About
Anatomy
Burrowing owl (Athene cunicularia)
Captive short-eared owl chick at about 18 days old
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 in order 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 head 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 degrees. Owls have fourteen 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: theforamina in their vertebrae through which the vertebral arteries pass are about ten times the diameter of the artery, instead of about the same size as the artery like 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.12
The smallest owl—weighing as little as 31 grams (1 oz) and measuring some 13.5 centimetres (5 in)—is the elf owl (Micrathene whitneyi).3Around the same diminutive length, although slightly heavier, are the lesser known long-whiskered owlet (Xenoglaux loweryi) and Tamaulipas pygmy owl (Glaucidium sanchezi).3 The largest owl by length is the great grey owl (Strix nebulosa), which measures around 70 cm (28 in) on average and can attain a length of 84 cm (33 in).3 However, the heaviest (and largest winged) owls are two similarly-sized eagle owls; theEurasian eagle-owl (Bubo bubo) and Blakiston's fish owl (B. blakistoni). These two species, which are on average about 2.53 cm (1.00 in) shorter in length than the great grey, can both attain a wingspan of 2 m (6.6 ft) and a weight of 4.5 kg (10 lb) in the largest females.34567
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, the facial disc helps 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 coloured 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.8
Sexual dimorphism
Sexual dimorphism is a physical difference between males and females of a species. Reverse sexual dimorphism is when females are larger than males and it has been observed across multiple owl species.9 The degree of size dimorphism varies across multiple populations and species, and is measured through various traits, such as wing span and body mass.9 Overall, female owls tend to be slightly larger than males. The exact explanation for this development in owls is unknown. However, there are several theories explaining the development of sexual dimorphism in owls.
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.10 However, if food is scarce, the male will first feed itself before feeding the female.11 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.11 Furthermore, owls have been observed to be roughly the same size as their prey.11 This has also been observed in other predatory birds.10 This 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.
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 without starving for a longer period of time. For example, one hypothesized sexual role is that larger females are more capable of dismembering prey and feeding it to their young, hence why female owls are larger than their male counterparts.9
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.11
Breeding and reproduction
Owl eggs typically have a white colour 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 life time. It is common for female burrowing owls to travel and find other mates, while the male stays in its territory and mates with other females.12
Behavior
The serrations on the leading edge of an owl's flight feathers reduce noise
Owl eyes each havenictitating membranes that can move independently of each other, as seen on this spotted eagle-owl in Johannesburg,South Africa
Owls yawn
Most owls are nocturnal, actively hunting their prey in darkness. Several types of owl, however, 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 theburrowing 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.13
Adaptations for hunting
All owls are carnivorous birds of prey and live mainly on a diet of insects and small rodents such as mice, rats and hares. 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 their prey that are listening for 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. While the morphological and biological mechanisms of this silent flight are more or less unknown, the structure of the feather has been heavily studied and accredited to a large portion of why they have this ability. Owls’ feathers are generally larger than the average birds’ feathers, have fewer radiates, longer pennulum, and achieve smooth edges with different rachis structures.14 Serrated edges along the owl’s remiges bring the flapping of the wing down to a nearly silent mechanism. Research has shown that the serrations are more likely reducing aerodynamic disturbances, rather than simply reducing noise.14 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,15 making the sound level emitted drop below the typical hearing spectrum of the owl’s usual prey1516 and also within the owl’s own best hearing range.citation needed This optimizes the owl’s ability to silently fly in order to capture prey without the prey hearing the owl first as it flies in. It also allows the owl to monitor the sound output from its flight pattern.
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 cattle 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.17
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 skull. 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.18 Since the eyes are fixed into these sclerotic tubes, they are unable to move the eyes in any direction.19Instead of moving their eyes, owls swivel their head to visualize their surroundings. Owls' heads are capable of swiveling through an angle of approximately 270°, easily enabling them to see behind them without relocating the torso.19 This ability keeps bodily movement at a minimum and 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. However, owls are farsighted and cannot focus on objects within a few centimeters of their eyes.1820 While it is commonly believed that owls have great nocturnal vision due to their large (and thus very light-gathering) eyes and pupils and/or extremely sensitive rod receptors, the true cause for their ability to see in the night is due to neural mechanisms which mediate the extraction of spatial information gathered from the retinal image throughout the nocturnal luminance range. These mechanisms are only able to function due to the large-sized retinal image.21 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.21 These attributes of the owl cause its nocturnal eyesight to be far superior to that of its average prey.21
Hearing
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.19 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.citation needed 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 a matter of about 0.00003 seconds, or 30 millionths of a second. Like the eyes, which utilize feather movements to focus light,citation neededdubious – discuss the ears are surrounded by feathers to maximize hearing capabilities. Behind the ear openings there are modified, dense feathers, densely packed to form a facial ruff, which creates an anteriorly-facing concave wall that cups the sound into the ear structure.22 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.19
It is a common misapprehension that the prominences above a great horned owl's head are 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 by using these talons to crush the skull and knead the body.19 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 of over 130 N to release prey in its talons.23 An owl’s talons, like those of most birds of prey, can seem massive in comparison to the body size outside of flight. The 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.24 An owl’s claws are sharp and curved. The family Tytonidae have inner and central toes of about equal length, while the family Strigidae have an inner toe that is distinctly shorter than the central one.23 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.citation needed
Snowy owl blends well with its snowy surroundings
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 colorations and sometimes even the texture patterns of their surroundings, the common barn owl being an exception. Nyctea scandiaca, or the snowy owl, appears nearly bleach-white in color with a few flecks of black, mimicking their snowy surroundings perfectly. 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 will be its vocalizations or its vividly colored eyes.
Evolution and systematics
Great horned owl (Bubo virginianus) sleeping during daytime in a hollow tree
The systematic placement of owls is disputed. For example, the Sibley-Ahlquist taxonomy finds that, based on DNA-DNA hybridization, owls are more closely related to the nightjars and their allies (Caprimulgiformes) than to the diurnal predators in the order Falconiformes; consequently, the Caprimulgiformes are placed in the Strigiformes, and the owls in general become a family Strigidae. A recent study indicates that the drastic rearrangement of the genome of the accipitrids may have obscured any close relationship of theirs with groups such as the owls.25 In any case, the relationships of the Caprimulgiformes, the owls, the falcons and the accipitrid raptors are not resolved to satisfaction; currently there is an increasing trend to consider each group (with the possible exception of the accipitrids) a distinct order.
There are some 220 to 225 extant species of owls, subdivided into two families: typical owls (Strigidae) and barn-owls (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 mya (million years ago), and, 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.26
During the Paleogene, the Strigiformes radiated into ecological niches now mostly filled by other groups of birds.clarification needed The owls as we know them today, on the other hand, 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) owl similar to today's North American spotted owl or the European tawny owl; the diversity in size and ecologyfound 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 typical owls, which for the most part seems to have taken place in Eurasia. In the Americas, there was rather an expansion of immigrant lineages of ancestral typical owls.
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.27 TheMesselasturidae, some of which were initially believed to be basal Strigiformes, are now generally accepted to be diurnal birds of prey showing some convergent evolutiontowards owls. The taxa often united under Strigogyps28 were formerly placed in part with the owls, specifically the Sophiornithidae; they appear to be Ameghinornithidaeinstead.293031
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