrical ear placement on the skull allows the owl to pinpoint the location of its prey. This is especially true for strictly nocturnal species such as the barn owls Tyto or Tengmalm's owl.[23] With ears set at different places on its skull, an owl is able to determine the direction from which the sound is coming by the minute difference in time that it takes for the sound waves to penetrate the left and right ears.The Hearing of the Barn Owl The owl turns its head until the sound reaches both ears at the same time, at which point it is directly facing the source of the sound. This time difference between ears is about 30 microseconds. Behind the ear openings are modified, dense feathers, densely packed to form a facial ruff, which creates an anterior-facing, concave wall that cups the sound into the ear structure.[26] This facial ruff is poorly defined in some species, and prominent, nearly encircling the face, in other species. The facial disk also acts to direct sound into the ears, and a downward-facing, sharply triangular beak minimizes sound reflection away from the face. The shape of the facial disk is adjustable at will to focus sounds more effectively.[23]
The prominences above a great horned owl's head are commonly mistaken as its ears. This is not the case; they are merely feather tufts. The ears are on the sides of the head in the usual location (in two different locations as described above).
Talons
While the auditory and visual capabilities of the owl allow it to locate and pursue its prey, the talons and beak of the owl do the final work. The owl kills its prey using these talons to crush the skull and knead the body.[23] The crushing power of an owl's talons varies according to prey size and type, and by the size of the owl. The burrowing owl (Athene cunicularia), a small, partly insectivorous owl, has a release force of only 5 N. The larger barn owl (Tyto alba) needs a force of 30 N to release its prey, and one of the largest owls, the great horned owl (Bubo virginianus) needs a force over 130 N to release prey in its talons.[27] An owl's talons, like those of most birds of prey, can seem massive in comparison to the body size outside of flight. The Tasmanian masked owl has some of the proportionally longest talons of any bird of prey; they appear enormous in comparison to the body when fully extended to grasp prey.[28] An owl's claws are sharp and curved. The family Tytonidae has inner and central toes of about equal length, while the family Strigidae has an inner toe that is distinctly shorter than the central one.[27] These different morphologies allow efficiency in capturing prey specific to the different environments they inhabit.
Beak
The beak of the owl is short, curved, and downward-facing, and typically hooked at the tip for gripping and tearing its prey. Once prey is captured, the scissor motion of the top and lower bill is used to tear the tissue and kill. The sharp lower edge of the upper bill works in coordination with the sharp upper edge of the lower bill to deliver this motion. The downward-facing beak allows the owl's field of vision to be clear, as well as directing sound into the ears without deflecting sound waves away from the face.[29]
Camouflage
The snowy owl has effective snow camouflage
The coloration of the owl's plumage plays a key role in its ability to sit still and blend into the environment, making it nearly invisible to prey. Owls tend to mimic the coloration and sometimes the texture patterns of their surroundings, the barn owl being an exception. The snowy owl (Bubo scandiacus) appears nearly bleach-white in color with a few flecks of black, mimicking their snowy surroundings perfectly, while the speckled brown plumage of the tawny owl (Strix aluco) allows it to lie in wait among the deciduous woodland it prefers for its habitat. Likewise, the mottled wood-owl (Strix ocellata) displays shades of brown, tan and black, making the owl nearly invisible in the surrounding trees, especially from behind. Usually, the only tell-tale sign of a perched owl is its vocalizations or its vividly colored eyes.
Behavior
Comparison of an owl (left) and hawk (right) remex.
The serrations on the leading edge of an owl's flight feathers reduce noise
Owl eyes each have nictitating membranes that can move independently of each other, as seen on this spotted eagle-owl in Johannesburg, South Africa.
Most owls are nocturnal, actively hunting their prey in darkness. Several types of owls are crepuscular—active during the twilight hours of dawn and dusk; one example is the pygmy owl (Glaucidium). A few owls are active during the day, also; examples are the burrowing owl (Speotyto cunicularia) and the short-eared owl (Asio flammeus).
Much of the owls' hunting strategy depends on stealth and surprise. Owls have at least two adaptations that aid them in achieving stealth. First, the dull coloration of their feathers can render them almost invisible under certain conditions. Secondly, serrated edges on the leading edge of owls' remiges muffle an owl's wing beats, allowing an owl's flight to be practically silent. Some fish-eating owls, for which silence has no evolutionary advantage, lack this adaptation.
An owl's sharp beak and powerful talons allow it to kill its prey before swallowing it whole (if it is not too big). Scientists studying the diets of owls are helped by their habit of regurgitating the indigestible parts of their prey (such as bones, scales, and fur) in the form of pellets. These "owl pellets" are plentiful and easy to interpret, and are often sold by companies to schools for dissection by students as a lesson in biology and ecology.[30]
Breeding and reproduction
Owl eggs typically have a white color and an almost spherical shape, and range in number from a few to a dozen, depending on species and the particular season; for most, three or four is the more common number. In at least one species, female owls do not mate with the same male for a lifetime. Female burrowing owls commonly travel and find other mates, while the male stays in his territory and mates with other females.[31]
Evolution and systematics
A great horned owl (Bubo virginianus) sleeping during daytime in a hollow tree
Recent phylogenetic studies place owls within the clade Telluraves, most closely related to the Accipitrimorphae and the Coraciimorphae,[32][33] although the exact placement within Telluraves is disputed.[34][35]
See below cladogram:
Telluraves
Accipitrimorphae
Cathartiformes (New World vultures)Vintage Vulture Drawing white background.jpg
Accipitriformes (hawks and relatives)Golden Eagle Illustration white background.jpg
Strigiformes (owls)Cuvier-12-Hibou à huppe courte.jpg
Coraciimorphae
Coliiformes (mouse birds)
Cavitaves
Leptosomiformes (cuckoo roller)
Trogoniformes (trogons and quetzals)Harpactes fasciatus 1838 white background.jpg
Picocoraciae
Bucerotiformes (hornbills and relatives)
Picodynastornithes
Coraciiformes (kingfishers and relatives)Cuvier-46-Martin-pêcheur d'Europe.jpg
Piciformes (woodpeckers and relatives)
Australaves
Cariamiformes (seriemas)Cariama cristata 1838 white background.jpg
Eufalconimorphae
Falconiformes (falcons)NewZealandFalconBuller white background.jpg
Psittacopasserae
Psittaciformes (parrots)Pyrrhura lucianii - Castelnau 2.jpg
Passeriformes (passerines)Cuvier-33-Moineau domestique.jpg
Cladogram of Telluraves relationships based on Braun & Kimball (2021)[36]
Some 220 to 225 extant species of owls are known, subdivided into two families: 1. true owls or typical owls family (Strigidae) and 2. barn-owls family (Tytonidae). Some entirely extinct families have also been erected based on fossil remains; these differ much from modern owls in being less specialized or specialized in a very different way (such as the terrestrial Sophiornithidae). The Paleocene genera Berruornis and Ogygoptynx show that owls were already present as a distinct lineage some 60–57 million years ago (Mya), hence, possibly also some 5 million years earlier, at the extinction of the non-avian dinosaurs. This makes them one of the oldest known groups of non-Galloanserae landbirds. The supposed "Cretaceous owls" Bradycneme and Heptasteornis are apparently non-avialan maniraptors.[37]
During the Paleogene, the Strigiformes radiated into ecological niches now mostly filled by other groups of birds.[clarification needed] The owls as known today, though, evolved their characteristic morphology and adaptations during that time, too. By the early Neogene, the other lineages had been displaced by other bird orders, leaving only barn-owls and typical owls. The latter at that time were usually a fairly generic type of (probably earless) owls similar to today's North American spotted owl or the European tawny owl; the diversity in size and ecology found in typical owls today developed only subsequently.
Around the Paleogene-Neogene boundary (some 25 Mya), barn-owls were the dominant group of owls in southern Europe and adjacent Asia at least; the distribution of fossil and present-day owl lineages indicates that their decline is contemporary with the evolution of the different major lineages of true owls, which for the most part seems to have taken place in Eurasia. In the Americas, rather, an expansion of immigrant lineages of ancestral typical owls occurred.
The supposed fossil herons "Ardea" perplexa (Middle Miocene of Sansan, France) and "Ardea" lignitum (Late Pliocene of Germany) were more probably owls; the latter was apparently close to the modern genus Bubo. Judging from this, the Late Miocene remains from France described as "Ardea" aureliensis should also be restudied.[38] The Messelasturidae, some of which were initially believed to be basal Strigiformes, are now generally accepted to be diurnal birds of prey showing some convergent evolution toward owls. The taxa often united under Strigogyps[39] were formerly placed in part with the owls, specifically the Sophiornithidae; they appear to be Ameghinornithidae instead.[40][41][42]
The ancient fossil owl Palaeoglaux artophoron
For fossil species and paleosubspecies of extant taxa, see the genus and species articles. For a full list of extant and recently extinct owls, see the article List of owl species.
Unresolved and basal forms (all fossil)
Berruornis (Late Paleocene of France) basal? Sophornithidae?
Strigiformes gen. et sp. indet. (Late Paleocene of Zhylga, Kazakhstan)[43]
Primoptynx (Early Eocene of Wyoming, U.S.)[44]
Palaeoglaux (Middle – Late Eocene of West-Central Europe) own family Palaeoglaucidae or Strigidae?
Palaeobyas (Late Eocene/Early Oligocene of Quercy, France) Tytonidae? Sophiornithidae?[citation needed]
Palaeotyto (Late Eocene/Early Oligocene of Quercy, France) Tytonidae? Sophiornithidae?[citation needed]
Strigiformes gen. et spp. indet. (Early Oligocene of Wyoming, U.S.)[38]
Ogygoptyngidae
Ogygoptynx (Middle/Late Paleocene of Colorado, U.S.)
Protostrigidae
Eostrix (Early Eocene of United States, Europe, and Mongolia). E. gulottai is the smallest known fossil (or living) owl.[45]
Minerva (Middle – Late Eocene of western U.S.) formerly Protostrix, includes "Aquila" ferox, "Aquila" lydekkeri, and "Bubo" leptosteus
Oligostrix (mid-Oligocene of Saxony, Germany)
Sophiornithidae
Sophiornis
Tytonidae
Genus Tyto – the barn owls, grass owls and masked owls, stand up to 500 mm (20 in) tall; some 15 extant species and possibly one recently extinct
Genus Phodilus – the bay owls, two to three extant species and possibly one recently extinct
Fossil genera
Nocturnavis (Late Eocene/Early Oligocene) includes "Bubo" incertus
Selenornis (Late Eocene/Early Oligocene) – includes "Asio" henrici
Necrobyas (Late Eocene/Early Oligocene – Late Miocene) includes "Bubo" arvernensis and Paratyto
Prosybris (Early Oligocene? – Early Miocene)
Placement unresolved
Tytonidae gen. et sp. indet. "TMT 164" (Middle Miocene) – Prosybris?
Strigidae
A long-eared owl (Asio otus) in an erect pose
The laughing owl (Ninox albifacies), last seen in 1914
Genus Aegolius – the saw-whet owls, four species
Genus Asio – the eared owls, eight species
Genus Athene – two to four species (depending on whether the genera Speotyto and Heteroglaux are included or not)
Genus Bubo – the horned owls, eagle-owls and fish-owls; paraphyletic with the genera Nyctea, Ketupa, and Scotopelia, some 25 species
Genus Glaucidium – the pygmy owls, about 30–35 species
Genus Gymnasio – the Puerto Rican owl
Genus Gymnoglaux – the bare-legged owl or Cuban screech-owl
Genus Lophostrix – the crested owl
Genus Jubula – the maned owl
Genus Megascops – the screech owls, some 20 species
Genus Micrathene – the elf owl
Genus Ninox – the Australasian hawk-owls or boobooks, some 20 species
Genus Otus – the scops owls; probably paraphyletic, about 45 species
Genus Pseudoscops – the Jamaican owl
Genus Psiloscops – the flammulated owl
Genus Ptilopsis – the white-faced owls, two species
Genus Pulsatrix – the spectacled owls, three species
Genus Strix – the earless owls, about 15 species, including four previously assigned to Ciccaba
Genus Surnia – the northern hawk-owl
Genus Taenioptynx - the collared owlet
Genus Uroglaux – the Papuan hawk-owl
Genus Xenoglaux – the long-whiskered owlet
Extinct genera
Genus Grallistrix – the stilt-owls, four species; prehistoric
Genus Ornimegalonyx – the Caribbean giant owls, one to two species; prehistoric
Fossil genera
Mioglaux (Late Oligocene? – Early Miocene of West-Central Europe) – includes "Bubo" poirreiri
Intutula (Early/Middle – ?Late Miocene of Central Europe) – includes "Strix/Ninox" brevis
Alasio (Middle Miocene of Vieux-Collonges, France) – includes "Strix" collongensis
Oraristrix – the Brea owl (Late Pleistocene)
Placement unresolved
"Otus/Strix" wintershofensis: fossil (Early/Middle Miocene of Wintershof West, Germany) – may be close to extant genus Ninox[38]
"Strix" edwardsi – fossil (Middle/Late? Miocene)
"Asio" pygmaeus – fossil (Early Pliocene of Odessa, Ukraine)
Strigidae gen. et sp. indet. UMMP V31030 (Late Pliocene) – Strix/Bubo?
the Ibizan owl, Strigidae gen. et sp. indet. – prehistoric[46]
Symbolism and mythology
African cultures
Among the Kikuyu of Kenya, it was believed that owls were harbingers of death. If one saw an owl or heard its hoot, someone was going to die. In general, owls are viewed as harbingers of bad luck, ill health, or death. The belief is widespread even today.[47]
Asia
In Mongolia, the owl is regarded as a benign omen. In one story, Genghis Khan was hiding from enemies in a small coppice when an owl roosted in the tree above him, which caused his pursuers to think no man could be hidden there.[48]
In modern Japan, owls are regarded as lucky and are carried in the form of a talisman or charm.[49]
Hootum Pyanchar Naksha by Kaliprasanna Singha (1841–1870), first published in 1861, is a book of social commentaries influential in Bengali literature. The name literally means "Sketches by a Watching Owl".
Sumerian and ancient Semitic cultures
In Sumerian, Akkadian, and Babylonian culture, the owl was associated with Lilith.[50] This association also occurs in the Bible (in some translations) in Isaiah 34:14.[51]
Ancient European and modern Western culture
The modern West generally associates owls with wisdom and vigilance. This link goes back at least as far as Ancient Greece, where Athens, noted for art and scholarship, and Athena, Athens' patron goddess and the goddess of wisdom, had the owl as a symbol.[52] Marija Gimbutas traces veneration of the owl as a goddess, among other birds, to the culture of Old Europe, long pre-dating Indo-European cultures.[53]
T. F. Thiselton-Dyer, in his 1883 Folk-lore of Shakespeare, says that "from the earliest period it has been considered a bird of ill-omen," and Pliny tells us how, on one occasion, even Rome itself underwent a lustration, because one of them strayed into the Capitol. He represents it also as a funereal bird, a monster of the night, the very abomination of human kind. Virgil describes its death-howl from the top of the temple by night, a circumstance introduced as a precursor of Dido's death. Ovid, too, constantly speaks of this bird's presence as an evil omen; and indeed the same notions respecting it may be found among the writings of most of the ancient poets."[54] A list of "omens drear" in John Keats' Hyperion includes the "gloom-bird's hated screech."[55] Pliny the Elder reports that owl's eggs were commonly used as a hangover cure.[56]
One of the etymologies offered for the name of the German folk hero Till Eulenspiegel is that it means "Mirror for Owls".
An owl-shaped protocorinthian aryballos, c. 640 BCE, from Greece
An owl-shaped protocorinthian aryballos, c. 640 BCE, from Greece
A Roman owl mosaic from Italica, Spain
A Roman owl mosaic from Italica, Spain
A Manises plate, c. 1535. A fantastical owl wearing a crown, a characteristic Manises design during the first half of the 16th century
A Manises plate, c. 1535. A fantastical owl wearing a crown, a characteristic Manises design during the first half of the 16th century
The Little Owl, 1506, by Albrecht Dürer
The Little Owl, 1506, by Albrecht Dürer
Wooden Owls of Natungram, West Bengal, India. The wooden owl is an integral part of an ancient and indigenous tradition and art form in Bengal along with its auspicious association with Goddess of wealth, Laxmi.
Wooden Owls of Natungram, West Bengal, India. The wooden owl is an integral part of an ancient and indigenous tradition and art form in Bengal along with its auspicious association with Goddess of wealth, Laxmi.
Hinduism
The Hindu goddess Lakshmi with the owl
In Hinduism, an owl is the vahana (mount) of the goddess Lakshmi, specially in eastern region of India.[57] Owl is considered a symbol of wealth, prosperity, wisdom, good luck and Fortune. This is the reason why Owl is seen with Godden Lakshmi, who is also the goddess the fortune, wealth and prosperity. The Goddess Lakshmi, is known to have a White Barn Owl as her vahana.
At the same time, owls are also associated with evil times in Hinduism. At times, Chamunda (fearsome form of Chandi) is depicted seated on an owl, her vahana (mount or vehicle). Hindus believed that owls are messengers of death.[58]
Native American cultures
People often allude to the reputation of owls as bearers of supernatural danger when they tell misbehaving children, "the owls will get you",[59] and in most Native American folklore, owls are a symbol of death.
According to the Apache and Seminole tribes, hearing owls hooting is considered the subject of numerous "bogeyman" stories told to warn children to remain indoors at night or not to cry too much, otherwise the owl may carry them away.[60][61] In some tribal legends, owls are associated with spirits of the dead, and the bony circles around an owl's eyes are said to comprise the fingernails of apparitional humans. Sometimes owls are said to carry messages from beyond the grave or deliver supernatural warnings to people who have broken tribal taboos.[62]
The Aztecs and the Maya, along with other natives of Mesoamerica, considered the owl a symbol of death and destruction. In fact, the Aztec god of death, Mictlantecuhtli, was often depicted with owls.[63] There is an old saying in Mexico that is still in use:[64] Cuando el tecolote canta, el indio muere ("When the owl cries/sings, the Indian dies"). The Popol Vuh, a Mayan religious text, describes owls as messengers of Xibalba (the Mayan "Place of Fright").[65]
The belief that owls are messengers and harbingers of the dark powers is also found among the Hočągara (Winnebago) of Wisconsin.[66] When in earlier days the Hočągara committed the sin of killing enemies while they were within the sanctuary of the chief's lodge, an owl appeared and spoke to them in the voice of a human, saying, "From now on, the Hočągara will have no luck." This marked the beginning of the decline of their tribe.[67] An owl appeared to Glory of the Morning, the only female chief of the Hočąk nation, and uttered her name. Soon after, she died.[68][69]
According to the culture of the Hopi, a Uto-Aztec tribe, taboos surround owls, which are associated with sorcery and other evils.
The Ojibwe tribes, as well as their Aboriginal Canadian counterparts, used an owl as a symbol for both evil and death. In addition, they used owls as a symbol of very high status of spiritual leaders of their spirituality.[70]
The Pawnee tribes viewed owls as the symbol of protection from any danger within their realms.[70]
The Puebloan peoples associated owls with Skeleton Man, the god of death and the spirit of fertility.[70]
The Yakama tribes use an owl as a totem, to guide where and how forests and natural resources are useful with management.[70]
Rodent control
A purpose-built owl-house or owlery at a farm near Morton on the Hill, England (2006)
Encouraging natural predators to control rodent population is a natural form of pest control, along with excluding food sources for rodents. Placing a nest box for owls on a property can help control rodent populations (one family of hungry barn owls can consume more than 3,000 rodents in a nesting season) while maintaining the naturally balanced food chain.[71]
Attacks on humans
Although humans and owls frequently live together in harmony, there have been incidents when owls have attacked humans. For example, in January 2013, a man from Inverness, Scotland suffered heavy bleeding and went into shock after being attacked by an owl, which was likely a 50-centimetre-tall (20 in) eagle-owl.[72] The photographer Eric Hosking lost his left eye after attempting to photograph a tawny owl, which inspired the title of his 1970 autobiography, An Eye for a Bird.
Conservation issues
See also: List of Strigiformes by population
The snowy owl is very endangered in Scandinavia[73] and Finland, where it is found only in northern Lapland.[74]
Almost all owls are listed in Appendix II of the international CITES treaty (the Convention on Illegal Trade in Endangered Species of Wild Fauna and Flora) with four species listed in Appendix I. Although owls have long been hunted, a 2008 news story from Malaysia indicates that the magnitude of owl poaching may be on the rise. In November 2008, TRAFFIC reported the seizure of 900 plucked and "oven-ready" owls in Peninsular Malaysia. Said Chris Shepherd, Senior Programme Officer for TRAFFIC's Southeast Asia office, "This is the first time we know of where 'ready-prepared' owls have been seized in Malaysia, and it may mark the start of a new trend in wild meat from the region. We will be monitoring developments closely." TRAFFIC commended the Department of Wildlife and National Parks in Malaysia for the raid that exposed the huge haul of owls. Included in the seizure were dead and plucked barn owls, spotted wood owls, crested serpent eagles, barred eagles, and brown wood owls, as well as 7,000 live lizards.[75]
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Székely T, Freckleton R. P., Reynolds J. D. (2004) Sexual selection explains Rensch's rule of size dimorphism in shorebirds. PNAS, 101, N. 33, p. 12224-12227.
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Stevenson, John (18 November 2020). "Small finlets on owl feathers point the way to less aircraft noise". Phys.org. Retrieved 20 November 2020.
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Willott J.F. (2001) Handbook of Mouse Auditory Research CRC Press ISBN 1420038737.
Dyson, M. L.; Klump, G. M.; Gauger, B. (April 1998). "Absolute hearing thresholds and critical masking ratios in the European barn owl: a comparison with other owls". Journal of Comparative Physiology. 182 (5): 695–702. doi:10.1007/s003590050214. S2CID 24641904.
Webster, Douglas B.; Fay, Richard R. (6 December 2012). "Hearing in Birds". The Evolutionary Biology of Hearing. Springer Science & Business Media. p. 547. ISBN 978-1-4612-2784-7.
Ian Hayward (27 July 2007). "Ask an expert: Are barn owl feathers waterproof?". RSPB. Retrieved 29 December 2015.
Walls G.L. (1942) The vertebrate eye and its adaptive radiation, Cranbook Institute of Science.
König, Claus, Friedhelm Weick & Jan-Hendrik Becking (1999). Owls: A guide to the owls of the world. Yale Univ Press, 1999. ISBN 0300079206
Hughes A. (1979). "A schematic eye for the rat". Vision Res. 19 (5): 569–588. doi:10.1016/0042-6989(79)90143-3. PMID 483586. S2CID 10317667.
Martin G.R. (1982). "An owl's eye: schematic optics and visual performance in Strix aluco L". J Comp Physiol. 145 (3): 341–349. doi:10.1007/BF00619338. S2CID 35039625.
Norberg, R.A. (31 August 1977). "Occurrence and independent evolution of bilateral ear asymmetry in owls and implications on owl taxonomy". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 280 (973): 375–408. Bibcode:1977RSPTB.280..375N. doi:10.1098/rstb.1977.0116.
Marti, C. D. (1974). "Feeding Ecology of Four Sympatric Owls" (PDF). The Condor. 76 (1): 45–61. doi:10.2307/1365983. JSTOR 1365983.
Einoder, Luke D. & Alastair M. M. Richardson (2007). "Aspects of the Hindlimb Morphology of Some Australian Birds of Prey: A Comparative and Quantitative Study". The Auk. 124 (3): 773–788. doi:10.1642/0004-8038(2007)124[773:AOTHMO]2.0.CO;2. S2CID 86011581.
Shamim1410 (22 April 2020). "Owl Bird Traits, Adaptations and Surprising Facts". Bird Baron. Retrieved 4 February 2022.
Owl Pellets in the Classroom: Safety Guidelines. carolina.com
Martin, Dennis J. (1973). "Selected Aspects of Burrowing Owl Ecology and Behavior". The Condor. 75 (4): 446–456. doi:10.2307/1366565. JSTOR 1366565. S2CID 55069283.
Jarvis, Erich D.; Mirarab, Siavash; Aberer, Andre J.; Li, Bo; Houde, Peter; Li, Cai; Ho, Simon Y. W.; Faircloth, Brant C.; Nabholz, Benoit; Howard, Jason T.; Suh, Alexander; Weber, Claudia C.; da Fonseca, Rute R.; Li, Jianwen; Zhang, Fang; Li, Hui; Zhou, Long; Narula, Nitish; Liu, Liang; Ganapathy, Ganesh; Boussau, Bastien; Bayzid, Md. Shamsuzzoha; Zavidovych, Volodymyr; Subramanian, Sankar; Gabaldón, Toni; Capella-Gutiérrez, Salvador; Huerta-Cepas, Jaime; Rekepalli, Bhanu; Munch, Kasper; et al. (12 December 2014). "Whole-genome analyses resolve early branches in the tree of life of modern birds". Science. 346 (6215): 1320–1331. Bibcode:2014Sci...346.1320J. doi:10.1126/science.1253451. PMC 4405904. PMID 25504713.
H Kuhl, C Frankl-Vilches, A Bakker, G Mayr, G Nikolaus, S T Boerno, S Klages, B Timmermann, M Gahr (2020) An unbiased molecular approach using 3’UTRs resolves the avian family-level tree of life. Molecular Biology and Evolution. https://doi.org/10.1093/molbev/msaa191
Prum, R.O. et al. (2015) A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526, 569–573.
Houde, Peter; Braun, Edward L.; Narula, Nitish; Minjares, Uriel; Mirarab, Siavash (2019). "Phylogenetic Signal of Indels and the Neoavian Radiation". Diversity. 11 (7): 108. doi:10.3390/d11070108. ISSN 1424-2818.
Braun, E.L. & Kimball, R.T. (2021) Data types and the phylogeny of Neoaves. Birds, 2(1), 1-22; https://doi.org/10.3390/birds2010001
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Mayr, Gerald (2005). ""Old World phorusrhacids" (Aves, Phorusrhacidae): a new look at Strigogyps ("Aenigmavis") sapea (Peters 1987)". PaleoBios. 25 (1): 11–16.
Alvarenga, Herculano M. F. & Höfling, Elizabeth (2003). "Systematic revision of the Phorusrhacidae (Aves: Ralliformes)". Papéis Avulsos de Zoologia. 43 (4): 55–91. doi:10.1590/S0031-10492003000400001.
Larco Herrera, Rafael and Berrin, Kathleen (1997) The Spirit of Ancient Peru Thames and Hudson, New York, ISBN 0500018022.
Peters, Dieter Stefan (January 2007). "The fossil family Ameghinornithidae (Mourer-Chauviré 1981): a short synopsis" (PDF). Journal of Ornithology. 148 (1): 25–28. doi:10.1007/s10336-006-0095-z. S2CID 27322057.
Mourer-Chauviré, Cécile (September 1994). "A large owl from the Paleocene of France". Palaeontology. 37 (2): 339–348. Retrieved 27 March 2021.
Mayr, Gerald; Gingerich, Philip; Smith, Thierry (July 2020). "Skeleton of a new owl from the early Eocene of North America (Aves, Strigiformes) with an accipitrid-like foot morphology". Journal of Vertebrate Paleontology. 40 (2): e1769116. doi:10.1080/02724634.2020.1769116. S2CID 222210173.
Gerald Mayr. "The world's smallest owl, the earliest unambiguous charadriiform bird, and other avian remains from the early Eocene Nanjemoy Formation of Virginia (USA)" (PDF). Phatfossils.com. Retrieved 24 March 2022.
Sánchez Marco, Antonio (2004). "Avian zoogeographical patterns during the Quaternary in the Mediterranean region and paleoclimatic interpretation" (PDF). Ardeola. 51 (1): 91–132.
"Owls in Lore and Culture – The Owl Pages". Owlpages.com.
Sparks & Soper (1979). Owls: their natural and unnatural history. p. 163. ISBN 0715349953.
"The Significance and Meaning of Owls in Japanese Culture". Owlcation.
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Isaiah 34:14
Deacy, Susan, and Villing, Alexandra (2001). Athena in the Classical World. Koninklijke Brill NV, Leiden, The Netherlands, ISBN 9004121420.
Gimbutas, Marija (2001) The living goddesses, University of California Press, p. 158. ISBN 0520927095.
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Dubow, Charles (1 January 2004). "Hangover Cures". Forbes. Archived from the original on 17 January 2003.
Chopra, Capt. Praveen (2017). Vishnu's Mount: Birds In Indian Mythology And Folklore. Notion Press. p. 109. ISBN 978-1-948352-69-7.
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Lenders, E. W. (1914). "The Myth of the 'Wah-ru-hap-ah-rah,' or the Sacred Warclub Bundle". Zeitschrift für Ethnologie. 46: 404–420 (409).
"Stikini, an owl monster of Seminole folklore". Native-languages.org. Retrieved 25 October 2015.
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"Native American Indian Owl Legends, Meaning and Symbolism from the Myths of Many Tribes". Native-languages.org. 25 July 2008. Retrieved 25 October 2015.
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"Cuando el tecolote canta, el indio muere". La Cronica. 27 July 2008. Archived from the original on 3 September 2010.
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"Owls". Hočąk Encyclopedia.
Radin, Paul (1990 [1923]) The Winnebago Tribe, Lincoln: University of Nebraska Press, pp. 7–9 ISBN 0803257104.
Smith, David Lee (1997) Folklore of the Winnebago Tribe, Norman: University of Oklahoma Press, p. 160
"Glory of the Morning", Hočąk Encyclopedia.
"Owls in Lore and Culture". The Owl Pages. 31 October 2012. p. 3. Retrieved 25 October 2015.
"The Hungry Owl Project". Hungryowl.org. Archived from the original on 13 August 2003. Retrieved 9 April 2010.
"Man needed hospital treatment after owl attack". Daily Telegraph (London). 25 January 2013.
Juvonen, Arto; Muukkonen, Tomi; Peltomäki, Jari; Varesvuo, Markku (2009). Linnut vauhdissa (in Finnish). Tammi. pp. 178, 187. ISBN 978-951-31-4604-7.
Harrison, Colin; Greensmith, Alan (1995). Koko maailman linnut (in Finnish). Translated by Laine, Lasse J.; Nikander, Pekka. Helsinki Media. p. 198. ISBN 951-875-637-6.
"Wildlife Trade News – Huge haul of dead owls and live lizards in Peninsular Malaysia". TRAFFIC. 12 November 2008.
Further reading
Calaprice, Alice & Heinrich, Bernd (1990): Owl in the House: A Naturalist's Diary. Joy Street Books, Boston. ISBN 0-316-35456-2.
Duncan, James (2013). The Complete Book of North American Owls. Thunder Bay Press, San Diego. ISBN 9781607107262.
Duncan, James (2003). Owls of the World. Key Porter Books, Toronto. ISBN 1552632148.
Heinrich, Bernd (1987): One Man's Owl. Princeton, N.J.: Princeton University Press. ISBN 9780691084701. OCLC 15486687.
Johnsgard, Paul A. (2002): North American Owls: Biology and Natural History, 2nd ed. Smithsonian Institution Press, Washington, D.C. ISBN 1-56098-939-4.
Maslow, Jonathan Evan (1983): The Owl Papers, 1st Vintage Books ed. Vintage Books, New York. ISBN 0-394-75813-7.
Sibley, Charles Gald & Monroe, Burt L. Jr. (1990): Distribution and taxonomy of the birds of the world: A Study in Molecular Evolution. Yale University Press, New Haven, CT. ISBN 0-300-04969-2
External links
Wikimedia Commons has media related to Strigiformes.
Wikispecies has information related to Strigiformes.
Look up owl in Wiktionary, the free dictionary.
The Owl Pages
Owl Brain Atlas
Smithsonian Snowy Owl Info
World Owl Trust
Athenian Owl coins
Eurasia:
World of Owls – Northern Ireland's only owl, bird of prey and exotic animal centre
Current Blakiston's Fish Owl Research in Russia
North America:
List of Owl Species Breeding In North American and Owl Photos
Oceania:
iprimus info. re Australian owls and frogmouths
Bird
Temporal range:
Late Cretaceous – present, 72–0 Ma[1][2]
PreꞒ
Ꞓ
O
S
D
C
P
T
J
K
Pg
N
Possible Early Cretaceous or early Late Cretaceous origin based on molecular clock[3][4][5]
Bird Diversity 2013.png
About this image
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Clade: Sauropsida
Clade: Avemetatarsalia
Clade: Ornithurae
Class: Aves
Linnaeus, 1758[6]
Extant clades
Palaeognathae (ratites and tinamou)
Neognathae
Pangalloanserae (fowl)
Neoaves
Synonyms
Neornithes Gadow, 1883
Birds are a group of warm-blooded vertebrates constituting the class Aves (/ˈeɪviːz/), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight skeleton. Birds live worldwide and range in size from the 5.5 cm (2.2 in) bee hummingbird to the 2.8 m (9 ft 2 in) ostrich. There are about ten thousand living species, more than half of which are passerine, or "perching" birds. Birds have wings whose development varies according to species; the only known groups without wings are the extinct moa and elephant birds. Wings, which evolved from forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in some birds, including ratites, penguins, and diverse endemic island species. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly seabirds and some waterbirds, have further evolved for swimming.
Birds are feathered theropod dinosaurs and constitute the only known living dinosaurs. Likewise, birds are considered reptiles in the modern cladistic sense of the term, and their closest living relatives are the crocodilians. Birds are descendants of the primitive avialans (whose members include Archaeopteryx) which first appeared about 160 million years ago (mya) in China. According to DNA evidence, modern birds (Neornithes) evolved in the Middle to Late Cretaceous, and diversified dramatically around the time of the Cretaceous–Paleogene extinction event 66 mya, which killed off the pterosaurs and all non-avian dinosaurs.[5]
Many social species pass on knowledge across generations, which is considered a form of culture. Birds are social, communicating with visual signals, calls, and songs, and participating in such behaviours as cooperative breeding and hunting, flocking, and mobbing of predators. The vast majority of bird species are socially (but not necessarily sexually) monogamous, usually for one breeding season at a time, sometimes for years, and rarely for life. Other species have breeding systems that are polygynous (one male with many females) or, rarely, polyandrous (one female with many males). Birds produce offspring by laying eggs which are fertilised through sexual reproduction. They are usually laid in a nest and incubated by the parents. Most birds have an extended period of parental care after hatching.
Many species of birds are economically important as food for human consumption and raw material in manufacturing, with domesticated and undomesticated birds being important sources of eggs, meat, and feathers. Songbirds, parrots, and other species are popular as pets. Guano (bird excrement) is harvested for use as a fertiliser. Birds figure throughout human culture. About 120 to 130 species have become extinct due to human activity since the 17th century, and hundreds more before then. Human activity threatens about 1,200 bird species with extinction, though efforts are underway to protect them. Recreational birdwatching is an important part of the ecotourism industry.
Evolution and classification
Main article: Evolution of birds
Slab of stone with fossil bones and feather impressions
Archaeopteryx lithographica is often considered the oldest known true bird.
The first classification of birds was developed by Francis Willughby and John Ray in their 1676 volume Ornithologiae.[7] Carl Linnaeus modified that work in 1758 to devise the taxonomic classification system currently in use.[8] Birds are categorised as the biological class Aves in Linnaean taxonomy. Phylogenetic taxonomy places Aves in the clade Theropoda.[9]
Definition
Aves and a sister group, the order Crocodilia, contain the only living representatives of the reptile clade Archosauria. During the late 1990s, Aves was most commonly defined phylogenetically as all descendants of the most recent common ancestor of modern birds and Archaeopteryx lithographica.[10] However, an earlier definition proposed by Jacques Gauthier gained wide currency in the 21st century, and is used by many scientists including adherents to the PhyloCode. Gauthier defined Aves to include only the crown group of the set of modern birds. This was done by excluding most groups known only from fossils, and assigning them, instead, to the broader group Avialae,[11] in part to avoid the uncertainties about the placement of Archaeopteryx in relation to animals traditionally thought of as theropod dinosaurs.[citation needed]
Gauthier and de Queiroz[12] identified four different definitions for the same biological name "Aves", which is a problem. The authors proposed to reserve the term Aves only for the crown group consisting of the last common ancestor of all living birds and all of its descendants, which corresponds to meaning number 4 below. He assigned other names to the other groups.[citation needed]
Crocodiles
Birds
Turtles
Lizards (including snakes)
The birds' phylogenetic relationships to major living reptile groups
Aves can mean all archosaurs closer to birds than to crocodiles (alternately Avemetatarsalia)
Aves can mean those advanced archosaurs with feathers (alternately Avifilopluma)
Aves can mean those feathered dinosaurs that fly (alternately Avialae)
Aves can mean the last common ancestor of all the currently living birds and all of its descendants (a "crown group", in this sense synonymous with Neornithes)
Under the fourth definition Archaeopteryx, traditionally considered one of the earliest members of Aves, is removed from this group, becoming a non-avian dinosaur instead. These proposals have been adopted by many researchers in the field of palaeontology and bird evolution, though the exact definitions applied have been inconsistent. Avialae, initially proposed to replace the traditional fossil content of Aves, is often used synonymously with the vernacular term "bird" by these researchers.[13]
Maniraptoromorpha
†Coelurus
†Ornitholestes
Maniraptoriformes
†Ornithomimosauria
Maniraptora
†Alvarezsauridae
Pennaraptora
†Oviraptorosauria
Paraves
Cladogram showing the results of a phylogenetic study by Cau, 2018.[14]
Most researchers define Avialae as branch-based clade, though definitions vary. Many authors have used a definition similar to "all theropods closer to birds than to Deinonychus",[15][16] with Troodon being sometimes added as a second external specifier in case it is closer to birds than to Deinonychus.[17] Avialae is also occasionally defined as an apomorphy-based clade (that is, one based on physical characteristics). Jacques Gauthier, who named Avialae in 1986, re-defined it in 2001 as all dinosaurs that possessed feathered wings used in flapping flight, and the birds that descended from them.[12][18]
Despite being currently one of the most widely used, the crown-group definition of Aves has been criticised by some researchers. Lee and Spencer (1997) argued that, contrary to what Gauthier defended, this definition would not increase the stability of the clade and the exact content of Aves will always be uncertain because any defined clade (either crown or not) will have few synapomorphies distinguishing it from its closest relatives. Their alternative definition is synonymous to Avifilopluma.[19]
Dinosaurs and the origin of birds
Main article: Origin of birds
Paraves
†Scansoriopterygidae
†Eosinopteryx
Eumaniraptora
†Jinfengopteryx
†Aurornis
†Dromaeosauridae
†Troodontidae
Avialae
Cladogram following the results of a phylogenetic study by Cau et al., 2015[20]
Anchiornis huxleyi is an important source of information on the early evolution of birds in the Late Jurassic period.[21]
Based on fossil and biological evidence, most scientists accept that birds are a specialised subgroup of theropod dinosaurs[22] and, more specifically, members of Maniraptora, a group of theropods which includes dromaeosaurids and oviraptorosaurs, among others.[23] As scientists have discovered more theropods closely related to birds, the previously clear distinction between non-birds and birds has become blurred. Recent discoveries in the Liaoning Province of northeast China, which demonstrate many small theropod feathered dinosaurs, contribute to this ambiguity.[24][25][26]
Simplified phylogenetic tree showing the relationship between modern birds and dinosaurs [27]
The consensus view in contemporary palaeontology is that the flying theropods, or avialans, are the closest relatives of the deinonychosaurs, which include dromaeosaurids and troodontids.[28] Together, these form a group called Paraves. Some basal members of Deinonychosauria, such as Microraptor, have features which may have enabled them to glide or fly. The most basal deinonychosaurs were very small. This evidence raises the possibility that the ancestor of all paravians may have been arboreal, have been able to glide, or both.[29][30] Unlike Archaeopteryx and the non-avialan feathered dinosaurs, who primarily ate meat, recent studies suggest that the first avialans were omnivores.[31]
The Late Jurassic Archaeopteryx is well known as one of the first transitional fossils to be found, and it provided support for the theory of evolution in the late 19th century. Archaeopteryx was the first fossil to display both clearly traditional reptilian characteristics—teeth, clawed fingers, and a long, lizard-like tail—as well as wings with flight feathers similar to those of modern birds. It is not considered a direct ancestor of birds, though it is possibly closely related to the true ancestor.[32]
Early evolution
See also: List of fossil bird genera
White slab of rock left with cracks and impression of bird feathers and bone, including long paired tail feathers
Confuciusornis sanctus, a Cretaceous bird from China that lived 125 million years ago, is the oldest known bird to have a beak.[33]
Over 40% of key traits found in modern birds evolved during the 60 million year transition from the earliest bird-line archosaurs to the first maniraptoromorphs, i.e. the first dinosaurs closer to living birds than to Tyrannosaurus rex. The loss of osteoderms otherwise common in archosaurs and acquisition of primitive feathers might have occurred early during this phase.[14][34] After the appearance of Maniraptoromorpha, the next 40 million years marked a continuous reduction of body size and the accumulation of neotenic (juvenile-like) characteristics. Hypercarnivory became increasingly less common while braincases enlarged and forelimbs became longer.[14] The integument evolved into complex, pennaceous feathers.[34]
The oldest known paravian (and probably the earliest avialan) fossils come from the Tiaojishan Formation of China, which has been dated to the late Jurassic period (Oxfordian stage), about 160 million years ago. The avialan species from this time period include Anchiornis huxleyi, Xiaotingia zhengi, and Aurornis xui.[13]
The well-known probable early avialan, Archaeopteryx, dates from slightly later Jurassic rocks (about 155 million years old) from Germany. Many of these early avialans shared unusual anatomical features that may be ancestral to modern birds, but were later lost during bird evolution. These features include enlarged claws on the second toe which may have been held clear of the ground in life, and long feathers or "hind wings" covering the hind limbs and feet, which may have been used in aerial manoeuvreing.[35]
Avialans diversified into a wide variety of forms during the Cretaceous period. Many groups retained primitive characteristics, such as clawed wings and teeth, though the latter were lost independently in a number of avialan groups, including modern birds (Aves).[36] Increasingly stiff tails (especially the outermost half) can be seen in the evolution of maniraptoromorphs, and this process culminated in the appearance of the pygostyle, an ossification of fused tail vertebrae.[14] In the late Cretaceous, about 100 million years ago, the ancestors of all modern birds evolved a more open pelvis, allowing them to lay larger eggs compared to body size.[37] Around 95 million years ago, they evolved a better sense of smell.[38]
A third stage of bird evolution starting with Ornithothoraces (the "bird-chested" avialans) can be associated with the refining of aerodynamics and flight capabilities, and the loss or co-ossification of several skeletal features. Particularly significant are the development of an enlarged, keeled sternum and the alula, and the loss of grasping hands. [14]
Avialae
†Anchiornis
†Archaeopteryx
†Xiaotingia
†Rahonavis
†Jeholornis
†Jixiangornis
Euavialae
†Balaur
Avebrevicauda
†Zhongjianornis
†Sapeornis
Pygostylia
†Confuciusornithiformes
†Protopteryx
†Pengornis
Ornithothoraces
Cladogram following the results of a phylogenetic study by Cau et al., 2015[20]
Early diversity of bird ancestors
See also: Protobirds and Avialae
Ornithothoraces
†Enantiornithes
Euornithes
†Archaeorhynchus
Ornithuromorpha
†Patagopteryx
†Vorona
†Schizooura
†Hongshanornithidae
†Jianchangornis
†Songlingornithidae
†Gansus
†Apsaravis
Ornithurae
†Hesperornithes
†Ichthyornis
†Vegavis
Aves
Mesozoic bird phylogeny simplified after Wang et al., 2015's phylogenetic analysis[39]
Ichthyornis, which lived 93 million years ago, was the first known prehistoric bird relative preserved with teeth.
The first large, diverse lineage of short-tailed avialans to evolve were the Enantiornithes, or "opposite birds", so named because the construction of their shoulder bones was in reverse to that of modern birds. Enantiornithes occupied a wide array of ecological niches, from sand-probing shorebirds and fish-eaters to tree-dwelling forms and seed-eaters. While they were the dominant group of avialans during the Cretaceous period, enantiornithes became extinct along with many other dinosaur groups at the end of the Mesozoic era.[36]
Many species of the second major avialan lineage to diversify, the Euornithes (meaning "true birds", because they include the ancestors of modern birds), were semi-aquatic and specialised in eating fish and other small aquatic organisms. Unlike the Enantiornithes, which dominated land-based and arboreal habitats, most early euornithes lacked perching adaptations and seem to have included shorebird-like species, waders, and swimming and diving species.[citation needed]
The latter included the superficially gull-like Ichthyornis[40] and the Hesperornithiformes, which became so well adapted to hunting fish in marine environments that they lost the ability to fly and became primarily aquatic.[36] The early euornithes also saw the development of many traits associated with modern birds, like strongly keeled breastbones, toothless, beaked portions of their jaws (though most non-avian euornithes retained teeth in other parts of the jaws).[41] Euornithes also included the first avialans to develop true pygostyle and a fully mobile fan of tail feathers,[42] which may have replaced the "hind wing" as the primary mode of aerial maneuverability and braking in flight.[35]
A study on mosaic evolution in the avian skull found that the last common ancestor of all Neornithes might have had a beak similar to that of the modern hook-billed vanga and a skull similar to that of the Eurasian golden oriole. As both species are small aerial and canopy foraging omnivores, a similar ecological niche was inferred for this hypothetical ancestor.[43]
Diversification of modern birds
See also: Sibley–Ahlquist taxonomy of birds and dinosaur classification
Aves
Palaeognathae
Struthioniformes
Tinamiformes
Neognathae
Other birds (Neoaves)
Galloanserae
Anseriformes
Galliformes
Basal divergences of modern birds
based on Sibley-Ahlquist taxonomy
All modern birds lie within the crown group Aves (alternately Neornithes), which has two subdivisions: the Palaeognathae, which includes the flightless ratites (such as the ostriches) and the weak-flying tinamous, and the extremely diverse Neognathae, containing all other birds.[44] These two subdivisions have variously been given the rank of superorder,[45] cohort,[9] or infraclass.[46] Depending on the taxonomic viewpoint, the number of known living bird species varies anywhere from 9,800[47] to 10,758.[48]
The discovery of Vegavis from the Maastrichtian, the last stage of the Late Cretaceous proved that the diversification of modern birds started before the Cenozoic era.[49] The affinities of an earlier fossil, the possible galliform Austinornis lentus, dated to about 85 million years ago,[50] are still too controversial to provide a fossil evidence of modern bird diversification. In 2020, Asteriornis from the Maastrichtian was described, it appears to be a close relative of Galloanserae, the earliest diverging lineage within Neognathae.[1]
Most studies agree on a Cretaceous age for the most recent common ancestor of modern birds but estimates range from the Early Cretaceous[3][51] to the latest Late Cretaceous.[52][4] Similarly, there is no agreement on whether most of the early diversification of modern birds occurred before or after the Cretaceous–Palaeogene extinction event.[53] This disagreement is in part caused by a divergence in the evidence; most molecular dating studies suggests a Cretaceous evolutionary radiation, while fossil evidence points to a Cenozoic radiation (the so-called 'rocks' versus 'clocks' controversy). Previous attempts to reconcile molecular and fossil evidence have proved controversial,[53][54] but more recent estimates, using a more comprehensive sample of fossils and a new way of calibrating molecular clocks, showed that while according to some studies, modern birds originated early in the Late Cretaceous in Western Gondwana, a pulse of diversification in all major groups occurred around the Cretaceous–Palaeogene extinction event. Modern birds expanded from West Gondwana to the Laurasia through two routes. One route was an Antarctic interchange in the Paleogene. This can be confirmed with the presence of multiple avian groups in Australia and New Zealand. The other route was probably through North America, via land bridges, during the Paleocene. This allowed the expansion and diversification of Neornithes into the Holarctic and Paleotropics.[55] On the other hand, the occurrence of Asteriornis in the Northern Hemisphere challenges biogeographical hypotheses of a Gondwanan origin of crown birds.[1]
Classification of bird orders
See also: List of birds
Cladogram of modern bird relationships based on Braun & Kimball (2021)[56]
Aves
Palaeognathae
Struthioniformes (ostriches) Struthio camelus - Etosha 2014 (1) white background.jpg
Rheiformes (rheas) Rhea white background.jpg
Apterygiformes (kiwis) Little spotted kiwi, Apteryx owenii, Auckland War Memorial Museum white background.jpg
Tinamiformes (tinamous) NothuraDarwiniiSmit white background.jpg
Casuariiformes (emu and cassowaries) Emu RWD2 white background.jpg
Neognathae
Galloanserae
Galliformes (chickens and relatives) Red Junglefowl by George Edward Lodge white background.png
Anseriformes (ducks and relatives) Cuvier-97-Canard colvert.jpg
Neoaves
Mirandornithes
Phoenicopteriformes (flamingos)Cuvier-87-Flamant rouge.jpg
Podicipediformes (grebes)Podiceps cristatus Naumann white background.jpg
Columbimorphae
Columbiformes (pigeons and doves) Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg
Mesitornithiformes (mesites)Monias benschi 1912 white background.jpg
Pterocliformes (sandgrouse)Pterocles quadricinctus white background.jpg
Passerea
Otidiformes (bustards)Cayley Ardeotis australis flipped.jpg
Cuculiformes (cuckoos)British birds in their haunts (Cuculus canorus).jpg
Musophagiformes (turacos)Planches enluminées d'histoire naturelle (1765) (Tauraco persa).jpg
Gruiformes (rails and cranes)Cuvier-72-Grue cendrée.jpg
Charadriiformes (waders and relatives)D'Orbigny-Mouette rieuse et Bec-en-ciseaux white background.jpg
Opisthocomiformes (hoatzin)Cuvier-59-Hoazin huppé.jpg
Strisores
Caprimulgiformes (nightjars) Chordeiles acutipennis texensisAQBIP06CA.jpg
Vanescaves
Nyctibiiformes (potoos) NyctibiusBracteatusSmit.jpg
Steatornithiformes (oilbird) Steatornis caripensis MHNT ZON STEA 1.jpg
Podargiformes (frogmouths) Batrachostomus septimus 01.jpg
Daedalornithes
Aegotheliformes (owlet-nightjars) Aegotheles savesi.jpg
Apodiformes (swifts, treeswifts and hummingbirds) White-eared Hummingbird (Basilinna leucotis) white background.jpg
Phaethoquornithes
Eurypygimorphae
Phaethontiformes (tropicbirds)Cuvier-95-Phaeton à bec rouge.jpg
Eurypygiformes (sunbittern and kagu)Cuvier-72-Caurale soleil.jpg
Aequornithes
Gaviiformes[57] (loons) Loon (PSF).png
Austrodyptornithes
Procellariiformes (albatrosses and petrels) Thalassarche chlororhynchos 1838.jpg
Sphenisciformes (penguins) Chinstrap Penguin white background.jpg
Ciconiiformes (storks) Weißstorch (Ciconia ciconia) white background.jpg
Suliformes (boobies, cormorants, etc.) Cormorant in Strunjan, white background.png
Pelecaniformes (pelicans, herons & ibises) Spot-billed pelican takeoff white background.jpg
(Ardeae)
Telluraves
Accipitrimorphae
Cathartiformes (New World vultures)Vintage Vulture Drawing white background.jpg
Accipitriformes (hawks and relatives)Golden Eagle Illustration white background.jpg
Strigiformes (owls)Cuvier-12-Hibou à huppe courte.jpg
Coraciimorphae
Coliiformes (mousebirds) ColiusCastanonotusKeulemans.jpg
Cavitaves
Leptosomiformes (cuckoo roller) Leptosomus discolor - 1825-1834 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - UBA01 IZ16700267.tif
Trogoniformes (trogons and quetzals)Harpactes fasciatus 1838 white background.jpg
Picocoraciae
Bucerotiformes (hornbills and relatives) A monograph of the Bucerotidæ, or family of the hornbills (Plate II) (white background).jpg
Picodynastornithes
Coraciiformes (kingfishers and relatives)Cuvier-46-Martin-pêcheur d'Europe.jpg
Piciformes (woodpeckers and relatives) Dendrocopos major -Durham, England -female-8 white background.jpg
Australaves
Cariamiformes (seriemas)Cariama cristata 1838 white background.jpg
Eufalconimorphae
Falconiformes (falcons)NewZealandFalconBuller white background.jpg
Psittacopasserae
Psittaciformes (parrots)Pyrrhura lucianii - Castelnau 2.jpg
Passeriformes (passerines)Cuvier-33-Moineau domestique.jpg
The classification of birds is a contentious issue. Sibley and Ahlquist's Phylogeny and Classification of Birds (1990) is a landmark work on the classification of birds,[58] although it is frequently debated and constantly revised. Most evidence seems to suggest the assignment of orders is accurate,[59] but scientists disagree about the relationships between the orders themselves; evidence from modern bird anatomy, fossils and DNA have all been brought to bear on the problem, but no strong consensus has emerged. More recently, new fossil and molecular evidence is providing an increasingly clear picture of the evolution of modern bird orders.[52][60]
Genomics
See also: list of sequenced animal genomes
As of 2010, the genome had been sequenced for only two birds, the chicken and the zebra finch. As of 2022 the genomes of 542 species of birds had been completed. At least one genome has been sequenced from every order.[61][62] These include at least one species in about 90% of extant avian families (218 out of 236 families recognised by the Howard and Moore Checklist).[63]
Being able to sequence and compare whole genomes gives researchers many types of information, about genes, the DNA that regulates the genes, and their evolutionary history. This has led to reconsideration of some of the classifications that were based solely on the identification of protein-coding genes. Waterbirds such as pelicans and flamingos, for example, may have in common specific adaptations suited to their environment that were developed independently.[61][62]
Distribution
See also: Lists of birds by region and List of birds by population
small bird withpale belly and breast and patterned wing and head stands on concrete
The range of the house sparrow has expanded dramatically due to human activities.[64]
Birds live and breed in most terrestrial habitats and on all seven continents, reaching their southern extreme in the snow petrel's breeding colonies up to 440 kilometres (270 mi) inland in Antarctica.[65] The highest bird diversity occurs in tropical regions. It was earlier thought that this high diversity was the result of higher speciation rates in the tropics; however recent studies found higher speciation rates in the high latitudes that were offset by greater extinction rates than in the tropics.[66] Many species migrate annually over great distances and across oceans; several families of birds have adapted to life both on the world's oceans and in them, and some seabird species come ashore only to breed,[67] while some penguins have been recorded diving up to 300 metres (980 ft) deep.[68]
Many bird species have established breeding populations in areas to which they have been introduced by humans. Some of these introductions have been deliberate; the ring-necked pheasant, for example, has been introduced around the world as a game bird.[69] Others have been accidental, such as the establishment of wild monk parakeets in several North American cities after their escape from captivity.[70] Some species, including cattle egret,[71] yellow-headed caracara[72] and galah,[73] have spread naturally far beyond their original ranges as agricultural expansion created alternative habitats although modern practices of intensive agriculture have negatively impacted farmland bird populations.[74]
Anatomy and physiology
Main articles: Bird anatomy and Bird vision
See also: Egg tooth
External anatomy of a bird (example: yellow-wattled lapwing): 1 Beak, 2 Head, 3 Iris, 4 Pupil, 5 Mantle, 6 Lesser coverts, 7 Scapulars, 8 Median coverts, 9 Tertials, 10 Rump, 11 Primaries, 12 Vent, 13 Thigh, 14 Tibio-tarsal articulation, 15 Tarsus, 16 Foot, 17 Tibia, 18 Belly, 19 Flanks, 20 Breast, 21 Throat, 22 Wattle, 23 Eyestripe
Compared with other vertebrates, birds have a body plan that shows many unusual adaptations, mostly to facilitate flight.
Skeletal system
Main article: Bird_anatomy § Skeletal_system
The skeleton consists of very lightweight bones. They have large air-filled cavities (called pneumatic cavities) which connect with the respiratory system.[75] The skull bones in adults are fused and do not show cranial sutures.[76] The orbital cavities that house the eyeballs are large and separated from each other by a bony septum (partition). The spine has cervical, thoracic, lumbar and caudal regions with the number of cervical (neck) vertebrae highly variable and especially flexible, but movement is reduced in the anterior thoracic vertebrae and absent in the later vertebrae.[77] The last few are fused with the pelvis to form the synsacrum.[76] The ribs are flattened and the sternum is keeled for the attachment of flight muscles except in the flightless bird orders. The forelimbs are modified into wings.[78] The wings are more or less developed depending on the species; the only known groups that lost their wings are the extinct moa and elephant birds.[79]
Excretory system
Like the reptiles, birds are primarily uricotelic, that is, their kidneys extract nitrogenous waste from their bloodstream and excrete it as uric acid, instead of urea or ammonia, through the ureters into the intestine. Birds do not have a urinary bladder or external urethral opening and (with exception of the ostrich) uric acid is excreted along with faeces as a semisolid waste.[80][81][82] However, birds such as hummingbirds can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia.[83] They also excrete creatine, rather than creatinine like mammals.[76] This material, as well as the output of the intestines, emerges from the bird's cloaca.[84][85] The cloaca is a multi-purpose opening: waste is expelled through it, most birds mate by joining cloaca, and females lay eggs from it. In addition, many species of birds regurgitate pellets.[86]
It is a common but not universal feature of altricial passerine nestlings (born helpless, under constant parental care) that instead of excreting directly into the nest, they produce a fecal sac. This is a mucus-covered pouch that allows parents to either dispose of the waste outside the nest or to recycle the waste through their own digestive system.[87]
Reproductive system
Males within Palaeognathae (with the exception of the kiwis), the Anseriformes (with the exception of screamers), and in rudimentary forms in Galliformes (but fully developed in Cracidae) possess a penis, which is never present in Neoaves.[88][89] The length is thought to be related to sperm competition.[90] When not copulating, it is hidden within the proctodeum compartment within the cloaca, just inside the vent. Female birds have sperm storage tubules[91] that allow sperm to remain viable long after copulation, a hundred days in some species.[92] Sperm from multiple males may compete through this mechanism. Most female birds have a single ovary and a single oviduct, both on the left side,[93] but there are exceptions: species in at least 16 different orders of birds have two ovaries. Even these species, however, tend to have a single oviduct.[93] It has been speculated that this might be an adaptation to flight, but males have two testes, and it is also observed that the gonads in both sexes decrease dramatically in size outside the breeding season.[94][95] Also terrestrial birds generally have a single ovary, as does the platypus, an egg-laying mammal. A more likely explanation is that the egg develops a shell while passing through the oviduct over a period of about a day, so that if two eggs were to develop at the same time, there would be a risk to survival.[93] While rare, mostly abortive, parthenogenesis is not unknown in birds and eggs can be diploid, automictic and results in male offspring.[96]
Birds are solely gonochoric.[97] Meaning they have two sexes: either female or male. The sex of birds is determined by the Z and W sex chromosomes, rather than by the X and Y chromosomes present in mammals. Male birds have two Z chromosomes (ZZ), and female birds have a W chromosome and a Z chromosome (WZ).[76]
In nearly all species of birds, an individual's sex is determined at fertilisation. However, one 2007 study claimed to demonstrate temperature-dependent sex determination among the Australian brushturkey, for which higher temperatures during incubation resulted in a higher female-to-male sex ratio.[98] This, however, was later proven to not be the case. These birds do not exhibit temperature-dependent sex determination, but temperature-dependent sex mortality.[99]
Respiratory and circulatory systems
Birds have one of the most complex respiratory systems of all animal groups.[76] Upon inhalation, 75% of the fresh air bypasses the lungs and flows directly into a posterior air sac which extends from the lungs and connects with air spaces in the bones and fills them with air. The other 25% of the air goes directly into the lungs. When the bird exhales, the used air flows out of the lungs and the stored fresh air from the posterior air sac is simultaneously forced into the lungs. Thus, a bird's lungs receive a constant supply of fresh air during both inhalation and exhalation.[100] Sound production is achieved using the syrinx, a muscular chamber incorporating multiple tympanic membranes which diverges from the lower end of the trachea;[101] the trachea being elongated in some species, increasing the volume of vocalisations and the perception of the bird's size.[102]
In birds, the main arteries taking blood away from the heart originate from the right aortic arch (or pharyngeal arch), unlike in the mammals where the left aortic arch forms this part of the aorta.[76] The postcava receives blood from the limbs via the renal portal system. Unlike in mammals, the circulating red blood cells in birds retain their nucleus.[103]
Heart type and features
Didactic model of an avian heart
The avian circulatory system is driven by a four-chambered, myogenic heart contained in a fibrous pericardial sac. This pericardial sac is filled with a serous fluid for lubrication.[104] The heart itself is divided into a right and left half, each with an atrium and ventricle. The atrium and ventricles of each side are separated by atrioventricular valves which prevent back flow from one chamber to the next during contraction. Being myogenic, the heart's pace is maintained by pacemaker cells found in the sinoatrial node, located on the right atrium.[citation needed]
The sinoatrial node uses calcium to cause a depolarising signal transduction pathway from the atrium through right and left atrioventricular bundle which communicates contraction to the ventricles. The avian heart also consists of muscular arches that are made up of thick bundles of muscular layers. Much like a mammalian heart, the avian heart is composed of endocardial, myocardial and epicardial layers.[104] The atrium walls tend to be thinner than the ventricle walls, due to the intense ventricular contraction used to pump oxygenated blood throughout the body. Avian hearts are generally larger than mammalian hearts when compared to body mass. This adaptation allows more blood to be pumped to meet the high metabolic need associated with flight.[105]
Organisation
Birds have a very efficient system for diffusing oxygen into the blood; birds have a ten times greater surface area to gas exchange volume than mammals. As a result, birds have more blood in their capillaries per unit of volume of lung than a mammal.[105] The arteries are composed of thick elastic muscles to withstand the pressure of the ventricular contractions, and become more rigid as they move away from the heart. Blood moves through the arteries, which undergo vasoconstriction, and into arterioles which act as a transportation system to distribute primarily oxygen as well as nutrients to all tissues of the body.[106] As the arterioles move away from the heart and into individual organs and tissues they are further divided to increase surface area and slow blood flow. Blood travels through the arterioles and moves into the capillaries where gas exchange can occur.[citation needed]
Capillaries are organised into capillary beds in tissues; it is here that blood exchanges oxygen for carbon dioxide waste. In the capillary beds, blood flow is slowed to allow maximum diffusion of oxygen into the tissues. Once the blood has become deoxygenated, it travels through venules then veins and back to the heart. Veins, unlike arteries, are thin and rigid as they do not need to withstand extreme pressure. As blood travels through the venules to the veins a funneling occurs called vasodilation bringing blood back to the heart.[106] Once the blood reaches the heart, it moves first into the right atrium, then the right ventricle to be pumped through the lungs for further gas exchange of carbon dioxide waste for oxygen. Oxygenated blood then flows from the lungs through the left atrium to the left ventricle where it is pumped out to the body.[citation needed]
The nictitating membrane as it covers the eye of a masked lapwing
Nervous system
The nervous system is large relative to the bird's size.[76] The most developed part of the brain is the one that controls the flight-related functions, while the cerebellum coordinates movement and the cerebrum controls behaviour patterns, navigation, mating and nest building. Most birds have a poor sense of smell[107] with notable exceptions including kiwis,[108] New World vultures[109] and tubenoses.[110] The avian visual system is usually highly developed. Water birds have special flexible lenses, allowing accommodation for vision in air and water.[76] Some species also have dual fovea. Birds are tetrachromatic, possessing ultraviolet (UV) sensitive cone cells in the eye as well as green, red and blue ones.[111] They also have double cones, likely to mediate achromatic vision.[112]
Many birds show plumage patterns in ultraviolet that are invisible to the human eye; some birds whose sexes appear similar to the naked eye are distinguished by the presence of ultraviolet reflective patches on their feathers. Male blue tits have an ultraviolet reflective crown patch which is displayed in courtship by posturing and raising of their nape feathers.[113] Ultraviolet light is also used in foraging—kestrels have been shown to search for prey by detecting the UV reflective urine trail marks left on the ground by rodents.[114] With the exception of pigeons and a few other species,[115] the eyelids of birds are not used in blinking. Instead the eye is lubricated by the nictitating membrane, a third eyelid that moves horizontally.[116] The nictitating membrane also covers the eye and acts as a contact lens in many aquatic birds.[76] The bird retina has a fan shaped blood supply system called the pecten.[76]
Eyes of most birds are large, not very round and capable of only limited movement in the orbits,[76] typically 10–20°.[117] Birds with eyes on the sides of their heads have a wide visual field, while birds with eyes on the front of their heads, such as owls, have binocular vision and can estimate the depth of field.[117][118] The avian ear lacks external pinnae but is covered by feathers, although in some birds, such as the Asio, Bubo and Otus owls, these feathers form tufts which resemble ears. The inner ear has a cochlea, but it is not spiral as in mammals.[119]
Defence and intraspecific combat
A few species are able to use chemical defences against predators; some Procellariiformes can eject an unpleasant stomach oil against an aggressor,[120] and some species of pitohuis from New Guinea have a powerful neurotoxin in their skin and feathers.[121]
A lack of field observations limit our knowledge, but intraspecific conflicts are known to sometimes result in injury or death.[122] The screamers (Anhimidae), some jacanas (Jacana, Hydrophasianus), the spur-winged goose (Plectropterus), the torrent duck (Merganetta) and nine species of lapwing (Vanellus) use a sharp spur on the wing as a weapon. The steamer ducks (Tachyeres), geese and swans (Anserinae), the solitaire (Pezophaps), sheathbills (Chionis), some guans (Crax) and stone curlews (Burhinus) use a bony knob on the alular metacarpal to punch and hammer opponents.[122] The jacanas Actophilornis and Irediparra have an expanded, blade-like radius. The extinct Xenicibis was unique in having an elongate forelimb and massive hand which likely functioned in combat or defence as a jointed club or flail. Swans, for instance, may strike with the bony spurs and bite when defending eggs or young.[122]
Feathers, plumage, and scales
Main articles: Feather, Flight feather, and Down feather
Owl with eyes closed in front of similarly coloured tree trunk partly obscured by green leaves
The disruptively patterned plumage of the African scops owl allows it to blend in with its surroundings.
Feathers are a feature characteristic of birds (though also present in some dinosaurs not currently considered to be true birds). They facilitate flight, provide insulation that aids in thermoregulation, and are used in display, camouflage, and signalling.[76] There are several types of feathers, each serving its own set of purposes. Feathers are epidermal growths attached to the skin and arise only in specific tracts of skin called pterylae. The distribution pattern of these feather tracts (pterylosis) is used in taxonomy and systematics. The arrangement and appearance of feathers on the body, called plumage, may vary within species by age, social status,[123] and sex.[124]
Plumage is regularly moulted; the standard plumage of a bird that has moulted after breeding is known as the "non-breeding" plumage, or—in the Humphrey–Parkes terminology—"basic" plumage; breeding plumages or variations of the basic plumage are known under the Humphrey–Parkes system as "alternate" plumages.[125] Moulting is annual in most species, although some may have two moults a year, and large birds of prey may moult only once every few years. Moulting patterns vary across species. In passerines, flight feathers are replaced one at a time with the innermost primary being the first. When the fifth of sixth primary is replaced, the outermost tertiaries begin to drop. After the innermost tertiaries are moulted, the secondaries starting from the innermost begin to drop and this proceeds to the outer feathers (centrifugal moult). The greater primary coverts are moulted in synchrony with the primary that they overlap.[126]
A small number of species, such as ducks and geese, lose all of their flight feathers at once, temporarily becoming flightless.[127] As a general rule, the tail feathers are moulted and replaced starting with the innermost pair.[126] Centripetal moults of tail feathers are however seen in the Phasianidae.[128] The centrifugal moult is modified in the tail feathers of woodpeckers and treecreepers, in that it begins with the second innermost pair of feathers and finishes with the central pair of feathers so that the bird maintains a functional climbing tail.[126][129] The general pattern seen in passerines is that the primaries are replaced outward, secondaries inward, and the tail from centre outward.[130] Before nesting, the females of most bird species gain a bare brood patch by losing feathers close to the belly. The skin there is well supplied with blood vessels and helps the bird in incubation.[131]
Red parrot with yellow bill and wing feathers in bill
Red lory preening
Feathers require maintenance and birds preen or groom them daily, spending an average of around 9% of their daily time on this.[132] The bill is used to brush away foreign particles and to apply waxy secretions from the uropygial gland; these secretions protect the feathers' flexibility and act as an antimicrobial agent, inhibiting the growth of feather-degrading bacteria.[133] This may be supplemented with the secretions of formic acid from ants, which birds receive through a behaviour known as anting, to remove feather parasites.[134]
The scales of birds are composed of the same keratin as beaks, claws, and spurs. They are found mainly on the toes and metatarsus, but may be found further up on the ankle in some birds. Most bird scales do not overlap significantly, except in the cases of kingfishers and woodpeckers. The scales of birds are thought to be homologous to those of reptiles and mammals.[135]
Flight
Main articles: Bird flight and Flightless birds
Black bird with white chest in flight with wings facing down and tail fanned and down pointing
Restless flycatcher in the downstroke of flapping flight
Most birds can fly, which distinguishes them from almost all other vertebrate classes. Flight is the primary means of locomotion for most bird species and is used for searching for food and for escaping from predators. Birds have various adaptations for flight, including a lightweight skeleton, two large flight muscles, the pectoralis (which accounts for 15% of the total mass of the bird) and the supracoracoideus, as well as a modified forelimb (wing) that serves as an aerofoil.[76]
Wing shape and size generally determine a bird's flight style and performance; many birds combine powered, flapping flight with less energy-intensive soaring flight. About 60 extant bird species are flightless, as were many extinct birds.[136] Flightlessness often arises in birds on isolated islands, most likely due to limited resources and the absence of mammalian land predators.[137] Flightlessnes is almost exclusively correlated with gigantism due to an island's inheren condition of isolation.[138] Although flightless, penguins use similar musculature and movements to "fly" through the water, as do some flight-capable birds such as auks, shearwaters and dippers.[139]
Behaviour
Most birds are diurnal, but some birds, such as many species of owls and nightjars, are nocturnal or crepuscular (active during twilight hours), and many coastal waders feed when the tides are appropriate, by day or night.[140]
Diet and feeding
Illustration of the heads of 16 types of birds with different shapes and sizes of beak
Feeding adaptations in beaks
Birds' diets are varied and often include nectar, fruit, plants, seeds, carrion, and various small animals, including other birds.[76] The digestive system of birds is unique, with a crop for storage and a gizzard that contains swallowed stones for grinding food to compensate for the lack of teeth.[141] Some species such as pigeons and some psittacine species do not have a gallbladder.[142] Most birds are highly adapted for rapid digestion to aid with flight.[143] Some migratory birds have adapted to use protein stored in many parts of their bodies, including protein from the intestines, as additional energy during migration.[144]
Birds that employ many strategies to obtain food or feed on a variety of food items are called generalists, while others that concentrate time and effort on specific food items or have a single strategy to obtain food are considered specialists.[76] Avian foraging strategies can vary widely by species. Many birds glean for insects, invertebrates, fruit, or seeds. Some hunt insects by suddenly attacking from a branch. Those species that seek pest insects are considered beneficial 'biological control agents' and their presence encouraged in biological pest control programmes.[145] Combined, insectivorous birds eat 400–500 million metric tons of arthropods annually.[146]
Nectar feeders such as hummingbirds, sunbirds, lories, and lorikeets amongst others have specially adapted brushy tongues and in many cases bills designed to fit co-adapted flowers.[147] Kiwis and shorebirds with long bills probe for invertebrates; shorebirds' varied bill lengths and feeding methods result in the separation of ecological niches.[76][148] Loons, diving ducks, penguins and auks pursue their prey underwater, using their wings or feet for propulsion,[67] while aerial predators such as sulids, kingfishers and terns plunge dive after their prey. Flamingos, three species of prion, and some ducks are filter feeders.[149][150] Geese and dabbling ducks are primarily grazers.[151][152]
Some species, including frigatebirds, gulls,[153] and skuas,[154] engage in kleptoparasitism, stealing food items from other birds. Kleptoparasitism is thought to be a supplement to food obtained by hunting, rather than a significant part of any species' diet; a study of great frigatebirds stealing from masked boobies estimated that the frigatebirds stole at most 40% of their food and on average stole only 5%.[155] Other birds are scavengers; some of these, like vultures, are specialised carrion eaters, while others, like gulls, corvids, or other birds of prey, are opportunists.[156]
Water and drinking
Water is needed by many birds although their mode of excretion and lack of sweat glands reduces the physiological demands.[157] Some desert birds can obtain their water needs entirely from moisture in their food. They may also have other adaptations such as allowing their body temperature to rise, saving on moisture loss from evaporative cooling or panting.[158] Seabirds can drink seawater and have salt glands inside the head that eliminate excess salt out of the nostrils.[159]
Most birds scoop water in their beaks and raise their head to let water run down the throat. Some species, especially of arid zones, belonging to the pigeon, finch, mousebird, button-quail and bustard families are capable of sucking up water without the need to tilt back their heads.[160] Some desert birds depend on water sources and sandgrouse are particularly well known for their daily congregations at waterholes. Nesting sandgrouse and many plovers carry water to their young by wetting their belly feathers.[161] Some birds carry water for chicks at the nest in their crop or regurgitate it along with food. The pigeon family, flamingos and penguins have adaptations to produce a nutritive fluid called crop milk that they provide to their chicks.[162]
Feather care
Main article: Preening
Feathers, being critical to the survival of a bird, require maintenance. Apart from physical wear and tear, feathers face the onslaught of fungi, ectoparasitic feather mites and bird lice.[163] The physical condition of feathers are maintained by preening often with the application of secretions from the preen gland. Birds also bathe in water or dust themselves. While some birds dip into shallow water, more aerial species may make aerial dips into water and arboreal species often make use of dew or rain that collect on leaves. Birds of arid regions make use of loose soil to dust-bathe. A behaviour termed as anting in which the bird encourages ants to run through their plumage is also thought to help them reduce the ectoparasite load in feathers. Many species will spread out their wings and expose them to direct sunlight and this too is thought to help in reducing fungal and ectoparasitic activity that may lead to feather damage.[164][165]
Migration
Main article: Bird migration
A flock of Canada geese in V formation
Many bird species migrate to take advantage of global differences of seasonal temperatures, therefore optimising availability of food sources and breeding habitat. These migrations vary among the different groups. Many landbirds, shorebirds, and waterbirds undertake annual long-distance migrations, usually triggered by the length of daylight as well as weather conditions. These birds are characterised by a breeding season spent in the temperate or polar regions and a non-breeding season in the tropical regions or opposite hemisphere. Before migration, birds substantially increase body fats and reserves and reduce the size of some of their organs.[166][167]
Migration is highly demanding energetically, particularly as birds need to cross deserts and oceans without refuelling. Landbirds have a flight range of around 2,500 km (1,600 mi) and shorebirds can fly up to 4,000 km (2,500 mi),[76] although the bar-tailed godwit is capable of non-stop flights of up to 10,200 km (6,300 mi).[168] Seabirds also undertake long migrations, the longest annual migration being those of sooty shearwaters, which nest in New Zealand and Chile and spend the northern summer feeding in the North Pacific off Japan, Alaska and California, an annual round trip of 64,000 km (39,800 mi).[169] Other seabirds disperse after breeding, travelling widely but having no set migration route. Albatrosses nesting in the Southern Ocean often undertake circumpolar trips between breeding seasons.[170]
A map of the Pacific Ocean with several coloured lines representing bird routes running from New Zealand to Korea
The routes of satellite-tagged bar-tailed godwits migrating north from New Zealand. This species has the longest known non-stop migration of any species, up to 10,200 km (6,300 mi).
Some bird species undertake shorter migrations, travelling only as far as is required to avoid bad weather or obtain food. Irruptive species such as the boreal finches are one such group and can commonly be found at a location in one year and absent the next. This type of migration is normally associated with food availability.[171] Species may also travel shorter distances over part of their range, with individuals from higher latitudes travelling into the existing range of conspecifics; others undertake partial migrations, where only a fraction of the population, usually females and subdominant males, migrates.[172] Partial migration can form a large percentage of the migration behaviour of birds in some regions; in Australia, surveys found that 44% of non-passerine birds and 32% of passerines were partially migratory.[173]
Altitudinal migration is a form of short-distance migration in which birds spend the breeding season at higher altitudes and move to lower ones during suboptimal conditions. It is most often triggered by temperature changes and usually occurs when the normal territories also become inhospitable due to lack of food.[174] Some species may also be nomadic, holding no fixed territory and moving according to weather and food availability. Parrots as a family are overwhelmingly neither migratory nor sedentary but considered to either be dispersive, irruptive, nomadic or undertake small and irregular migrations.[175]
The ability of birds to return to precise locations across vast distances has been known for some time; in an experiment conducted in the 1950s, a Manx shearwater released in Boston in the United States returned to its colony in Skomer, in Wales within 13 days, a distance of 5,150 km (3,200 mi).[176] Birds navigate during migration using a variety of methods. For diurnal migrants, the sun is used to navigate by day, and a stellar compass is used at night. Birds that use the sun compensate for the changing position of the sun during the day by the use of an internal clock.[76] Orientation with the stellar compass depends on the position of the constellations surrounding Polaris.[177] These are backed up in some species by their ability to sense the Earth's geomagnetism through specialised photoreceptors.[178]
Communication
See also: Bird vocalisation
Bird song
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Song of the house wren, a common North American songbird
Mimicry
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A tooth-billed bowerbird mimicking a spangled drongo
Drumming
0:03
A woodpecker drumming on wood
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Large brown patterned ground bird with outstretched wings each with a large spot in the centre
The startling display of the sunbittern mimics a large predator.
Birds communicate primarily using visual and auditory signals. Signals can be interspecific (between species) and intraspecific (within species).
Birds sometimes use plumage to assess and assert social dominance,[179] to display breeding condition in sexually selected species, or to make threatening displays, as in the sunbittern's mimicry of a large predator to ward off hawks and protect young chicks.[180]
Visual communication among birds may also involve ritualised displays, which have developed from non-signalling actions such as preening, the adjustments of feather position, pecking, or other behaviour. These displays may signal aggression or submission or may contribute to the formation of pair-bonds.[76] The most elaborate displays occur during courtship, where "dances" are often formed from complex combinations of many possible component movements;[181] males' breeding success may depend on the quality of such displays.[182]
Bird calls and songs, which are produced in the syrinx, are the major means by which birds communicate with sound. This communication can be very complex; some species can operate the two sides of the syrinx independently, allowing the simultaneous production of two different songs.[101] Calls are used for a variety of purposes, including mate attraction,[76] evaluation of potential mates,[183] bond formation, the claiming and maintenance of territories,[76] the identification of other individuals (such as when parents look for chicks in colonies or when mates reunite at the start of breeding season),[184] and the warning of other birds of potential predators, sometimes with specific information about the nature of the threat.[185] Some birds also use mechanical sounds for auditory communication. The Coenocorypha snipes of New Zealand drive air through their feathers,[186] woodpeckers drum for long-distance communication,[187] and palm cockatoos use tools to drum.[188]
Flocking and other associations
massive flock of tiny birds seen from distance so that birds appear as specks
Red-billed queleas, the most numerous species of wild bird,[189] form enormous flocks – sometimes tens of thousands strong.
While some birds are essentially territorial or live in small family groups, other birds may form large flocks. The principal benefits of flocking are safety in numbers and increased foraging efficiency.[76] Defence against predators is particularly important in closed habitats like forests, where ambush predation is common and multiple eyes can provide a valuable early warning system. This has led to the development of many mixed-species feeding flocks, which are usually composed of small numbers of many species; these flocks provide safety in numbers but increase potential competition for resources.[190] Costs of flocking include bullying of socially subordinate birds by more dominant birds and the reduction of feeding efficiency in certain cases.[191]
Birds sometimes also form associations with non-avian species. Plunge-diving seabirds associate with dolphins and tuna, which push shoaling fish towards the surface.[192] Some species of hornbills have a mutualistic relationship with dwarf mongooses, in which they forage together and warn each other of nearby birds of prey and other predators.[193]
Resting and roosting
"Roosting" redirects here. For other uses, see Roost.
Pink flamingo with grey legs and long neck pressed against body and head tucked under wings
Many birds, like this American flamingo, tuck their head into their back when sleeping.
The high metabolic rates of birds during the active part of the day is supplemented by rest at other times. Sleeping birds often use a type of sleep known as vigilant sleep, where periods of rest are interspersed with quick eye-opening "peeks", allowing them to be sensitive to disturbances and enable rapid escape from threats.[194] Swifts are believed to be able to sleep in flight and radar observations suggest that they orient themselves to face the wind in their roosting flight.[195] It has been suggested that there may be certain kinds of sleep which are possible even when in flight.[196]
Some birds have also demonstrated the capacity to fall into slow-wave sleep one hemisphere of the brain at a time. The birds tend to exercise this ability depending upon its position relative to the outside of the flock. This may allow the eye opposite the sleeping hemisphere to remain vigilant for predators by viewing the outer margins of the flock. This adaptation is also known from marine mammals.[197] Communal roosting is common because it lowers the loss of body heat and decreases the risks associated with predators.[198] Roosting sites are often chosen with regard to thermoregulation and safety.[199] Unusual mobile roost sites include large herbivores on the African savanna that are used by oxpeckers.[200]
Many sleeping birds bend their heads over their backs and tuck their bills in their back feathers, although others place their beaks among their breast feathers. Many birds rest on one leg, while some may pull up their legs into their feathers, especially in cold weather. Perching birds have a tendon-locking mechanism that helps them hold on to the perch when they are asleep. Many ground birds, such as quails and pheasants, roost in trees. A few parrots of the genus Loriculus roost hanging upside down.[201] Some hummingbirds go into a nightly state of torpor accompanied with a reduction of their metabolic rates.[202] This physiological adaptation shows in nearly a hundred other species, including owlet-nightjars, nightjars, and woodswallows. One species, the common poorwill, even enters a state of hibernation.[203] Birds do not have sweat glands, but can lose water directly through the skin, and they may cool themselves by moving to shade, standing in water, panting, increasing their surface area, fluttering their throat or using special behaviours like urohidrosis to cool themselves.[204][205]
Breeding
See also: Category:Avian sexuality, Animal sexual behaviour § Birds, Seabird breeding behaviour, and Sexual selection in birds
Social systems
Bird faces up with green face, black breast and pink lower body. Elaborate long feathers on the wings and tail.
Like others of its family, the male Raggiana bird-of-paradise has elaborate breeding plumage used to impress females.[206]
Ninety-five per cent of bird species are socially monogamous. These species pair for at least the length of the breeding season or—in some cases—for several years or until the death of one mate.[207] Monogamy allows for both paternal care and biparental care, which is especially important for species in which care from both the female and the male parent is required in order to successfully rear a brood.[208] Among many socially monogamous species, extra-pair copulation (infidelity) is common.[209] Such behaviour typically occurs between dominant males and females paired with subordinate males, but may also be the result of forced copulation in ducks and other anatids.[210]
For females, possible benefits of extra-pair copulation include getting better genes for her offspring and insuring against the possibility of infertility in her mate.[211] Males of species that engage in extra-pair copulations will closely guard their mates to ensure the parentage of the offspring that they raise.[212]
Other mating systems, including polygyny, polyandry, polygamy, polygynandry, and promiscuity, also occur.[76] Polygamous breeding systems arise when females are able to raise broods without the help of males.[76] Mating systems vary across bird families[213] but variations within species are thought to be driven by environmental conditions.[214]
Breeding usually involves some form of courtship display, typically performed by the male.[215] Most displays are rather simple and involve some type of song. Some displays, however, are quite elaborate. Depending on the species, these may include wing or tail drumming, dancing, aerial flights, or communal lekking. Females are generally the ones that drive partner selection,[216] although in the polyandrous phalaropes, this is reversed: plainer males choose brightly coloured females.[217] Courtship feeding, billing and allopreening are commonly performed between partners, generally after the birds have paired and mated.[218]
Homosexual behaviour has been observed in males or females in numerous species of birds, including copulation, pair-bonding, and joint parenting of chicks.[219] Over 130 avian species around the world engage in sexual interactions between the same sex or homosexual behaviours. "Same-sex courtship activities may involve elaborate displays, synchronized dances, gift-giving ceremonies, or behaviors at specific display areas including bowers, arenas, or leks."[220]
Territories, nesting and incubation
See also: Bird nest
Many birds actively defend a territory from others of the same species during the breeding season; maintenance of territories protects the food source for their chicks. Species that are unable to defend feeding territories, such as seabirds and swifts, often breed in colonies instead; this is thought to offer protection from predators. Colonial breeders defend small nesting sites, and competition between and within species for nesting sites can be intense.[221]
All birds lay amniotic eggs with hard shells made mostly of calcium carbonate.[76] Hole and burrow nesting species tend to lay white or pale eggs, while open nesters lay camouflaged eggs. There are many exceptions to this pattern, however; the ground-nesting nightjars have pale eggs, and camouflage is instead provided by their plumage. Species that are victims of brood parasites have varying egg colours to improve the chances of spotting a parasite's egg, which forces female parasites to match their eggs to those of their hosts.[222]
Yellow weaver (bird) with black head hangs an upside-down nest woven out of grass fronds.
Male golden-backed weavers construct elaborate suspended nests out of grass.
Bird eggs are usually laid in a nest. Most species create somewhat elaborate nests, which can be cups, domes, plates, mounds, or burrows.[223] Some bird nests can be a simple scrape, with minimal or no lining; most seabird and wader nests are no more than a scrape on the ground. Most birds build nests in sheltered, hidden areas to avoid predation, but large or colonial birds—which are more capable of defence—may build more open nests. During nest construction, some species seek out plant matter from plants with parasite-reducing toxins to improve chick survival,[224] and feathers are often used for nest insulation.[223] Some bird species have no nests; the cliff-nesting common guillemot lays its eggs on bare rock, and male emperor penguins keep eggs between their body and feet. The absence of nests is especially prevalent in open habitat ground-nesting species where any addition of nest material would make the nest more conspicuous. Many ground nesting birds lay a clutch of eggs that hatch synchronously, with precocial chicks led away from the nests (nidifugous) by their parents soon after hatching.[225]
Nest made of straw with five white eggs and one grey speckled egg
Nest of an eastern phoebe that has been parasitised by a brown-headed cowbird
Incubation, which regulates temperature for chick development, usually begins after the last egg has been laid.[76] In monogamous species incubation duties are often shared, whereas in polygamous species one parent is wholly responsible for incubation. Warmth from parents passes to the eggs through brood patches, areas of bare skin on the abdomen or breast of the incubating birds. Incubation can be an energetically demanding process; adult albatrosses, for instance, lose as much as 83 grams (2.9 oz) of body weight per day of incubation.[226] The warmth for the incubation of the eggs of megapodes comes from the sun, decaying vegetation or volcanic sources.[227] Incubation periods range from 10 days (in woodpeckers, cuckoos and passerine birds) to over 80 days (in albatrosses and kiwis).[76]
The diversity of characteristics of birds is great, sometimes even in closely related species. Several avian characteristics are compared in the table below.[228][229]
Species Adult weight
(grams) Incubation
(days) Clutches
(per year) Clutch size
Ruby-throated hummingbird (Archilochus colubris) 3 13 2.0 2
House sparrow (Passer domesticus) 25 11 4.5 5
Greater roadrunner (Geococcyx californianus) 376 20 1.5 4
Turkey vulture (Cathartes aura) 2,200 39 1.0 2
Laysan albatross (Diomedea immutabilis) 3,150 64 1.0 1
Magellanic penguin (Spheniscus magellanicus) 4,000 40 1.0 1
Golden eagle (Aquila chrysaetos) 4,800 40 1.0 2
Wild turkey (Meleagris gallopavo) 6,050 28 1.0 11
Parental care and fledging
Main article: Parental care in birds
At the time of their hatching, chicks range in development from helpless to independent, depending on their species. Helpless chicks are termed altricial, and tend to be born small, blind, immobile and naked; chicks that are mobile and feathered upon hatching are termed precocial. Altricial chicks need help thermoregulating and must be brooded for longer than precocial chicks. The young of many bird species do not precisely fit into either the precocial or altricial category, having some aspects of each and thus fall somewhere on an "altricial-precocial spectrum".[230] Chicks at neither extreme but favouring one or the other may be termed semi-precocial[231] or semi-altricial.[232]
Hummingbird perched on edge of tiny nest places food into mouth of one of two chicks
A female calliope hummingbird feeding fully grown chicks
Looking down on three helpless blind chicks in a nest within the hollow of a dead tree trunk
Altricial chicks of a white-breasted woodswallow
The length and nature of parental care varies widely amongst different orders and species. At one extreme, parental care in megapodes ends at hatching; the newly hatched chick digs itself out of the nest mound without parental assistance and can fend for itself immediately.[233] At the other extreme, many seabirds have extended periods of parental care, the longest being that of the great frigatebird, whose chicks take up to six months to fledge and are fed by the parents for up to an additional 14 months.[234] The chick guard stage describes the period of breeding during which one of the adult birds is permanently present at the nest after chicks have hatched. The main purpose of the guard stage is to aid offspring to thermoregulate and protect them from predation.[235]
In some species, both parents care for nestlings and fledglings; in others, such care is the responsibility of only one sex. In some species, other members of the same species—usually close relatives of the breeding pair, such as offspring from previous broods—will help with the raising of the young.[236] Such alloparenting is particularly common among the Corvida, which includes such birds as the true crows, Australian magpie and fairy-wrens,[237] but has been observed in species as different as the rifleman and red kite. Among most groups of animals, male parental care is rare. In birds, however, it is quite common—more so than in any other vertebrate class.[76] Although territory and nest site defence, incubation, and chick feeding are often shared tasks, there is sometimes a division of labour in which one mate undertakes all or most of a particular duty.[238]
The point at which chicks fledge varies dramatically. The chicks of the Synthliboramphus murrelets, like the ancient murrelet, leave the nest the night after they hatch, following their parents out to sea, where they are raised away from terrestrial predators.[239] Some other species, such as ducks, move their chicks away from the nest at an early age. In most species, chicks leave the nest just before, or soon after, they are able to fly. The amount of parental care after fledging varies; albatross chicks leave the nest on their own and receive no further help, while other species continue some supplementary feeding after fledging.[240] Chicks may also follow their parents during their first migration.[241]
Brood parasites
Main article: Brood parasite
Small brown bird places an insect in the bill of much larger grey bird in nest
Reed warbler raising a common cuckoo, a brood parasite
Brood parasitism, in which an egg-layer leaves her eggs with another individual's brood, is more common among birds than any other type of organism.[242] After a parasitic bird lays her eggs in another bird's nest, they are often accepted and raised by the host at the expense of the host's own brood. Brood parasites may be either obligate brood parasites, which must lay their eggs in the nests of other species because they are incapable of raising their own young, or non-obligate brood parasites, which sometimes lay eggs in the nests of conspecifics to increase their reproductive output even though they could have raised their own young.[243] One hundred bird species, including honeyguides, icterids, and ducks, are obligate parasites, though the most famous are the cuckoos.[242] Some brood parasites are adapted to hatch before their host's young, which allows them to destroy the host's eggs by pushing them out of the nest or to kill the host's chicks; this ensures that all food brought to the nest will be fed to the parasitic chicks.[244]
Sexual selection
The peacock tail in flight, the classic example of a Fisherian runaway
Main article: Sexual selection in birds
Birds have evolved a variety of mating behaviours, with the peacock tail being perhaps the most famous example of sexual selection and the Fisherian runaway. Commonly occurring sexual dimorphisms such as size and colour differences are energetically costly attributes that signal competitive breeding situations.[245] Many types of avian sexual selection have been identified; intersexual selection, also known as female choice; and intrasexual competition, where individuals of the more abundant sex compete with each other for the privilege to mate. Sexually selected traits often evolve to become more pronounced in competitive breeding situations until the trait begins to limit the individual's fitness. Conflicts between an individual fitness and signalling adaptations ensure that sexually selected ornaments such as plumage colouration and courtship behaviour are "honest" traits. Signals must be costly to ensure that only good-quality individuals can present these exaggerated sexual ornaments and behaviours.[246]
Inbreeding depression
Main article: Inbreeding depression
Inbreeding causes early death (inbreeding depression) in the zebra finch Taeniopygia guttata.[247] Embryo survival (that is, hatching success of fertile eggs) was significantly lower for sib-sib mating pairs than for unrelated pairs.[citation needed]
Darwin's finch Geospiza scandens experiences inbreeding depression (reduced survival of offspring) and the magnitude of this effect is influenced by environmental conditions such as low food availability.[248]
Inbreeding avoidance
Main article: Inbreeding avoidance
Incestuous matings by the purple-crowned fairy wren Malurus coronatus result in severe fitness costs due to inbreeding depression (greater than 30% reduction in hatchability of eggs).[249] Females paired with related males may undertake extra pair matings (see Promiscuity#Other animals for 90% frequency in avian species) that can reduce the negative effects of inbreeding. However, there are ecological and demographic constraints on extra pair matings. Nevertheless, 43% of broods produced by incestuously paired females contained extra pair young.[249]
Inbreeding depression occurs in the great tit (Parus major) when the offspring produced as a result of a mating between close relatives show reduced fitness. In natural populations of Parus major, inbreeding is avoided by dispersal of individuals from their birthplace, which reduces the chance of mating with a close relative.[250]
Southern pied babblers Turdoides bicolor appear to avoid inbreeding in two ways. The first is through dispersal, and the second is by avoiding familiar group members as mates.[251]
Cooperative breeding in birds typically occurs when offspring, usually males, delay dispersal from their natal group in order to remain with the family to help rear younger kin.[252] Female offspring rarely stay at home, dispersing over distances that allow them to breed independently, or to join unrelated groups. In general, inbreeding is avoided because it leads to a reduction in progeny fitness (inbreeding depression) due largely to the homozygous expression of deleterious recessive alleles.[253] Cross-fertilisation between unrelated individuals ordinarily leads to the masking of deleterious recessive alleles in progeny.[254][255]
Ecology
Gran Canaria blue chaffinch, an example of a bird highly specialised in its habitat, in this case in the Canarian pine forests
Birds occupy a wide range of ecological positions.[189] While some birds are generalists, others are highly specialised in their habitat or food requirements. Even within a single habitat, such as a forest, the niches occupied by different species of birds vary, with some species feeding in the forest canopy, others beneath the canopy, and still others on the forest floor. Forest birds may be insectivores, frugivores, or nectarivores. Aquatic birds generally feed by fishing, plant eating, and piracy or kleptoparasitism. Many grassland birds are granivores. Birds of prey specialise in hunting mammals or other birds, while vultures are specialised scavengers. Birds are also preyed upon by a range of mammals including a few avivorous bats.[256] A wide range of endo- and ectoparasites depend on birds and some parasites that are transmitted from parent to young have co-evolved and show host-specificity.[257][258]
Some nectar-feeding birds are important pollinators, and many frugivores play a key role in seed dispersal.[259] Plants and pollinating birds often coevolve,[260] and in some cases a flower's primary pollinator is the only species capable of reaching its nectar.[261]
Birds are often important to island ecology. Birds have frequently reached islands that mammals have not; on those islands, birds may fulfil ecological roles typically played by larger animals. For example, in New Zealand nine species of moa were important browsers, as are the kererū and kokako today.[259] Today the plants of New Zealand retain the defensive adaptations evolved to protect them from the extinct moa.[262]
Many birds act as ecosystem engineers through the construction of nests, which provide important microhabitats and food for hundreds of species of invertebrates.[263][264] Nesting seabirds may affect the ecology of islands and surrounding seas, principally through the concentration of large quantities of guano, which may enrich the local soil[265] and the surrounding seas.[266]
A wide variety of avian ecology field methods, including counts, nest monitoring, and capturing and marking, are used for researching avian ecology.[267]
Relationship with humans
Main article: Birds in culture
Two rows of cages in a dark barn with many white chickens in each cage
Industrial farming of chickens
Since birds are highly visible and common animals, humans have had a relationship with them since the dawn of man.[268] Sometimes, these relationships are mutualistic, like the cooperative honey-gathering among honeyguides and African peoples such as the Borana.[269] Other times, they may be commensal, as when species such as the house sparrow[270] have benefited from human activities. Several bird species have become commercially significant agricultural pests,[271] and some pose an aviation hazard.[272] Human activities can also be detrimental, and have threatened numerous bird species with extinction (hunting, avian lead poisoning, pesticides, roadkill, wind turbine kills[273] and predation by pet cats and dogs are common causes of death for birds).[274]
Birds can act as vectors for spreading diseases such as psittacosis, salmonellosis, campylobacteriosis, mycobacteriosis (avian tuberculosis), avian influenza (bird flu), giardiasis, and cryptosporidiosis over long distances. Some of these are zoonotic diseases that can also be transmitted to humans.[275]
Economic importance
See also: Pet § Birds
Illustration of fisherman on raft with pole for punting and numerous black birds on raft
The use of cormorants by Asian fishermen is in steep decline but survives in some areas as a tourist attraction.
Domesticated birds raised for meat and eggs, called poultry, are the largest source of animal protein eaten by humans; in 2003, 76 million tons of poultry and 61 million tons of eggs were produced worldwide.[276] Chickens account for much of human poultry consumption, though domesticated turkeys, ducks, and geese are also relatively common.[citation needed] Many species of birds are also hunted for meat. Bird hunting is primarily a recreational activity except in extremely undeveloped areas. The most important birds hunted in North and South America are waterfowl; other widely hunted birds include pheasants, wild turkeys, quail, doves, partridge, grouse, snipe, and woodcock.[citation needed] Muttonbirding is also popular in Australia and New Zealand.[277] Although some hunting, such as that of muttonbirds, may be sustainable, hunting has led to the extinction or endangerment of dozens of species.[278]
Other commercially valuable products from birds include feathers (especially the down of geese and ducks), which are used as insulation in clothing and bedding, and seabird faeces (guano), which is a valuable source of phosphorus and nitrogen. The War of the Pacific, sometimes called the Guano War, was fought in part over the control of guano deposits.[279]
Birds have been domesticated by humans both as pets and for practical purposes. Colourful birds, such as parrots and mynas, are bred in captivity or kept as pets, a practice that has led to the illegal trafficking of some endangered species.[280] Falcons and cormorants have long been used for hunting and fishing, respectively. Messenger pigeons, used since at least 1 AD, remained important as recently as World War II. Today, such activities are more common either as hobbies, for entertainment and tourism,[281]
Amateur bird enthusiasts (called birdwatchers, twitchers or, more commonly, birders) number in the millions.[282] Many homeowners erect bird feeders near their homes to attract various species. Bird feeding has grown into a multimillion-dollar industry; for example, an estimated 75% of households in Britain provide food for birds at some point during the winter.[283]
In religion and mythology
Woodcut of three long-legged and long-necked birds
The 3 of Birds by the Master of the Playing Cards, 15th-century Germany
Birds play prominent and diverse roles in religion and mythology. In religion, birds may serve as either messengers or priests and leaders for a deity, such as in the Cult of Makemake, in which the Tangata manu of Easter Island served as chiefs[284] or as attendants, as in the case of Hugin and Munin, the two common ravens who whispered news into the ears of the Norse god Odin. In several civilisations of ancient Italy, particularly Etruscan and Roman religion, priests were involved in augury, or interpreting the words of birds while the "auspex" (from which the word "auspicious" is derived) watched their activities to foretell events.[285]
They may also serve as religious symbols, as when Jonah (Hebrew: יונה, dove) embodied the fright, passivity, mourning, and beauty traditionally associated with doves.[286] Birds have themselves been deified, as in the case of the common peacock, which is perceived as Mother Earth by the people of southern India.[287] In the ancient world, doves were used as symbols of the Mesopotamian goddess Inanna (later known as Ishtar),[288][289] the Canaanite mother goddess Asherah,[288][289][290] and the Greek goddess Aphrodite.[288][289][291][292][293] In ancient Greece, Athena, the goddess of wisdom and patron deity of the city of Athens, had a little owl as her symbol.[294][295][296] In religious images preserved from the Inca and Tiwanaku empires, birds are depicted in the process of transgressing boundaries between earthly and underground spiritual realms.[297] Indigenous peoples of the central Andes maintain legends of birds passing to and from metaphysical worlds.[297]
In culture and folklore
Painted tiles with design of birds from Qajar dynasty
Birds have featured in culture and art since prehistoric times, when they were represented in early cave paintings.[298] Some birds have been perceived as monsters, including the mythological Roc and the Māori's legendary Pouākai, a giant bird capable of snatching humans.[299] Birds were later used as symbols of power, as in the magnificent Peacock Throne of the Mughal and Persian emperors.[300] With the advent of scientific interest in birds, many paintings of birds were commissioned for books.[citation needed]
Among the most famous of these bird artists was John James Audubon, whose paintings of North American birds were a great commercial success in Europe and who later lent his name to the National Audubon Society.[301] Birds are also important figures in poetry; for example, Homer incorporated nightingales into his Odyssey, and Catullus used a sparrow as an erotic symbol in his Catullus 2.[302] The relationship between an albatross and a sailor is the central theme of Samuel Taylor Coleridge's The Rime of the Ancient Mariner, which led to the use of the term as a metaphor for a 'burden'.[303] Other English metaphors derive from birds; vulture funds and vulture investors, for instance, take their name from the scavenging vulture.[304]
Perceptions of bird species vary across cultures. Owls are associated with bad luck, witchcraft, and death in parts of Africa,[305] but are regarded as wise across much of Europe.[306] Hoopoes were considered sacred in Ancient Egypt and symbols of virtue in Persia, but were thought of as thieves across much of Europe and harbingers of war in Scandinavia.[307] In heraldry, birds, especially eagles, often appear in coats of arms.[308]
In music
Main article: Birds in music
In music, birdsong has influenced composers and musicians in several ways: they can be inspired by birdsong; they can intentionally imitate bird song in a composition, as Vivaldi, Messiaen, and Beethoven did, along with many later composers; they can incorporate recordings of birds into their works, as Ottorino Respighi first did; or like Beatrice Harrison and David Rothenberg, they can duet with birds.[309][310][311][312]
Conservation
Main article: Bird conservation
See also: Late Quaternary prehistoric birds, List of extinct birds, and Raptor conservation
Large black bird with featherless head and hooked bill
The California condor once numbered only 22 birds, but conservation measures have raised that to over 500 today.
Although human activities have allowed the expansion of a few species, such as the barn swallow and European starling, they have caused population decreases or extinction in many other species. Over a hundred bird species have gone extinct in historical times,[313] although the most dramatic human-caused avian extinctions, eradicating an estimated 750–1800 species, occurred during the human colonisation of Melanesian, Polynesian, and Micronesian islands.[314] Many bird populations are declining worldwide, with 1,227 species listed as threatened by BirdLife International and the IUCN in 2009.[315][316]
The most commonly cited human threat to birds is habitat loss.[317] Other threats include overhunting, accidental mortality due to collisions with buildings or vehicles, long-line fishing bycatch,[318] pollution (including oil spills and pesticide use),[319] competition and predation from nonnative invasive species,[320] and climate change.
Governments and conservation groups work to protect birds, either by passing laws that preserve and restore bird habitat or by establishing captive populations for reintroductions. Such projects have produced some successes; one study estimated that conservation efforts saved 16 species of bird that would otherwise have gone extinct between 1994 and 2004, including the California condor and Norfolk parakeet.[321]
See also
Animal track
Avian sleep
Bat
Climate change and birds
Glossary of bird terms
Ornithology
Paleocene dinosaurs
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Further reading
Library resources about
Bird
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Resources in your library
Resources in other libraries
Roger Lederer und Carol Burr: Latein für Vogelbeobachter: über 3000 ornithologische Begriffe erklärt und erforscht, aus dem Englischen übersetzt von Susanne Kuhlmannn-Krieg, Verlag DuMont, Köln 2014, ISBN 978-3-8321-9491-8.
del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi (eds.): Handbook of the Birds of the World (17-volume encyclopaedia), Lynx Edicions, Barcelona, 1992–2010. (Vol. 1: Ostrich to Ducks: ISBN 978-84-87334-10-8, etc.).
All the Birds of the World, Lynx Edicions, 2020.
National Geographic Field Guide to Birds of North America, National Geographic, 7th edition, 2017. ISBN 9781426218354
National Audubon Society Field Guide to North American Birds: Eastern Region, National Audubon Society, Knopf.
National Audubon Society Field Guide to North American Birds: Western Region, National Audubon Society, Knopf.
Svensson, Lars: Birds of Europe, Princeton University Press, second edition, 2010. ISBN 9780691143927
Svensson, Lars: Collins Bird Guide: The Most Complete Guide to the Birds of Britain and Europe, Collins, 2nd edition, 2010. ISBN 978-0007268146
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Birds (class: Aves)
Outline
Anatomy
Beak Crop
Milk Dactyly Eggs Feathers Flight Preen gland Plumage Vision
Behaviour
Singing Intelligence Migration Foraging Sexual selection Lek mating Seabird breeding Incubation Brood parasites Nesting Hybrids
Evolution
Origin of birds
Theropoda dinosaurs Origin of flight Evolution of birds Darwin's finches Seabirds
Fossil birds
Archaeopteryx Omnivoropterygiformes Confuciusornithiformes Enantiornithes Chaoyangiformes Patagopterygiformes Ambiortiformes Songlingornithiformes Hongshanornithidae Gansuiformes Ichthyornithiformes Hesperornithes Lithornithiformes Dinornithiformes Aepyornithiformes Gastornithiformes
Human interaction
Ringing Ornithology Ornithomancy Bird collections Birdwatching
Big year Bird feeding Conservation Aviculture Waterfowl hunting Cockfighting Pigeon racing Falconry Pheasantry Imping Egg collecting
Lists
Families and orders Genera Glossary of bird terms List by population Lists by region Recently extinct birds Late Quaternary prehistoric birds Notable birds
Individuals Fictional
Neornithes
Palaeognathae
Struthioniformes (ostriches) Rheiformes (rheas) Tinamiformes (tinamous) Apterygiformes (kiwis) Casuariiformes (emus and cassowaries)
Neognathae
Galloanserae
(fowls)
Anseriformes
(waterfowls)
Anatidae
(ducks)
Anatinae
Aythyini Mergini Oxyurini Anserinae
swans true geese Dendrocygninae Stictonettinae Tadorninae
Anhimidae
Anhima Chauna
Anseranatidae
Anseranas
Galliformes
(landfowls-
gamebirds)
Cracidae
Cracinae Oreophasinae Penelopinae
Megapodidae
Aepypodius Alectura Eulipoa Leipoa Macrocephalon Megapodius Talegalla
Numididae
Acryllium Agelastes Guttera Numida
Odontophoridae
Callipepla Colinus Cyrtonyx Dactylortyx Dendrortyx Odontophorus Oreortyx Philortyx Rhynchortyx
Phasianidae
Meleagridinae (turkeys) Perdicinae Phasianinae (pheasants and relatives) Tetraoninae
Neoaves
Columbea
Columbimorphae
Columbiformes (doves and pigeons) Mesitornithiformes (mesites) Pterocliformes (sandgrouse)
Mirandornithes
Phoenicopteriformes (flamingos) Podicipediformes (grebes)
Passerea
Otidimorphae
Cuculiformes (cuckoos) Musophagiformes (turacos) Otidiformes (bustards)
Strisores
Caprimulgiformes (nightjars and relatives) Steatornithiformes Podargiformes Apodiformes (swifts and hummingbirds)
Opisthocomiformes
Opisthocomiformes (hoatzin)
Cursorimorphae
Charadriiformes (gulls and relatives) Gruiformes (cranes and relatives)
Phaethontimorphae
Phaethontiformes (tropicbirds) Eurypygiformes (kagu and sunbittern)
Aequornithes
Gaviiformes (loons or divers) Sphenisciformes (penguins) Procellariiformes (albatrosses and petrels) Ciconiiformes (storks) Suliformes (cormorants and relatives) Pelecaniformes (pelicans and relatives)
Australaves
Cariamiformes (seriemas and relatives) Falconiformes (falcons and relatives) Psittaciformes (parrots) Passeriformes (perching birds)
Afroaves
Cathartiformes (New World vultures and condors) Accipitriformes (eagles and hawks) Strigiformes (owls) Coliiformes (mousebirds) Trogoniformes (trogons and quetzals) Leptosomiformes (cuckoo-roller) Bucerotiformes (hornbills and hoopoes) Coraciiformes (kingfishers and rollers) Piciformes (woodpeckers and relatives)
Category Commons Portal WikiProject
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Extant chordate classes
Kingdom Animalia (unranked) Bilateria Superphylum Deuterostomia
Cephalochordata
Leptocardii (lancelets)
Olfactores
Tunicata
(Urochordata)
Ascidiacea (sea squirts) Appendicularia (larvaceans) Thaliacea (pyrosomes, salps, doliolids)
Vertebrata
Cyclostomata
Myxini (hagfish) Hyperoartia (lampreys)
Gnathostomata
(jawed vertebrates)
Chondrichthyes (cartilaginous fish: sharks, rays, chimaeras)
Euteleostomi
(bony vertebrates)
Actinopterygii (ray-finned fish)
Sarcopterygii
(lobe-finned fish)
Actinistia (coelacanths)¹
Rhipidistia
Dipnoi (lungfish)¹
Tetrapoda
Lissamphibia (modern amphibians: frogs, salamanders, caecilians)
Amniota
Mammalia (mammals)
Sauria
Lepidosauria
Rhynchocephalia (tuatara)² Squamata (scaled reptiles)²
Archelosauria
Testudines (turtles)²
Archosauria
Crocodilia (crocodilians)² Aves (birds)
¹subclasses of Sarcopterygii ²orders of class Reptilia (reptiles) italics denote paraphyletic groups
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Maniraptora
Kingdom: Animalia Phylum: Chordata Clade: Dinosauria Clade: Theropoda Clade: Maniraptoriformes
Avemetatarsalia
see Avemetatarsalia
Theropoda
see Theropoda
Maniraptora
see below↓
Maniraptora
Maniraptora
†Elopteryx? †Fukuivenator? †Kakuru? †Yaverlandia?
†Alvarezsauroidea
Alnashetri Aorun? Bannykus Haplocheirus Shishugounykus Tugulusaurus? Xiyunykus Patagonykinae?
Alvarezsauridae
Achillesaurus Alvarezsaurus Bradycneme Heptasteornis
Patagonykinae
Achillesaurus? Bonapartenykus Patagonykus
Parvicursorinae
Dzharaonyx Khulsanurus Kol? Nemegtonykus Ondogurvel Parvicursor Qiupanykus Trierarchuncus
Ceratonykini
Albinykus Ceratonykus Xixianykus
Mononykini
Albertonykus Linhenykus Mononykus Shuvuuia
†Therizinosauria
Eshanosaurus? Falcarius Fukuivenator? Jianchangosaurus Lingyuanosaurus
Therizinosauroidea
Alxasaurus Beipiaosaurus Enigmosaurus Martharaptor Suzhousaurus
Therizinosauridae
Erliansaurus Erlikosaurus Nanshiungosaurus Neimongosaurus Nothronychus Paralitherizinosaurus Segnosaurus Therizinosaurus
Pennaraptora
see below↓
Patagonykus puertai
Mononykus olecranus
Therizinosaurus cheloniformis
Pennaraptora
†Oviraptorosauria
Incisivosaurus Ningyuansaurus Protarchaeopteryx Scansoriopterygidae?
Caudipteridae
Caudipteryx Similicaudipteryx Xingtianosaurus
Caenagnathoidea
Avimimus Kol?
Caenagnathidae
Anomalipes Beibeilong Chirostenotes Gigantoraptor Leptorhynchos Hagryphus Microvenator Nomingia? Ojoraptorsaurus
Elmisaurinae
Citipes Elmisaurus
Caenagnathinae
Anzu Apatoraptor Caenagnathasia Caenagnathus Epichirostenotes
Oviraptoridae
Luoyanggia Nankangia Nomingia? Tongtianlong Yulong
Oviraptorinae
Citipati Corythoraptor? Huanansaurus? Oviraptor Rinchenia?
Heyuanninae
Banji? Conchoraptor Ganzhousaurus? Gobiraptor Heyuannia Jiangxisaurus Khaan Machairasaurus Nemegtomaia Oksoko Shixinggia?
Paraves
†Imperobator †Palaeopteryx? †Pneumatoraptor †Rahonavis
†Scansoriopterygidae?
Ambopteryx Epidexipteryx Scansoriopteryx Yi
†Anchiornithidae
Anchiornis Aurornis Caihong Eosinopteryx Liaoningvenator? Ostromia Pedopenna Serikornis Xiaotingia Yixianosaurus
Eumaniraptora
see below↓
Apatoraptor pennatus
Ambopteryx longibrachium
Anchiornis huxleyi
Eumaniraptora
†Deinonychosauria?
Anchiornithidae? Archaeopterygidae? Troodontidae? Unenlagiidae?
Dromaeosauridae
Pyroraptor Shanag Variraptor Zhenyuanlong Unenlagiidae?
Microraptoria
Changyuraptor Graciliraptor Hesperonychus Microraptor Sinornithosaurus Tianyuraptor Wulong Zhongjianosaurus
Eudromaeosauria
Bambiraptor Tianyuraptor? Vectiraptor Zhenyuanlong?
Saurornitholestinae?
Atrociraptor Bambiraptor? Saurornitholestes
Dromaeosaurinae
Achillobator Dakotaraptor? Deinonychus? Dromaeosauroides Dromaeosaurus Itemirus Saurornitholestes? Utahraptor Yurgovuchia Zapsalis
Velociraptorinae
Acheroraptor Adasaurus Boreonykus? Deinonychus? Dineobellator Kansaignathus Kuru Linheraptor Luanchuanraptor? Nuthetes? Saurornitholestes? Shri Tsaagan Velociraptor
†Troodontidae
Albertavenator Archaeornithoides? Geminiraptor Hesperornithoides Jianianhualong Koparion? Liaoningvenator Papiliovenator Paronychodon? Polyodontosaurus? Sinornithoides Talos Tochisaurus Xixiasaurus Anchiornithidae?
Jinfengopteryginae
Almas? Jinfengopteryx Liaoningvenator? Philovenator? Tamarro
Sinovenatorinae
Daliansaurus Mei Sinovenator Sinusonasus
Troodontinae
Borogovia Byronosaurus? Gobivenator Latenivenatrix Linhevenator Pectinodon Philovenator? Saurornithoides Stenonychosaurus Troodon Urbacodon Zanabazar
†Unenlagiidae
Halszkaraptorinae?
Halszkaraptor Hulsanpes Mahakala
Unenlagiinae
Austroraptor Buitreraptor Dakotaraptor? Neuquenraptor Ornithodesmus? Pamparaptor Pyroraptor? Rahonavis? Unenlagia Unquillosaurus? Ypupiara
Averaptora?
Overoraptor Rahonavis? †Microraptoria? †Troodontidae? †Unenlagiidae?
Avialae
see below↓
Microraptor gui
Utahraptor ostrommaysorum Zanabazar junior
Austroraptor cabazai
Avialae
Avialae
†Alcmonavis †Balaur †Cretaaviculus? †Fukuipteryx †Rahonavis? †Yandangornis †Anchiornithidae? †Scansoriopterygidae?
†Archaeopterygidae?
Alcmonavis? Archaeopteryx Wellnhoferia Anchiornithidae?
†Jeholornithiformes
Dalianraptor? Jeholornis Jixiangornis? Kompsornis Neimengornis
Euavialae
†Jixiangornis?
Avebrevicauda
†Zhongornis
†Omnivoropterygidae
Omnivoropteryx Sapeornis
Pygostylia
†"Proornis" †Omnivoropterygidae?
†Confuciusornithidae
Changchengornis Confuciusornis Eoconfuciusornis Yangavis
†Jinguofortisidae
Chongmingia Jinguofortis
Ornithothoraces
see below↓
Archaeopteryx lithographica Confuciusornis sp.
Ornithothoraces
†Enantiornithes
Brevirostruavis Dalingheornis Elsornis Eoalulavis Eocathayornis? Falcatakely Feitianius Fortipesavis Houornis Ilerdopteryx Liaoningornis Liaoxiornis? Microenantiornis Mirusavis Paraprotopteryx Praeornis? Protopteryx Yuanjiawaornis Yuornis
Iberomesornithidae
Iberomesornis Noguerornis
Pengornithidae
Chiappeavis Eopengornis Parapengornis Pengornis Yuanchuavis
Longipterygidae
Boluochia Camptodontornis Dapingfangornis Evgenavis? Longipteryx Longirostravis Rapaxavis Shanweiniao Shengjingornis
Euenantiornithes
Abavornis Alethoalaornis Alexornis Avimaia Catenoleimus Cathayornis Cratoavis Cruralispennia Cuspirostrisornis Dunhuangia Elbretornis Elektorornis Enantiornis Eoenantiornis Explorornis Flexomornis Fortunguavis Grabauornis Gracilornis Gurilynia Holbotia Huoshanornis Incolornis Junornis Kizylkumavis Kuszholia? Largirostrornis Lectavis Lenesornis Longchengornis Martinavis Monoenantiornis Musivavis Nanantius Orienantius Otogornis Parvavis Piscivorenantiornis Platanavis? Pterygornis Qiliania Sazavis Shangyang Sinornis Xiangornis Yungavolucris
Bohaiornithidae
Beiguornis Bohaiornis Gretcheniao Linyiornis Longusunguis Parabohaiornis Shenqiornis Sulcavis Zhouornis
Gobipterygidae
Gobipteryx Jibeinia? Vescornis?
Avisauridae
(sensu Cau & Arduini, 2008)
Bauxitornis? Concornis? Enantiophoenix Halimornis Mystiornis
Avisauridae (sensu Chiappe, 1992)
Avisaurus Gettyia Intiornis Mirarce Neuquenornis Soroavisaurus
Euornithes
see below↓
Longipteryx chaoyangensis Cruralispennia multidonta
Euornithes
Euornithes
†Archaeorhynchus †Chaoyangia †Gargantuavis? †Horezmavis †Jianchangornis †Platanavis †Wyleyia? †Xinghaiornis †Zhongjianornis
Ornithuromorpha
†Bellulornis †Brevidentavis †Changmaornis †Changzuiornis †Dingavis †Eogranivora †Gansus †Hollanda †Iteravis †Jiuquanornis †Juehuaornis †Kaririavis †Khinganornis †Meemannavis †Vorona †Yumenornis
†Schizoouridae
Mengciusornis Schizooura
†Patagopterygiformes
Alamitornis Kuszholia? Patagopteryx
†Ambiortiformes
Ambiortus Apsaravis? Palintropus?
†Hongshanornithidae
Archaeornithura Hongshanornis Longicrusavis Parahongshanornis Tianyuornis
†Songlingornithidae
Hollanda? Piscivoravis? Songlingornis Yanornis? Yixianornis?
†Yanornithidae?
Abitusavis Similiyanornis Yanornis
Ornithurae
†Antarcticavis? †Apatornis †Cerebavis †Gallornis †Guildavis †Iaceornis †Ichthyornis †Kookne †Limenavis †Qinornis †Tingmiatornis
†Hesperornithes
Baptornis Brodavis Chupkaornis Enaliornis Judinornis Pasquiaornis Potamornis
Hesperornithidae
Asiahesperornis Canadaga Fumicollis Hesperornis Parahesperornis
†Vegaviidae
Australornis? Maaqwi Neogaeornis? Polarornis Vegavis
†Cimolopterygidae
Ceramornis Cimolopteryx Lamarqueavis?
Aves / Neornithes
Palaeognathae
see Palaeognathae
Neognathae
Pangalloanserae Panneoaves
Patagopteryx deferrariisi Ichthyornis dispar
See also: Archaeornithes Carinatae Ichthyornithes Odontognathae Odontornithes Sauriurae Category
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Human uses of birds
Activities
Aviculture Birdwatching
Big year Bird conservation Fletching In sport
Cockfighting Falconry Pigeon racing Vinkensport In science
Model organism Ornithology In mythology and religion
Augury Sacred ibis Sky burial In hunting
Cormorant fishing Driven grouse shooting Plume hunting Wildfowling
Falconry Book of Frederick II 1240s detail falconers.jpg
Products
Chicken Down Egg Feather Guano Poultry
In the arts
In art
Bird-and-flower painting Feather tights In heraldry
Avalerion Crow/Raven Eagle Gallic rooster Martlet Turul In poetry
The Conference of the Birds Ode to a Nightingale To a Skylark Crow In prose
A History of British Birds The Tale of Jemima Puddle-Duck The Ugly Duckling Jonathan Livingston Seagull In theatre and ballet
The Birds Swan Lake The Firebird In film
The Birds Kes The Big Year Animated films Chicken films Horror films In music In fashion
Aigrette Feather boa Feather cloak In dance
Cendrawasih Chicken dance
Species
Golden eagle Penguin Pigeon/Dove Raven
of the Tower of London
People
Illustrators
John James Audubon (The Birds of America) Thomas Bewick John Gould Lars Jonsson John Gerrard Keulemans Edward Lear Richard Lewington Roger Tory Peterson Henry Constantine Richter Joseph Smit Archibald Thorburn Joseph Wolf Conservationists
Niels Krabbe Peter Scott Organisations
BirdLife International Royal Society for the Protection of Birds Wildfowl & Wetlands Trust
Related
Category:Birds and humans Zoomusicology
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Lists of dinosaurs by continent
Taxon identifiers
Wikidata: Q5113 Wikispecies: Aves ADW: Aves AFD: Aves BOLD: 51 CoL: V2 EoL: 695 EPPO: 1AVESC Fauna Europaea: 10699 Fauna Europaea (new): f2fd1555-ab1f-40f7-9cbf-abebff1ffbda Fossilworks: 36616 GBIF: 212 iNaturalist: 3 IRMNG: 1142 ITIS: 174371 NCBI: 8782 NZOR: dcced4e7-06b1-466e-9d85-5a384501dac2 Plazi: E1E0B077-76F6-D736-3B27-36617A705C73 uBio: 21646 WoRMS: 1836 ZooBank: AAFCA22F-1980-4B62-9149-8887F1C1FDC1
Authority control Edit this at Wikidata
National libraries
Spain France (data) Germany Israel United States Latvia Japan Czech Republic
Other
National Archives (US)
Categories:
BirdsAnimal classesDinosaursExtant Late Cretaceous first appearancesFeathered dinosaursSantonian first appearancesTaxa named by Carl Linnaeus
List of birds
This article lists living orders and families of birds. The links below should then lead to family accounts and hence to individual species.
The passerines (perching birds) alone account for well over 5,000 species. In total there are about 10,000 species of birds described worldwide, though one estimate of the real number places it at almost twice that.[1]
Taxonomy is very fluid in the age of DNA analysis, so comments are made where appropriate, and all numbers are approximate. In particular see Sibley-Ahlquist taxonomy for a very different classification.
Phylogeny
Cladogram of modern bird relationships based on Jarvis, E.D. et al. (2014)[2] with some clade names after Yuri, T. et al. (2013).[3]
Aves
Palaeognathae
Struthionimorphae
Struthioniformes (ostriches)[4]
Notopalaeognathae
Rheimorphae
Rheiformes (rheas)
Novaeratitae
Casuariiformes (cassowaries & emus)
Apterygiformes (kiwi)
†Aepyornithiformes (elephant birds)
Tinamimorphae
†Dinornithiformes (moas)
†Lithornithiformes (false tinamous)
Tinamiformes (tinamous)
Neognathae
Galloanserae
Gallomorphae
Galliformes (landfowl)
Odontoanserae
†Odontopterygiformes
Anserimorphae
†Vegaviiformes[5]
†Gastornithiformes
Anseriformes (waterfowl)
Neoaves
Columbea
Mirandornithes
Phoenicopteriformes (flamingoes)
Podicipediformes (grebes)
Columbimorphae
Mesitornithiformes (mesites)
Pterocliformes (sandgrouse)
Columbiformes (pigeons)
Passerea
Otidae
Otidimorphae
Cuculiformes (cuckoos)
Otidiformes (bustards)
Musophagiformes (turacos)
Cypselomorphae
Caprimulgiformes (nightjars)
Nyctibiiformes (oilbirds & potoos)
Podargiformes (frogmouths)
Aegotheliformes (owlet-nightjars)
Apodiformes (hummingbirds & swifts)
Gruae
Opisthocomiformes (hoatzin)
Cursorimorphae
Gruiformes (rails and cranes)
Charadriiformes (shorebirds)
Ardeae
Phaethontimorphae
Eurypygiformes (sunbittern, kagu)
Phaethontiformes (tropicbirds)
Aequornithes
Gaviiformes (loons)
Austrodyptornithes
Procellariiformes (albatross and petrels)
Sphenisciformes (penguins)
Ciconiiformes (storks)
Suliformes (boobies, cormorants, etc.)
Pelecaniformes (pelicans, herons & egrets)
Telluraves
Afroaves
Accipitrimorphae
Cathartiformes (condors and New World vultures)
Accipitriformes (hawks, eagles, vultures, etc.)
Strigiformes (owls)
Coraciimorphae
Coliiformes (mousebirds)
Leptosomiformes (cuckoo roller)
Trogoniformes (trogons)
Bucerotiformes (hornbills, hoopoe and wood hoopoes)
Coraciiformes (kingfishers etc.)
Piciformes (woodpeckers etc.)
Picodynastornithes
Picocoraciae
Eucavitaves
Cavitaves
Australaves
Cariamiformes (seriemas)
Eufalconimorphae
Falconiformes (falcons)
Psittacopasserae
Psittaciformes (parrots)
Passeriformes (songbirds and kin)
Paleognathae
The Paleognathae, or "old jaws", are one of the two superorders recognized within the taxonomic class Aves and consist of the ratites and tinamous. The ratites are mostly large and long-legged, flightless birds, lacking a keeled sternum. Traditionally, all the ratites were place in the order Struthioniformes. However, recent genetic analysis has found that the group is not monophyletic, as it is paraphyletic with respect to the tinamous, so the ostriches are classified as the only members of the order Struthioniformes and other rattites placed in other orders.[6][7]
Struthioniformes
Greater rhea pair
Eudromia elegans
Casuarius casuarius
Africa; 2 species
Struthionidae: ostrich
Notopalaeognathae
Rheiformes
South America; 2 species
†Opisthodactylidae
Rheidae: rheas
Casuariiformes
Australasia; 4 species
Casuariidae: cassowaries and emu
Apterygiformes
Australasia; 5 species
Apterygidae: kiwis
Aepyornithiformes
Madagascar
†Aepyornithidae: elephant birds
Dinornithiformes
New Zealand
†Megalapteryidae: upland moas
†Dinornithidae: great moas
†Emeidae: lesser moas
Tinamiformes
South America; 45 species
Tinamidae: tinamous
Neognathae
Nearly all living birds belong to the superorder Neognathae or "new jaws". With their keeled sternum (breastbone), unlike the ratites, they are known as carinatae.
Galloanserae
Galliformes
Australian brush turkey
Worldwide; 250 species
†Sylviornithidae
Megapodii
Megapodidae: megapodes
Craci
Cracidae: chachalacas, curassows, and guans
Phasiani
Numidioidea
Numididae: guineafowl
Phasianoidea: pheasants and allies
Odontophoridae: New World quail
Phasianidae: pheasants and relatives
Gastornithiformes
†Gastornithidae
†Dromornithidae: mihirungs
Anseriformes
Worldwide; 150 species
Anhimidae: screamers
Anseranatidae: magpie-goose
Anatidae: ducks, geese, and swans
Mirandornithes
Podicipediformes
Worldwide; 19 species
Podicipedidae: grebes
Phoenicopteriformes
Worldwide; 6 species
†Palaelodidae: swimming flamingos
Phoenicopteridae: flamingos
Columbimorphae
Columbiformes
Worldwide; 300 species
Columbidae: pigeons and doves
Pterocliformes
Africa, Europe, Asia; 16 species
Pteroclidae: sandgrouse
Mesitornithiformes
Madagascar; 3 species
Mesitornithidae: mesites
Cypselomorphae
Caprimulgiformes
Worldwide; 97 species
Caprimulgidae: nightjars
Steatornithiformes
South America; 1 species
Steatornithidae: oilbird
Nyctibiiformes
Americas; 7 species
Nyctibiidae: potoos
Podargiformes
Tawny frogmouth
Asia and Australasia; 16 species
Podargidae: frogmouths
Aegotheliformes
Australasia; 9 species
Aegothelidae: owlet-nightjars
Apodiformes
Worldwide; 478 species
Hemiprocnidae: treeswifts
Apodidae: swifts
Trochilidae: hummingbirds
Otidimorphae
Cuculiformes
Worldwide; 126 species
Cuculidae: cuckoos and relatives
Musophagiformes
Africa; 23 species
Musophagidae: turacos and relatives
Otidiformes
Africa and Eurasia; 27 species
Otididae: bustards
Gruae
Opisthocomiformes
South America; 1 species
Opisthocomidae: hoatzin
Gruiformes
Worldwide; 164 species
Grui: cranes and allies
Gruidae: cranes
Aramidae: limpkin
Psophiidae: trumpeters
Ralli: rails and allies
†Aptornithidae: adzebills
Heliornithidae: finfoots
Sarothruridae: flufftails
Rallidae: rails and relatives
Charadriiformes
Worldwide; 350 species
Charadrii
Chionida: thick-knees and allies
Burhinidae: thick-knees and relatives
Chionididae: sheathbills
Pluvianellidae: Magellanic plover
Charadriida: plover-like waders
Pluvialidae: golden plovers
Ibidorhynchidae: ibisbill
Haematopodidae: oystercatchers
Recurvirostridae: avocets and stilts
Charadriidae: plovers and lapwings
Scolopaci
Jacanida: jacana-like waders
Greater painted-snipe
Rostratulidae: painted snipes
Egyptian plover
Pluvianidae: Egyptian plover
Jacanidae: jacanas
Thinocoridae: seedsnipes
Plains-wanderer
Pedionomidae: plains-wanderer
Scolopacida
Scolopacidae: sandpipers and relatives
Lari
Turnicida
Turnicidae: buttonquail
Larida: gulls and allies
Glareolidae: coursers and pratincoles
Dromadidae: crab-plover
Stercorariidae: skuas and jaegers
Alcidae: auks and puffins
Laridae: gulls, skimmers and terns
Phaethontimorphae
Eurypygiformes
Neotropics and New Caledonia; 2 species
Rhynochetidae: kagu
Sunbittern
Eurypygidae: sunbittern
Phaethontiformes
Oceanic; 3 species
Phaethontidae: tropicbirds
Aequornithes
Gaviiformes
North America, Eurasia; 5 species
Gaviidae: loons
Sphenisciformes
Antarctic and southern waters; 17 species
Spheniscidae: penguins
Procellariiformes
Pan-oceanic; 120 species
Diomedeidae: albatrosses
Oceanitidae: austral storm petrels
Hydrobatidae: northern storm petrels
Procellariidae: petrels and relatives
Ciconiiformes
Worldwide; 19 species
White stork
Ciconiidae: storks
Suliformes
Worldwide; 59 species
Fregatae
Fregatidae: frigatebirds
Sulae
Sulidae: boobies and gannets
Anhingidae: darters
Phalacrocoracidae: cormorants and shags
Pelecaniformes
Hamerkop
Worldwide; 108 species
Threskiornithes
Threskiornithidae: ibises and spoonbills
Pelecani
Scopidae: hamerkop
Balaenicipitidae: shoebill
Pelecanidae: pelicans
Ardeae
Ardeidae: herons and relatives
Afroaves
Accipitriformes
Osprey
Worldwide; 200 species
Cathartae
Cathartidae: New World vultures
Accipitres
Sagittariidae: secretarybird
Pandionidae: osprey
Accipitridae: hawks, eagles, buzzards, harriers, kites and Old World vultures
Strigiformes
Worldwide; 130 species
Tytonidae: barn owls
Strigidae: true owls
Coliiformes
Blue-naped mousebird
Sub-Saharan Africa; 6 species
Coliidae: mousebirds
Leptosomiformes
Madagascar; 1 species
Leptosomidae: cuckoo-roller
Trogoniformes
Sub-Saharan Africa, Americas, Asia; 35 species
Trogonidae: trogons and quetzals
Bucerotiformes
Old World, New Guinea; 64 species
Buceroidea
Bucerotidae: hornbills
Upupoidea
Upupidae: hoopoe
Phoeniculidae: woodhoopoes
Coraciiformes
Worldwide; 144 species
Meropi
Meropidae: bee-eaters
Coracii
Coraciidae: rollers
Brachypteraciidae: ground rollers
Coracii
Todidae: todies
Momotidae: motmots
Alcedinidae: kingfishers
Kingfisher
Piciformes
Worldwide except Australasia; 400 species
Galbuli
Galbulidae: jacamars
Bucconidae: puffbirds
Pici
Lybiidae: African barbets
Megalaimidae: Asian barbets
Ramphastidae: toucans
Semnornithidae: toucan barbets
Capitonidae: American barbets
Picidae: woodpeckers
Indicatoridae: honeyguides
Australaves
Cariamiformes
South America; 2 species
Cariamidae: seriemas
Falconiformes
Worldwide; 60 species
Falconidae: falcons and relatives
Psittaciformes
Pan-tropical, southern temperate zones; 330 species
Strigopoidea
Strigopidae: kakapo, kea and kakas
Cacatuoidea
Cacatuidae: cockatoos and cockatiels
Psittacoidea
Psittacidae: African and American parrots
Psittaculidae: Australasian parrots, Pesquet's parrot, vasa parrots
Passeriformes
Rock wren
Eurylaimus javanicus
Pitta cyanea
Pachyramphus castaneus
Lyrebird
Worldwide; 5,000 species
Acanthisitti
Acanthisittidae: New Zealand wrens
Tyranni: suboscines
Eurylaimides: Old World suboscines
Sapayoidae: sapayoa
Calyptomenidae: Calyptomenid broadbills
Pittidae: pittas
Eurylaimidae: broadbills
Philepittidae: asities
Tyrannides: New World suboscines
Tyrannida: bronchophones
Pipridae: manakins
Cotingidae: cotingas
Oxyruncidae: sharpbills
Onychorhynchidae: royal flycatchers and allies
Tityridae: becards and tityras
Pipritidae: pipriteses
Platyrinchidae: spadebills
Tachurididae: many-colored rush tyrants
Rhynchocyclidae: mionectine flycatchers
Tyrannidae: tyrant flycatchers
Furnariida: tracheophones
Melanopareiidae: crescent-chests
Conopophagidae: gnateaters
Thamnophilidae: antbirds
Grallariidae: antpittas
Rhinocryptidae tapaculos
Formicariidae: ground antbirds
Furnariidae: ovenbirds
Passeri: oscines
Menurides
Atrichornithidae: scrub-birds
Menuridae: lyrebirds
Climacterides
Ptilonorhynchidae: bowerbirds
Climacteridae: Australasian treecreepers
Meliphagides
Maluridae: Australasian wrens
Dasyornithidae: bristlebirds
Pardalotidae: gerygones and allies (Acanthizidae)
Meliphagidae: honeyeaters and relatives
Orthonychides
Pomatostomidae: Australasian babblers
Orthonychidae: logrunners
Corvides
Cinclosomatoidea
Cinclosomatidae: quail-thrushes and jewel-babblers
Campephagoidea
Campephagidae: cuckoo-shrikes
Mohouoidea
Mohouidae: whitehead and allies
Neosittoidea
Neosittidae: sittellas
Orioloidea
Eulacestomidae: wattled ploughbills
Psophodidae: whipbirds and quail-thrushes
Oreoicidae: Australo-Papuan bellbirds
Falcunculidae: crested shriketits
Paramythiidae: painted berrypeckers
Pteruthiidae: shrike-babblers
Vireonidae: vireos and relatives
Pachycephalidae: whistlers and relatives (Colluricinclidae)
Oriolidae: Old World orioles
Malaconotoidea
Machaerirhynchidae: boatbills
Artamidae: woodswallows and butcherbirds
Rhagologidae: mottled berryhunter
Aegithinidae: ioras
Pityriaseidae: bristlehead
Malaconotidae: bushshrikes and relatives
Platysteiridae: wattle-eyes and batises
Vangidae: vangas (Tephrodornithidae; Prionopidae)
Corvoidea
Rhipiduridae: fantails
Lamproliidae: silktail, drongo fantail
Dicruridae: drongos
Ifritidae: blue-capped ifrits
Melampittidae: melampittas
Corcoracidae: Australian mudnesters
Paradisaeidae: birds-of-paradise
Monarchidae: monarch flycatchers
Laniidae: shrikes
Corvidae: jays and crows
Passerides
Melanocharitida
Melanocharitidae: berrypeckers
Cnemophilida
Cnemophilidae: satinbirds
Petroicida
Petroicidae: Australasian robins
Notiomystidae: stitchbird
Callaeidae: wattlebirds
Eupetida
Picathartidae: rockfowl
Chaetopidae: rock-jumpers
Eupetidae: rail-babbler
Sylviida
Paroidea
Stenostiridae: fairy warblers
Hyliotidae: hyliotas
Remizidae: penduline tits
Paridae: chickadees and true tits
Alaudoidea
Nicatoridae: nicators
Panuridae: bearded reedling
Alaudidae: larks
Macrosphenidae: African warblers
Locustelloidea
Cisticolidae: cisticolas and relatives
Acrocephalidae: marsh warblers
Pnoepygidae: pygmy wren-babblers
Locustellidae: grass warblers
Donacobiidae: donacobius
Bernieridae: Malagasy warblers
Hirundinidae: swallows and martins
Pycnonotidae: bulbuls
Aegithaloidea
Phylloscopidae: leaf warblers
Cettiidae: bush warblers (Erythrocercidae; Scotocercidae)
Hyliidae: hylias
Aegithalidae: bushtits
Sylvioidea
Sylviidae: true warblers
Paradoxornithidae: parrotbills, fulvettas
Zosteropidae: white-eyes
Timaliidae: babblers and relatives
Pellorneidae: fulvettas, ground babblers
Leiothrichidae: laughing thrushes
Muscicapida
Reguloidea
Regulidae: kinglets
Bombycilloidea
Elachuridae: spotted wren-babblers
†Mohoidae: Hawaiian honeyeaters
Ptiliogonatidae: silky-flycatchers
Bombycillidae: waxwings
Dulidae: palmchat
Hypocoliidae: hypocolius
Certhioidea
Tichodromidae: wallcreeper
Sittidae: nuthatches
Certhiidae: treecreepers
Troglodytidae: wrens
Polioptilidae: gnatcatchers
Muscicapoidea
Cinclidae: dippers
Turdidae: thrushes and relatives
Muscicapidae: flycatchers and relatives
Buphagidae: oxpeckers
Mimidae: mockingbirds and thrashers
Sturnidae: starlings and mynas (Rhabdornithidae)
Passerida
Promeropidae: sugarbirds
Arcanatoridae: dapplethroat and allies
Dicaeidae: flowerpeckers
Nectariniidae: sunbirds
Irenidae: fairy-bluebirds
Chloropseidae: leafbirds
Peucedramidae: olive warbler
Prunellidae: accentors
Ploceoidea
Urocynchramidae: pink-tailed bunting
Ploceidae: weavers and relatives
Viduidae: whydahs and indigobirds
Estrildidae: weaver finches
Passerid clade
Passeridae: Old World sparrows
Motacillidae: wagtails and pipits
Fringillidae: finches and relatives
Calcariidae: longspurs, snow buntings
Rhodinocichlidae: rosy thrush-tanagers
Emberizidae: Old World buntings
Passerellidae: American sparrows
Phaenicophilidae: palm-tanager and allies
Icteridae: New World blackbirds and New World orioles
Teretistridae: Cuban warblers
Parulidae: wood warblers
Mitrospingidae
Cardinalidae: cardinals, grosbeaks, and New World buntings
Thraupidae: tanagers and relatives (Coerebidae)
See also
Lists of animals
List of chicken breeds
List of birds by common name
List of individual birds
Lists by continent
List of birds of Africa
List of birds of Antarctica
List of birds of Asia
List of birds of Australia
List of birds of Europe
List of birds of North America
List of birds of South America
Lists by smaller geographic unit
Lists of birds by region
Extinct birds
List of recently extinct bird species
List of Late Quaternary prehistoric bird species
List of fossil bird genera
List of fictional birds
References
Barrowclough, GF; Cracraft, J; Klicka, J; Zink, RM (2016). "How Many Kinds of Birds Are There and Why Does It Matter?". PLOS ONE. 11 (11): e0166307. Bibcode:2016PLoSO..1166307B. doi:10.1371/journal.pone.0166307. PMC 5120813. PMID 27880775.
Jarvis, E.D.; et al. (2014). "Whole-genome analyses resolve early branches in the tree of life of modern birds". Science. 346 (6215): 1320–1331. Bibcode:2014Sci...346.1320J. doi:10.1126/science.1253451. PMC 4405904. PMID 25504713.
Yuri, T.; et al. (2013). "Parsimony and Model-Based Analyses of Indels in Avian Nuclear Genes Reveal Congruent and Incongruent Phylogenetic Signals". Biology. 2 (1): 419–444. doi:10.3390/biology2010419. PMC 4009869. PMID 24832669.
Boyd, John (2007). "NEORNITHES: 46 Orders" (PDF). John Boyd's website. Retrieved 30 December 2017.[unreliable source?]
Worthy, T.H.; Degrange, F.J.; Handley, W.D.; Lee, M.S.Y. (2017). "The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres)". Royal Society Open Science. 11 (10): 170975. Bibcode:2017RSOS....470975W. doi:10.1098/rsos.170975. PMC 5666277. PMID 29134094.
Hackett, S.J.; et al. (2008). "A Phylogenomic Study of Birds Reveals Their Evolutionary History". Science. 320 (5884): 1763–1768. Bibcode:2008Sci...320.1763H. doi:10.1126/science.1157704. PMID 18583609. S2CID 6472805.
Yuri, T (2013). "Parsimony and model-based analyses of indels in avian nuclear genes reveal congruent and incongruent phylogenetic signals". Biology. 2 (1): 419–44. doi:10.3390/biology2010419. PMC 4009869. PMID 24832669.
Table of Contents
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Famous Rabbits In Movies or Television
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A Famous Rabbit From Literature & From Movies or Television
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Cadbury Bunny (Cadbury Commercials)
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Captain Carrot (Zoo Crew Comics)
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Crusader Rabbit (Crusader Rabbit)
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Daisy The Rabbit with Longest Tail
Famous Real Life Rabbit
Darius the Largest Rabbit in the World
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Easter Bunny
Famous Rabbit From Literature
Edward Tulane (The Miraculous Journey of Edward Tulane)
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Energizer Bunny (TV Commercial)
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Flopsy the Oldest Rabbit Ever
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Harvey (Harvey)
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Hodge Podge (Bloom County Comics)
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Honey Bunny (Looney Tunes)
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Hoppy The Marvel Bunny (Hoppy the Marvel Bunny)
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Jackalope of the Smithsonian
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Jazz Jackrabbit (Jazz JackRabbit Video Game)
Famous Rabbit From Video Games
Judy Hops (Zootopia)
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Lola Bunny (Looney Tunes)
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March Hare (Alice in Wonderland)
Famous Rabbit From Literature And Movies
Marlon Bundo (A Day in The Life of Marlon Bundo)
A Famous Rabbit From Literature & From Real Life
Mick The Agouti Rabbit
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Miffy (Nijntje)
Famous Rabbit From Literature And Movies And Television
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Famous Rabbit From Comics
Mr. Bunny Rabbit (Captain Kangaroo)
Famous Rabbits In Movies or Television
Quicky The Nesquik Bunny
Famous Rabbits In Movies or Television
Oswald the Lucky Rabbit
Famous Rabbit From Comic Books And Television
Peppy Hare (Star Fox Game)
Famous Rabbits In Video Games
Peter Rabbit (The Tale of Peter Rabbit)
A Famous Rabbit From Literature & From Movies or Television
Playboy Bunny
Famous Rabbit From Real Life
Rabbit (Winnie The Pooh)
A Famous Rabbit From Literature & From Movies or Television
Rabbit of Caerbanog (Monty Phyton)
Famous Rabbit From Movies And Television
Ricochet Rabbit (The Magilla Gorilla Show)
Famous Rabbits In Movies or Television
Roger Rabbit (Who Framed Roger Rabbit)
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Schnuffel
Famous Rabbit From Movies & Television
Skippy Rabbit (Robin Hood)
Famous Rabbit From Movies & Television
Snowball The Rabbit (Secret Life Of Pets)
Famous Rabbits In Movies or Television
Taawi The Multi Talented Rabbit
Famous Real Life Rabbit
The White Rabbit (Alice in Wonderland)
A Famous Rabbit From Literature
The Velveteen Rabbit (The Velveteen Rabbit)
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Thunder Bunny (Thunder Bunny Comics)
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Trix Rabbit (Trix Cereals)
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