great horned owl

The great horned owl (''Bubo virginianus''), also known as the tiger owl (originally derived from early naturalists' description as the "winged tiger" or "tiger of the air"), or the hoot owl, is a large owl native to the Americas. It is an extremely adaptable bird with a vast range and is the most widely distributed true owl in the Americas. Its primary diet is rabbits and hares, rats and mice, and voles, although it freely hunts any animal it can overtake, including rodents and other small mammals, larger mid-sized mammals, birds, reptiles, amphibians, and invertebrates. In ornithological study, the great horned owl is often compared to the Eurasian eagle-owl (''Bubo bubo''), a closely related species, which despite the latter's notably larger size, occupies the same ecological niche in Eurasia, and the red-tailed hawk (''Buteo jamaicensis''), with which it often shares similar habitat, prey, and nesting habits by day, thus is something of a diurnal ecological equivalent. The great horned owl is one of the earliest nesting birds in North America, often laying eggs weeks or even months before other raptorial birds.

Taxonomy

The great horned owl was formally described in 1788 by the German naturalist Johann Friedrich Gmelin in his revised and expanded edition of Carl Linnaeus's ''Systema Naturae''. He placed it with the other owls in the genus '' Strix'' and coined the binomial name ''Strix virginia''. Gmelin based his description on that of English naturalist George Edwards who had described and illustrated the great horned owl in 1747 in the second volume of his ''A Natural History of Uncommon Birds''. Edwards had seen a live specimen from Virginia at the house of the Earl of Burlington in Chiswick. Edwards also owned a preserved specimen, and another specimen formed part of the Leverian collection. The great horned owl is now placed in the genus '' Buco'' that was introduced in 1805 by André Duméril.

The great horned owl represents one of the one or two radiations of the genus across the Bering land bridge to the Americas. Whereas the Magellanic horned owl clearly divided once the owl had spread through the Americas, the consensus seems to be that the snowy owl and the great horned owl divided back in Eurasia and the snowy then spread back over the Arctic through northernmost North America separately from the radiation of the horned owl. The great horned and Eurasian eagle-owls may in fact be conspecifics, based on similarities in life history, geographic distribution, and appearance. In one case, a zoo-kept male great horned owl and female Eurasian eagle-owl produced an apparently healthy hybrid. Genetic testing indicates that the snowy owl, not the Eurasian eagle-owl, is the most closely related living species. Pleistocene era fossils have been found of ''Bubo'' owls in North America, which may either be distinct species or paleosubspecies, from as far east as Georgia, but predominantly in the Rocky Mountains and to the west of them. Almost all fossils indicate these owls were larger than modern great horned owls.

A large number of subspecies, more than 20 altogether, have been named. However, many of these are not true subspecies and only examples of individual or clinal variation. Subspecies differences are mainly in color and size and generally follow Gloger's and Bergmann's rules: The most conservative treatments of great horned owl subspecies may describe as few as 10, although an intermediate number is typical in most writings.

Fifteen subspecies are currently recognised:
* ''B. v. algistus'' ( Oberholser, 1904) – west Alaska
* ''B. v. lagophonus'' (Oberholser, 1904) – central Alaska to northeast Oregon, Idaho and northwest Montana (USA)
* ''B. v. saturatus'' Ridgway, 1877 – coastal southeast Alaska to coastal north California (USA)
* ''B. v. pacificus'' Cassin, 1854 – coastal central California (USA) to northwest Baja California (Mexico)
* ''B. v. subarcticus'' Hoy, PR, 1853 – central west Canada to north Idaho (USA)
* ''B. v. pallescens'' Stone, 1897 – southwest USA to south Mexico
* ''B. v. pinorum'' Dickerman & Johnson, AB, 2008 – south Idaho to north Arizona and north New Mexico (USA)
* ''B. v. heterocnemis'' (Oberholser, 1904) – northeast Canada to the Great Lakes region
* ''B. v. virginianus'' ( Gmelin, JF, 1788) – southeast Canada to central, east USA
* ''B. v. elachistus'' Brewster, 1902 – south Baja California (Mexico)
* ''B. v. mayensis'' Nelson, 1901 – Yucatán Peninsula (southeast Mexico)
* ''B. v. mesembrinus'' (Oberholser, 1904) – south Mexico to west Panama
* ''B. v. nigrescens'' Berlepsch, 1884 – Colombia to northwest Peru
* ''B. v. nacurutu'' ( Vieillot, 1817) – east Colombia through the Guianas to north, east Brazil, Argentina, Bolivia and central Peru
* ''B. v. deserti'' Reiser, 1905 – northeast Brazil (central north Bahia)

Description

The great horned owl is generally colored for camouflage. The underparts of the species are usually light with some brown horizontal barring; the upper parts and upper wings are generally a mottled brown usually bearing heavy, complex, darker markings. All subspecies are darkly barred to some extent along the sides, as well.

A variable-sized white patch is seen on the throat. The white throat may continue as a streak running down the middle of the breast even when the birds are not displaying, which in particularly pale individuals can widen at the belly into a large white area. South American great horned owls typically have a smaller white throat patch, often unseen unless actively displaying, and rarely display the white area on the chest. Individual and regional variations in overall color occur, with birds from the subarctic showing a washed-out, light-buff color, while those from the Pacific Coast of North America, Central America, and much of South America can be a dark brownish color overlaid with blackish blotching. The skin of the feet and legs, though almost entirely obscured by feathers, is black. Even tropical great horned owls have feathered legs and feet. The feathers on the feet of the great horned owl are the second-longest known in any owl (after the snowy owl). The bill is dark gunmetal-gray, as are the talons.

All great horned owls have a facial disc. This can be reddish, brown, or gray in color (depending on geographical and racial variation) and is demarked by a dark rim culminating in bold, blackish side brackets. This species' "horns" are tufts of feathers, called plumicorns. The purpose of plumicorns is not fully understood, but the hypothesis that they serve as a visual cue in territorial and sociosexual interactions with other owls is generally accepted.

Physiology and measurements

The great horned owl is the heaviest extant owl in Central and South America and is the second-heaviest owl in North America, after the closely related, but very different-looking snowy owl. It is heavily built, with a barrel-shaped body, a large head, and broad wings. Its size can vary considerably across its range, with populations in interior Alaska and Ontario being largest and populations in California and Texas being smallest, though those from the Yucatán Peninsula and Baja California appear to be even smaller. Adult great horned owls range in length from , with an average of , and possess a wingspan of , with an average of . Females are somewhat larger than males. Mean body weight is for females and for males. Depending on subspecies, maximum weight can reach ..

The wing chord length is . The wing loading, the measured wing area compared to weight, is high, meaning the wings are relatively small in surface area for the bird's weight; the species' wing loading has been described as proportionately the highest among raptors. The tail, being relatively short as is typical of most owls, is long. Like other owl species, the great horned owl is capable of “silent flight”, which is the way owls fly while making almost no discernable noise, despite their large size. This is made possible thanks to three main components of the owl's wing structure. The leading edge of their feathers have serrations that help to disrupt the turbulence generated by wing flapping, then the softer feathers help deaden the sound, and finally the trailing fringe of the feathers that works to finish cutting the sounds made by flight. The structure of the great horned owl wing also allows it to fly at a very low speeds for the size of the species, as slow as 2 miles per hour when they are gliding on breezes.

The legs, feet, and talons are large and powerful. Tarsal length is . The average foot span of a fully spread foot, from talon to talon, is around , as compared to in long-eared owls, in barn owls, and in the great grey owl. Great horned owls can apply at least 300 lb/in² of crushing power in their talons, a pressure considerably greater than the human hand is capable of exerting. In some big females, the gripping power of the great horned owl may be comparable to much larger raptor species such as the golden eagle.

The hard, inflexible bill of the great horned owl is long, although the culmen, the exposed bill portion as measured along the top of the beak, is only .

The outer ear openings, which are concealed by feathers on the sides of the head, are relatively smaller than those of the Eurasian eagle owl, being in vertical axis, with the left ear slightly larger than the right. Like most exclusively (or near exclusively) nocturnal species, the great horned owl has asymmetrical ear holes that allow for the triangulation of sounds when hunting in the dark. The different-height holes, while still close together, are differentiated enough that the owl is able to use the timing and direction of the sound waves hitting each hole to precisely locate prey even if the prey is located under cover such as snow. The disc-like shape of their faces also helps to direct the sounds they hear toward their ears. While the true nature/purpose of the ear tufts that are present on the great horned owl is unknown, researchers agree that the tufts do not play any role in the hearing ability of the owl. It is estimated that their hearing is up to ten times that of a human being.

The great horned owl's eyes, just slightly smaller than the eyes of a human being, are large even for an owl and rank proportionately among the largest eyes of all terrestrial vertebrates. The great horned owl has cylindrical eyes which creates more distance from the lens of the eye to the retina, which allows it to act more like a telephoto lens for farther distance sight compared to that possible from round eyes. They are visually highly adapted for nocturnal hunting and provide a wide, almost completely binocular field of view, a large corneal surface and a predominantly rod retina. The great horned owl's eye contains both rods and cones like most species that see in color, but the vision of a great horned owl closely resembles that of many other nocturnal species. The peak wavelengths that are observed by the cones is 555 nm and the research suggests that the great horned owl has relatively weak color vision, especially compared to other bird species. Despite (or perhaps as a result of) the poorer sense of color vision, the owl manages to have excellent night vision. Instead of turning its eyes, an owl must turn its whole head, and the great horned owl can rotate its neck 270°. The iris is yellow, except in the amber-eyed South American great horned owl (''B. v. nacurutu'').

Calls

The great horned owl's song is normally a low-pitched but loud (or also transcribed as , or , , ) and can last for four or five syllables. The call is resonant and has warranted descriptions as varied as "solemn" and "terrifying". The female's call is higher and rises in pitch at the end of the call. Female vocalizations are higher in pitch because of a smaller syrinx in the larger sex. Calling seems to peak after rather than before midnight. Usually, territorial hooting decreases in February or March at the onset of egg laying. On occasion, this species exhibits "an indescribable assemblage of hoots, chuckles, screeches, and squawks, given so rapidly and disconnectedly that the effect is both startling and amusing". Descriptions of some of these odd sounds including a growling note pair, a laughing , a high-pitched ; a weak, soft , a cat-like , a hawk-like note of , and a nighthawk-like . These vocalizations may be variously uttered when the birds are disturbed and angered at the nest (frequently preceding an attack on an interloping human or other animal), represent the vocal development of young owls, or are given during courtship and during territorial disputes with other owls. Young owls still in the care of their parents make loud, persistent hissing or loud, piercing screeching sounds that are often confused with the calls of the barn owl.

Subspecies

* Common/eastern great horned owl (''Bubo virginianus virginianus'')
:Eastern United States eastwards from at least as far west as Minnesota to Texas; northeastwards in distribution to southern Quebec, southern Ontario, Nova Scotia and Prince Edward Island. Resident all-year. The race ''B. v. mesembrinus'' from Central America south of the Isthmus of Tehuantepec, may be merely a southerly leg of this race, as its coloring is almost exactly the same as ''virginianus'', although with rather smaller body size. However, ''mesembrinus'' type owls are discontinuous in range, with only paler owls from ''pallescens'' and ''mayensis'' found in the in-between range before the northern ''virginianus'' reappears back up in South Texas.
:The "nominate race" is a medium-hued form, neither darkly saturated nor strikingly pale. Darker gray or somewhat paler individuals are known. It tends to be richly tinged with rufous and barred distinctly blackish-brown below with rather soft contrast. The feet can range from tawny to buff to creamy and are the legs are typically barred dark to a moderate extent. The facial disc is often a solid cinnamon-red color. This is mid-to-large race, with a wing chord length of , averaging , in males and , averaging , in females. Unexpectedly, although it is not the longest-winged, the nominate is the heaviest known race as males weigh from , averaging , and females weigh from , averaging ; the prior figures originally from a huge sample around Michigan. In comparison, ''B. v. subarcticus'', though averaging longer in wing length averages somewhat less heavy. On the contrary, in Costa Rica, within the range of the possibly synonymous ''mesembrinus'', great horned owls average approximately , the lightest average mass reported anywhere for this species. Other standard measurements of this race are a tail length of , a tarsus length of about and a bill length of . ''B. v. virginianus'' also tends to have relatively the longest ear tufts on any race.
* South American great horned owl (''Bubo virginianus nacurutu'')
:A lowland form occurring in disjunct populations from eastern Colombia to the Guyanas; also from Brazil and Uruguay south of the Amazon Basin to Bolivia, the Buenos Aires Province in northern Argentina and western Paraguay; resident all-year. Includes the proposed subspecies ''scotinus'', ''elutus'', and ''deserti''. The status of this form, especially the relationships between the scattered subpopulations and with ssp. ''nigrescens'' and the Magellanic horned owl, deserves more study.
:Dull, earthy brownish color is typical; birds from the semiarid interior of Brazil often have much white on uppertail- and ear-coverts against a dull gray background (sometimes separated as ''deserti''). This race is less fuscous than ''nigrescens''. It is the only subspecies where the iris is amber, not yellow. The Magellanic horned owl, while somewhat similar in coloring, has yellow eyes like other horned owls, not amber eyes. ''B. v. nacurutu'' is a medium-sized race, smaller than most in North America but not as small as some of the Mexican races. The wing chord length is in males and in females. The tail in both sexes can range from . Only three birds have had published weights, two males scaling and one female weighing . The most notable feature of this race is its large bill, at , which is the biggest of any horned owl race despite the otherwise moderate size of ''B. v. nacurutu''.
* Northern/sub-Arctic great horned owl (''Bubo virginianus subarcticus'')
:Breeding range from Mackenzie, British Columbia region east to the southern Hudson Bay; southern limit unclear but at least reaches to Montana and North Dakota. Non-breeding birds are regularly found south to latitude 45°S, i.e. Wyoming or South Dakota, occasionally ranging beyond this limit especially in years with low prey in the north. This race includes the birds described as ''occidentalis'' (based on a wintering individual, as was the original ''subarcticus'') and ''sclariventris''. The older name ''wapacuthu'' was occasionally used for this subspecies, but it cannot with certainty be assigned to a recognizable taxon and is thus considered a ''nomen dubium''. The population described as ''algistus'' is probably based on wandering individuals and/or various intergrades of ''subarcticus'' with other races.
:This is the palest form of horned owl, with the ground color essentially whitish with a faint buff tinge above; black underside barring variable from indistinct to pronounced, being most often prominent on the upper chest amongst otherwise pale plumage. This race shows little to no reddish coloration. ''B. v. subarcticus'' shows a very high degree of clinal variation, ranging from in the Contiguous United States where owls are often medium-grayish and more heavily marked to the subarctic zone in Canada where very pale birds with almost non-existent markings are prominent. Very pale birds are similar to a young female snowy owl from a distance. In this race, the feet range from immaculate white to buff, with little or no mottling. In the west of Canada, ''subarcticus'' may hybridize with the dark ''saturatus'' and may do the same with this similarly dark ''heterocnemis'' in the east. In both cases, they may produce intermediate looking hybrids of reddish tone, like a ''virginianus'' but with sharper contrasting of colors. This is one of the largest-bodied subspecies. The wing chord length is , averaging , in males and , averaging , in females. Body mass ranges from , averaging , in males and from , averaging , in females. Tail length is and in males and females, respectively. Bill length is in both and one bird had a tarsus of .
* California great horned owl (''Bubo virginianus pacificus'')
:Central and southern California west of the Sierra Nevada except for the San Joaquin Valley, south to Northwestern Baja California, Mexico. Intergrades with ''pallescens'' in San Diego County, California (see also below). Resident all-year.
:Very rich brown, dark underside barring distinct, less pronounced than in ''saturatus'' but more pronounced than in ''pallescens''. Humeral area is black. Feet are mottled dark. The facial disc is often even darkly mottled. This is a fairly small-bodied race, in fact including the lightest wild great horned owl adult ever weighed. The wing chord length is , averaging , in males and , averaging , in females. Body mass ranges from , averaging , in males and from , averaging , in females. Tail length is and in males and females, respectively. Bill length is and one bird had a tarsus of .
* Coastal great horned owl (''Bubo virginianus saturatus'')
:Pacific coast from southeastern Alaska to northern California. Includes the previously described form of ''B. v. leucomelas''. The often-recognized race ''B. v. lagophonus'' is often now considered a mere clinal variation of the same race from interior Alaska to Oregon, the Snake River, and northwestern Montana. These forms may be reported in winter as far south as Colorado and Texas, but these are in times of irruption.
:A dark and overall brownish form with heavily barred and mottled underside, with a dull tawny base. Inland birds (''lagophonus'') tend to have a more grayish base, the coastal owls being more richly brown. Otherwise, inland and coastal owls are practically the same. The facial disc can range from gray to reddish-gray to dark rufous. The feet are fairly dusky gray typically, although some buff-footed individuals are known, and legs are more prominently barred with black than in other North American races. This is a large race going on linear measurements, which in Alaskan owls outmatch those of all other races but for ''heterocnemis'' in Ontario (which may itself be a discontinuous eastern wing of this race). The wing chord length is , averaging , in males and , averaging , in females. Tail length is and in males and females. In both sexes, known bill and tarsal lengths are and . No weights are known to have been published.
* North Andean great horned owl (''Bubo virginianus nigrescens'')
: Andes; arid temperate and puna zones from Colombia to northwestern Peru. Resident all-year round. Presumably synonymous with the described ''columbianus''.
:A dark, cold gray-brown form with heavy fuscous blotching. Arguably this is the darkest colored race on average, though it could be rivaled by individuals from ''saturatus'' and ''elachistus''. This race has only minimally the rufous tinge seen in other darkish races, although some ''nigrescens'' may have a cinnamon facial disc. The largest winged race of owl in South America, this owl has a wing chord length of in males and in females. The tail in both sexes can range from . Bill length is , again relatively long as in ''nacurutu'' and one bird had a tarsal length of , indicating relatively long legs in the race. No published weights are known. Apparently, despite its sizeable wing area, ''nigrescens'' is notably smaller overall when specimens are compared side-by-side with those from ''saturatus''.
* Desert great horned owl (''Bubo virginianus pallescens'')
:San Joaquin Valley southeastwards through arid regions of southeastern California and southern Utah eastwards to western Kansas and southwards to Guerrero and western Veracruz in Mexico; intergrades with ''pacificus'' in San Diego County, if not elsewhere; vagrant individuals of ''saturatus'' and the Rocky Mountains population, which look similar to intergrades, also seem to occur in its range. Resident all-year. This race is likely synonymous with ''melanocercus''.
:A pale dusky buff form with indistinct barring, especially on the underside. Darker individuals tend to have a deeper fulvous base to their upper sides. Humeral area is umber in color and the feathers of the feet are white and usually unmarked. A small race, it averages slightly larger in wing length than ''pacificus'' but weighs less on average. The wing chord length is , averaging , in males and , averaging , in females. Body mass ranges from , averaging , in males and from , averaging , in females. In both sexes, tail length is and bill length is .
* Yucatán great horned owl (''Bubo virginianus mayensis'')
:Endemic to the southern two-thirds of the Yucatán Peninsula. Resident all-year.
:A medium-pale form, fairly similar to ''pallescens'' in both hue and ventral markings. Going on linear measurements, ''B. v. mayensis'' is smaller than all North American horned owls, even the smallish ''pallescens'', and is only slightly larger at median than the following race. This race has wing chord and tail lengths of and in males and and in females. In both sexes, the bill length is and tarsal length is .
* Baja California great horned owl (''Bubo virginianus elachistus'')
:Southern Baja California, Mexico. Resident all-year.
:Similar in color to ''pacificus'' but even darker and more heavily barred, like a miniaturized ''saturatus''. It is considerably (5–10%) smaller than ''pacificus'' linearly; some size overlap does occur though. On average, it is the smallest known subspecies. Males have a wing chord length of , tail length of and bill length of . A single female had a wing chord of and tail of .
* Northeastern great horned owl (''Bubo virginianus heterocnemis'')
:Breeds in eastern Canada (northern Quebec, Labrador, Newfoundland). Its southern breeding range seems to be delineated by the Saint Lawrence River. In winter, this race may disperses southwards throughout Ontario to as far as the Northeastern United States. This subspecies may be synonymous with ''saturatus'', although it is distributed far to the east of that race. ''B. v. heterocnemis'' is surrounded by the much paler ''subarcticus'' to its west and quite differently marked ''virginianus'' to its south, the latter two overlap and possibly hybridize in some of the northeast.
:A fairly dark and grey, heavily barred form. Feet pale with dusky mottling. Going on median reported linear measurements (since body mass is unknown), this is the largest bodied subspecies on average. Males have a wing chord length of and females range from . In both sexes, the tail is and the bill is .
* Rocky Mountains great horned owl (''Bubo virginianus pinorum'')
:The Rocky Mountains population breeds south of the Snake River in Idaho south to Arizona, New Mexico, and the Guadalupe Mountains of Texas. Westwards, it is presumed to occur to the Modoc Plateau and Mono Lake of California. This race was included in the presumed subspecies ''occidentalis'', but was recently first described and named as a distinct subspecies and makes up the missing piece in the once-muddled distribution of great horned owls in the West and Rockies. Downslope movements into valleys occupied by ''pallescens'' might occur, but this needs study.
:A medium gray form, intermediate in coloring between ''saturatus'' and ''pallescens''. Moderately barred and tinged buff or ochraceous on the underside. Feet mottled. This is a largish race, wing chord lengths being inexplicably greater in males, at , than in females, at . Tail length can range from and a female weighed .

Species identification

The combination of the species' bulk, prominent ear tufts and barred plumage distinguishes it through much of the range, but it may be easily confused with the lesser or Magellanic horned owl (''B. magellanicus''), which may overlap in range. The Magellanic horned owl was once considered a subspecies of the great horned, but is now almost universally considered a distinct species, as is supported by genetic materials, with the great horned being the paraspecies. Overall coloration is similar, but the Magellanic is markedly smaller with smaller feet and a smaller head, with finer, but more numerous brownish bars on the underside, rather than the blotchy, irregular barring typical of great horned owls. Other eagle-owls may superficially be somewhat similar, but the species is generically allopatric with the exception of wintering snowy owls. More tropical species with ear tufts, the stygian owl (''A. stygius'') and striped owl (''A. clamator''), are much smaller. Other large owls lack ear tufts.

Distribution and habitat

The breeding habitat of the great horned owl extends high into the subarctic of North America, where they are found up to the northwestern and southern Mackenzie Mountains, Keewatin, Ontario, northern Manitoba, Fort Chimo in Ungava, Okak, Newfoundland and Labrador, Anticosti Island and Prince Edward Island. They are distributed throughout most of North America and very spottily in Central America and then down into South America south to upland regions of Argentina, Bolivia and Peru, before they give way to the Magellanic horned owl, which thence ranges all the way to Tierra del Fuego, the southern tip of the continent. It is absent or rare from southern Guatemala, El Salvador, Nicaragua, and Costa Rica to Panama (where only two records) in Central America and the mangrove forests of northwestern South America. The species is also absent from the West Indies, the Haida Gwaii and almost all off-shore islands in the Americas, its ability to colonize islands apparently being considerably less than those of barn owls and short-eared owls. Since the division into two species, the great horned owl is the second most widely distributed owl in the Americas, just after the barn owl.

The great horned owl is among the world's most adaptable owls or even bird species in terms of habitat. The great horned owl can take up residence in trees that border all manner of deciduous, coniferous, and mixed forests, tropical rainforests, pampas, prairie, mountainous areas, deserts, subarctic tundra, rocky coasts, mangrove swamp forests, and some urban areas. It is less common in the more extreme areas of the Americas. In the Mojave and Sonora Deserts, they are absent from the heart of the deserts and are only found on the vegetated or rocky fringes. Even in North America, they are rare in landscapes including more than 70% old-growth forest, such as the aspen forest of the Rockies. They have only been recorded a handful of times in true rainforests such as the Amazon rainforest. In the Appalachian Mountains, they appear to use old-growth forest but in Arkansas are actually often found near temporary agricultural openings in the midst of large areas of woodland. Similarly in south-central Pennsylvania, the owls use cropland and pasture more than deciduous and total forest cover, indicating preference for fragmented landscapes. In prairies, grasslands and deserts, they can successfully live year round as long as there are rocky canyons, steep gullies and/or wooded coulees with shade-giving trees to provide them shelter and nesting sites.

In mountainous areas of North America, they are usually absent above the tree line, but great horned owls can be found up to in California and in the Rockies. In the Andean Mountains, on the other hand, they have adapted to being a true montane species, often found at least above sea level and are regularly recorded in treeless Puna grassland zones at in Ecuador and Peru. They are generally rare in non-tidal wetland habitat, and are replaced in the high Arctic tundra by snowy owls. They prefer areas where open habitats, which they often hunt in, and woods, where they tend to roost and nest, are juxtaposed. Thus lightly populated rural regions can be ideal. This species can occasionally be found in urban or suburban areas. However, they seem to prefer areas with less human activity and are most likely to be found in park-like settings in such developed areas, unlike eastern and western screech owls (''Megascops asio'' & ''M. kennicottii'') which may regularly occur in busy suburban settings. All mated great horned owls are permanent residents of their territories, but unmated and younger birds move freely in search of company and a territory, and leave regions with little food in winter.

Behavior

In most aspects of their behavior, great horned owls are typical of owls and most birds of prey. From experimentally raising young owls in captivity, Paul L. Errington felt that they were a bird of "essentially low intelligence" that could only hunt when partially wild and instinctually driven by hunger to hunt whatever they first encounter. He showed captive birds that were provided strips of meat from hatching, rather than having to hunt or to simulate hunting to obtain food, had no capacity to hunt. On the contrary, William J. Baerg compared behaviorally his captive-raised great horned owls to parrots, which are famously intelligent birds, although not as often playful "it knows its keeper and usually accepts whatever he wishes to do with a good deal of tolerance". Arthur C. Bent also noted the variability in temperaments of great horned owls to their handlers, some generally pleasant, though most are eventually aggressive. Most captive specimens, once mature, seem to resent attempts at contact and are often given to attacking their keepers. They will only follow cues when conditioned from an early age but rarely with the same level of success seen in some diurnal birds of prey trained for falconry or entertainment, although this does not necessarily correlate with intelligence as posited by Errington. Carl D. Marti also disagrees with Errington's assessments, noting that their prey selection is not as "completely random as Errington suggested"; while "Great Horned Owls appeared to select their mammalian prey in general relation to the prey populations...cottontails, appeared to be selected as prey out of relation to their population status."

Like most owls, the great horned owl makes great use of secrecy and stealth. Due to its natural-colored plumage, it is well camouflaged both while active at night and while roosting during the day. During the daytime it roosts usually in large trees (including snags & large hollows but usually thick branches) but may occasionally be in crevices or small caves in rocks or in dense shrubbery. Pine and other coniferous trees may be preferred where available since they are particularly dense and provide cover throughout the year. Typically, males have a favorite roosting site not far from the nest, sometimes used over successive years. While roosting, great horned owls may rest in the "tall-thin" position, where they sit as erect and hold themselves as slim as is possible. The kind of posture is well known as a further method of camouflage for other owls, like long-eared owls or great grey owls, especially if humans or other potential mammalian carnivores approach them. The Eurasian eagle owl rarely, if ever, assumes the tall-thin position. Outside of the nesting season, great horned owls may roost wherever their foraging path ends at dawn. Generally great horned owls are active at night, although in some areas they may be active in the late afternoon or early morning. At dusk, the owl utters a few calls before flying to a more open sing-post, i.e. large bare branch or large rocks to deliver song. Normally several perches are used to mark occupied territory or to attract a female. Despite its camouflage and cryptic locations, this species can still sometimes be spotted on its daytime roosts, especially by American crows (''Corvus brachyrhynchos''). Since owls are, next to red-tailed hawks, perhaps the main predator of crows and their young, crows sometimes congregate from considerable distances to mob owls and caw angrily at them for hours on end. When the owls try to fly off to avoid this harassment, they are often followed by the corvids.

Territoriality and movements

Typically, great horned owls are highly sedentary, often capable of utilizing a single territory throughout their mature lives. Although some species such as snowy owls, northern saw-whet owls, long-eared and short-eared owls are true migrants, most North American owls are not migratory and will generally show fidelity to a single territory year around. In great horned owls, mated pairs occupy territories year-round and long-term. Territories are established and maintained through hooting, with highest activity before egg-laying and second peak in autumn when juveniles disperse, and can range from an average of in Yukon to an average of in Wyoming.

Most territorial defense is performed by males, but females frequently assist their partners in hooting contests with neighbors or intruders, even during incubation. On occasion, although territory borders may be successfully maintained via vocalizations alone without even seeing the competing owl, such confrontations may turn physical, with various levels of threats distinguished. The highest threat level involves the spreading of wings, bill-clapping, hissing, higher-pitched screams of longer duration, with general body poised to strike with its feet at intruder. If the intruder continues to press the confrontation, the defending owl will "hop" forward and strike it with feet, attempting to grasp and rake with claws.

Territoriality appears to place a limit on the number of breeding pairs in a given area. Individuals prevented from establishing a territory live a silent existence as "floaters". Radio-telemetry revealed that such floaters concentrate along boundaries of established territories. At Kluane Lake in Yukon, incursions into neighboring territories were observed only twice—by females when a neighboring female had died or emigrated, suggesting that territorial defense may be sex-specific. At least four dead great horned owls in Kluane were apparently killed by others of their own species in territorial conflicts. Owls killed by other horned owls are sometimes cannibalized, although the origin of the killing may have been territorial aggression. Northern populations occasionally irrupt south during times of food shortage, but there is no annual migration even at the northern limits of the great horned owl's range.

Hunting behavior

Hunting tends to peak between 8:30 pm and midnight and then can resume from 4:30 am to sunrise. Hunting tends to be most prolonged during winter by virtue of prey being more scarce. However, great horned owls can learn to target certain prey during daylight in the afternoon when it is more vulnerable, such as eastern fox squirrels (''Sciurus niger'') while they're building their leaf nests and chuckawallas (''Sauromalus ater'') sunning themselves on desert rocks. Owls hunt mainly by watching from a snag, pole or other high perch. During hunting forays, they often fly about from perch to perch, stopping to survey for food at each, until they sense a prey item below. From such vantage points, owls dive down to the ground, often with wings folded, to ambush their prey. Effective maximum hunting distance of an owl from an elevated perch is . Due to their short but broad wings, great horned owls are ideally suited for low speed and maneuverability. Despite reports that they do not hunt on the wing, they also sometimes hunt by flying low over openings on the ground, scanning below for prey activity. Great horned owls can fly at speeds of more than in level flight. Hunting flights are slow, often quartering low above the ground where prey is likely to occur in open country or open woodland. Brief hovering flight (for about 6–18 seconds) has been described, especially in windy areas. On occasion owls may actually walk on the ground in pursuit of small prey or, rarely, inside a chicken coop to prey on the fowl within. Rodents and invertebrates may be caught on foot around the base of bushes, through grassy areas, and near culverts and other human structures in range and farm habitat. The great horned owl is generally a poor walker on the ground; it walks like a starling, with a pronounced side-to-side gait. They have been known to wade into shallow water for aquatic prey, although this has been only rarely reported. Owls can snatch birds and some arboreal mammals directly from tree branches in a glide as well. The stiff feathering of their wings allows owls to produce minimal sound in flight while hunting.

Almost all prey are killed by crushing with the owl's feet or by incidentally stabbing of the talons, though some may be bitten about the face as well. Prey is swallowed whole when possible. When prey is swallowed whole, owls regurgitate pellets of bone and other non-digestible bits about 6 to 10 hours later, usually in the same location where the prey was consumed. Great horned owl pellets are dark gray or brown in color and very large, long and thick, and have been known to contain skulls up to in width inside them. However, not all prey can be swallowed at once, and owls will also fly with prey to a perch and tear off pieces with their bill. Most dietary studies focus on pellets found under perches and around nests, since they provide a more complete picture of the diversity of prey consumed, but prey remains outside of pellets may provide clues to prey excluded from the pellets and a combination of both is recommended. Many large prey items are dismembered. Great horned owls may behead large prey before taking it to its nest or eating perch. The legs may also be removed, as may (in some bird prey) the wings. The great horned owl will also crush the bones of its prey to make it more compact for carrying. On occasion, the owls may return to the kill site to continue eating if the prey is too heavy to fly with after dismemberment. Many owls will accrue a cache of prey, especially those who are nesting. Caches must be at a safe location, usually the crotch of a tall tree. In northern regions, where large prey is prevalent, an owl may let uneaten food freeze and then thaw it out later using its own body heat. Hunting success seems to require fairly open understory, and experimental testing of microhabitat proved that open areas provided more hunting success on five species of rodent, with cloudy nights and denser bush foliage both decreasing success.

Prey and trophic ecology

Prey can vary greatly based on opportunity. According to one author, "Almost any living creature that walks, crawls, flies, or swims, except the large mammals, is the great horned owl's legitimate prey". In fact, the great horned owl has the most diverse prey profile of any raptor in the Americas. Over 500 species have been identified as great horned owl prey, with dozens more identified only to genus or general type (especially numerous invertebrates) and presumably several more unknown from their relatively little-studied populations in the Neotropics. Mammals (more than 200 species) and birds (nearly 300 species) make up the majority of their diet. Their diet in North America is made up of 87.6% mammals, 6.1% birds, 1.6% reptiles and amphibians with the remaining 4.7% being made up by insects, other assorted invertebrates and fish. Estimated mass of individual prey for the owls has ranged from as little as to as much as Most prey is in the range of (shrews) to (jackrabbits). A single owl requires about of food per day and can subsist on a large kill over several days. Despite the great diversity of prey taken by these predators, in most of the continental United States from the East to the Midwest as well as Canada and Alaska, great horned owls largely live off just a handful of prey species: three species of lagomorph: the eastern cottontail (''Sylvilagus floridanus''), the snowshoe hare (''Lepus americanus'') and the black-tailed jackrabbit (''Lepus californicus''); two species of New World mice: the white-footed mouse and the North American deermouse (''Peromyscus leucopus & maniculatus''); approximately three species of vole: the meadow, prairie and woodland voles (''Microtus pennsylvanicus, ochrogaster & pinetorum''); and one introduced pest, the brown rat.

Rodents

Small rodents form the great majority of great horned owl prey by number. Weighing a mere and on average, the nine species of New World mice in ''Peromyscus'' and eight species voles in ''Microtus'' recorded in the diet would appear to be overly small to be as important as they are to a predatory bird of this size. The prominence of these genera is undoubtedly due to the abundance of both genera in the wooded edge habitats frequented by great horned owls. It is estimated that a family of owls with two offspring would need to take about a half dozen (voles) to a dozen (mice) of these rodents every night to satisfy their dietary requirements but apparently the accessibility and abundance of these foods is irresistible as their numeric dominance is indisputable. By winter in areas that hold heavy snow, ''Peromyscus'' mice often come to outnumber the voles in the diet since the mice tend to travel over the surface of the snow while the voles make tunnels underneath the snow. In fact, a healthy family of great horned owls can decimate a colony of field rats, thus potentially performing a key role in controlling a highly destructive pest. Great horned owls living in the timbered fringes of garbage or refuse dumps may subsist mostly on rats.

In the Rockies, California, the Southwestern United States and northern Mexico, the diversity of this species' diet rises, in sync with the diversity of rodents. Especially important, from Colorado to Washington state is the northern pocket gopher (''Thomomys talpoides''), although assorted other pocket gophers (''Geomys, Cratogeomys, Zygogeomys, Pappogeomys'' and other '' Thomomys'' ssp.) are readily taken. While the northern weighs from , other pocket gophers hunted average from in mass. From Washington to Baja California a very important food is the pocket mice, primarily the Great Basin pocket mouse (''Perognathus parvus''). While the Great Basin species is a relative giant at , other hunted pocket mice (which may include both ''Perognathus'' and ''Chaetodipus'' ssp.) can average nearly as light as . In East Texas, the hispid cotton rat (''Sigmodon hispidus'') is the most commonly recorded prey species. The same species constituted 75% by number of a small sampling in Oklahoma. In semi-desert and other arid habitats, kangaroo rats become increasingly important prey, ten species have been reported in the diet but most prominently the Ord's and Merriam's kangaroo rats (''Dipodomys ordii & merriami''), both being widespread, numerous and relatively diminutive (at ). Eight known larger species of kangaroo rats, including the giant kangaroo rat (''Dipodomys ingens'') averaging at , are also taken.

The squirrels, including ground squirrels, marmots (''Marmota''), prairie dogs (''Cynomys''), chipmunks and tree squirrels, are diurnal and so are largely unavailable to great horned owls as prey. Occasionally though, one will be caught from their leaf nest, nest hole or burrow entrance first thing in the morning or in the late afternoon and approximately 35 species have been successfully predated by these owls. In general larger sized than other rodent families, the species hunting range from the gray-collared chipmunk (''Tamias cinereicollis'') to the hoary marmot (''Marmota caligata''); thus, squirrels can be provide a very fulfilling meal. An even larger rodent is sometimes attacked as prey by great horned owls, the North American porcupine (''Erethizon dorsatum''), in which average adults range from . This has been determined from owls who have porcupine quills imbedded in them, sometimes resulting in death. On occasion, they are successful in killing porcupine, even adults as determined by the size of the quills left behind and prey remains at bloodied kill sites. Other rodents recorded as secondary prey in North America include flying squirrels (''Glaucomys'' ssp.), the golden mouse (''Ochrotomys nuttalli''), red-backed voles & bog lemmings (''Myodes'' & ''Synaptomys'' ssp.), the muskrat (''Ondatra zibethicus''), the northern grasshopper mouse (''Onychomys leucogaster''), the northern pygmy mouse (''Baiomys taylori'') and jumping mice (''Zapus'' & ''Napaeozapus'' ssp.).

Lagomorphs

Although generally no match for rodents in sheer quantity of individuals, in terms of prey biomass, the most significant prey of North American great horned owls are hares and rabbits. About a dozen lagomorphs species are known to be hunted by the owl, from the relatively tiny pygmy rabbit to several hares weighing more than . Two hare species, the black-tailed jackrabbit and snowshoe hare, are so important to the owls as a food source that the local owl populations sharply rise and fall in sync with the hares' cyclical population trends. With adult weights of in adult cottontails, in snowshoe hares and in black-tailed jackrabbits, these species are overall the largest regular prey for this species. In Utah, where great horned owls are dependent on the jackrabbits, average brood size rose from 2 at jackrabbit population lows to 3.3 when the jackrabbits were at their peak. At the peak of population cycle, jackrabbits accounted for 90.2% and desert cottontails (''Sylvilagus audubonii'') for another 8.7% of prey biomass. In the short-grass prairie of Colorado, mountain cottontail (''Sylvilagus nuttallii'') and black-tailed jackrabbits predominated in October to December, making up 42.9% by number (and nearly all the biomass), thence dropping to 9.3% by number in April, while voles rose to 32.2% peak in May, down to a minimum of 10.2% by number in June. Further north in Colorado, in the absence of jackrabbits, the mountain cottontails falls to third place by number (12.9%) behind the northern pocket gopher (36.5%) and prairie vole (24.7%) but still dominates the biomass, making up about half. In central Utah, the lagomorphs (black-tailed jackrabbit/desert cottontail) and Ord's kangaroo rat each made up 39% of the food by number, respectively. The mountain cottontail dominates the biomass of prey in the Sierran foothills of California, making up 61.1% of the biomass, although are numerically secondary to desert woodrat. Remarkably, in the Morley Nelson Snake River Birds of Prey National Conservation Area of Idaho, individual rodents (1159 counted) were more than 10 times more numerous than lagomorphs (114 counted) by quantity and yet the jackrabbit and mountain cottontail still made up approximately half of the biomass. The dependence on lagomorphs also extends into Mexico, as in Baja California about a quarter of identified prey was black-tailed jackrabbit and either desert or the larger Mexican cottontail (''Sylvilagus cunicularius''). In the northern boreal forest, great horned owls are even more dependent on the snowshoe hare. At the peak of the 10 year hare cycle, snowshoe hares were by far the largest component of both summer and winter diets (77–81% and 90–99%, respectively, in Alberta; 83–86% and 75–98%, respectively, in Yukon). At the lowest point of the hare's cycle, summer diets consisted of only 0–16% snowshoe hare in Alberta and 12.7% in Yukon. When hares were scarce, great horned owls in these regions fed mostly on large rodents, mice and voles, grouse and ducks. Because fewer of these alternative prey species are available in boreal forest during winter, owls had to emigrate or suffer high mortalities if they stayed. In Alberta, the local population of great horned owls can increase threefold from hare population lows to peaks. The dependency on the snowshoe hare by the great horned owl extends into Alaska as well.

Other mammals

Other mammals are taken readily as well. From both the tropics and the United States, several species of opossum may be taken, down to the size of the tiny dwarf fat-tailed mouse opossum (''Thylamys velutinus''). In Brazil, white-eared opossum (''Didelphis albiventris'') were found in 12% of pellets, but all specimens appeared to be juveniles each weighing about . Quite differently, in Pennsylvania, Virginia opossum (''Didelphis virginianus'') made up 6% by number of prey but due to their large size (approximately ) and that all specimens were adults, they occupied the highest percentage of biomass of any species in a wide study from that state. At least eight species of shrews are taken by opportunity and make up the smallest mammalian prey taken by great horned owls, as specimens of least shrew (''Cryptotis parva'') or masked shrew (''Sorex cinereus'') have had an estimated weight of only . One of the more regularly taken shrews, though, is the larger northern short-tailed shrew (''Blarina brevicauda''), which was represented in more than 2% of pellets in the Upper Midwest. Moles, of at least four or five species, are also widely but lightly reported as prey. Remnants of armadillo, presumably nine-banded armadillo (''Dasypus novemcinctus''), have been found around owl nests in the south. 11 species of bat are known to be hunted by great horned owls. One pellet in Texas was found to be composed entirely of Mexican free-tailed bats (''Tadarida brasiliensis''). Smaller species of mammalian carnivore, such as ringtail (''Bassariscus astutus''), American mink (''Neogale vison''), black-footed ferret (''Mustela nigripes'') and various other small mustelids (''Mustela'' ssp.), are sometimes taken as prey. Prey in the form of canids, like foxes or coyotes (''Canis latrans'') are often juveniles presumably snatched from the mouths of dens by night. Kit and swift foxes of up to adult size may taken. Surprisingly, at least two cases of a great horned owl preying on an adult raccoon (''Procyon lotor'') have been reported. One instance of an owl taking a bobcat (''Lynx rufus'') as prey was also reportedly observed. In one case, a great horned owl was the likely killer of an adult female fisher (''Martes pennanti''), though young ones are typically taken. Occasionally, domestic carnivores are also prey. A few cases of young or small dogs (''Canis lupus familiaris'') and several of juvenile and adult cats (''Felis silvestris catus'') being killed by great horned owls have been reported. The most infamous predatory association amongst relatively larger carnivores is that with skunks. Due to their poor sense of smell, great horned owls are the only predators to routinely attack these bold mammals with impunity. All six skunk species found in North America are reported as prey, including adult striped skunks (''Mephitis mephitis''), which can be three times as heavy as the attacking owl. In one single nest, the remains of 57 striped skunks were found. Due to the proclivity of skunk predation, great horned owls nests frequently smell strongly of skunk and occasionally stink so powerfully of skunk that they leave the smell at kill sites or on prey remains.

Birds

After mammals, birds rank as the next most important general prey group. Birds are usually considerably secondary in the diet but outnumber the mammals in the diet by diversity, as more than 250 species have been killed in North America alone. Statistically, the most significant avian prey seems to be galliforms, of which they are known to have preyed on 23 species, basically consisting of all of the native species found in the United States. In the Upper Midwest, the ring-necked pheasant (''Phasianus colchicus'') and northern bobwhite (''Colinus virginianus'') were the fifth and sixth (out of 124 identified species) most significant prey species in 4838 pellets. Errington characterized the predatory pressure exerted on bobwhites by great horned owls as "light but continuous pressure", which may be considered characteristic of the species' hunting of all galliforms. Usually coveys of quail are partially protected by spending the night roosting communally in dense thickets but should a hunting owl be able to track down the communal roost, losses can be fairly heavy until the roost relocates. Similarly, owls may track down sleeping grouse, which also roost in vegetation but more openly than quail. Some grouse, such as greater prairie chicken (''Tympanuchus cupido''), sharp-tailed grouse (''Tympanuchus phasianellus'') and greater sage-grouse (''Centrocercus urophasianus''), may also been vulnerable to great horned owls while displaying conspicuously in openings on a lek first thing in the morning. In the boreal forest, especially in years where the snowshoe hare experiences population decreases, great horned owls prey fairly heavily (approximately 25% of biomass) on ruffed grouse (''Bonasa umbellus'') and spruce grouse (''Falcipennis canadensis''), enough so in the earlier bird to possibly contribute to population reductions. Larger species of galliform are not immune to predation either. On Protection Island in Washington state, introduced common peafowl (''Pavo cristatus'') are an important prey item. The wild turkey (''Meleagris gallopavo''), on average between the sexes, is probably the largest bird the great horned owl hunts in which they kill adults. Both full-grown wild turkeys and adult domestic turkeys have been hunted and killed. Under normal circumstances, domestic chicken (''Gallus gallus domesticus'') will be ignored in favor of wild prey. On occasion, individual owls, especially inexperienced juveniles, will become habitual fowl killers. These errant owls mainly hunt chickens, though will also take domestic guineafowl, turkeys and anything else available. In general, chickens kept in locked cages with enclosed tops overnight are safe from great horned owls; not so chickens left free range or in open enclosures.

While galliforms are widely reported, the few cases where great horned owls locally turn to birds as the primary food source over mammals, these may often be local responses to the abundance of breeding water birds or concentrations of roosting water birds, since they tend to roost in relatively open spots. They have been known to predate more than 110 different species of assorted water bird. In prairie wetlands of North Dakota, avian prey, primarily represented by ducks and the American coot (''Fulica americana'') came to represent 65% by number and 83% by biomass of the diet of the local owls, also including secondarily grebes, smaller rails and shorebirds as well as upland-based species like grey partridge (''Perdix perdix''), sharp-tailed grouse and passerines. 77% of the ducks in that study were juveniles, the largest duck being a male mallard (''Anas platyrhnychos'') weighing approximately , but nearly all the coots were adults. On Protection Island, Washington, where they are no native land mammals, rhinoceros auklets (''Cerorhinca monocerata''), both adults and nestlings, were the most numerous prey, present in 93% of 120 pellets. Species as large as adult Canada geese, snow goose and great blue herons have been successfully killed. The nestlings of even larger species like trumpeter swans (''Cygnus buccinator''), American white pelicans (''Pelecanus eryhtrorhynchos''), brown pelicans (''Pelecanus occidentalis'') and sandhill cranes (''Grus canadensis'') have also been killed by these owls.

Other assorted birds are taken seemingly at random opportunity. The predatory effect of this species on other raptorial birds, which is often considerable, is explored in the following section. In Brazil, it was found in a small study that birds overall outnumbered mammals in pellets, although most were not determined to species and the ones that were shown a tremendously diverse assemblage of birds with no obvious dietary preference. Although not usually numerically significant, 86 species of passerine have been taken by great horned owls. Members from most North American families are known as prey, although among smaller types such as chickadees, warblers, sparrows, cardinals, wrens and most tyrant flycatchers only a few species from each have been recorded. Nonetheless, an occasionally unlucky migrant or local breeder is sometimes snatched. Fledgling songbirds are regularly taken in spring and summer. The smallest avian prey known for great horned owls are the blue-grey gnatcatcher (''Polioptila caerulea'') and the ruby-crowned kinglet (''Regulus calendula''). Somewhat larger bodied families are more prominent, i.e. the corvids (14 species) and icterids (14 species) and, secondarily, the kingbirds (''Tyrannus'' ssp.), thrushes, mimids and European starling (''Sturnus vulgaris''). This is likely due to the fact that the larger passerines usually roost in relatively open spots and have larger, more conspicuous nests. Crows and ravens tend to be grabbed off of their communal roosts by night.

Other prey

The great horned owl rarely misses an opportunity to hunt reptiles and amphibians. However, lizards are largely unavailable as prey due to their typically diurnal periods of activity. However, some snakes are partially or largely nocturnal, and more than a dozen species are hunted in North America. Snakes hunted range from small, innocuous garter snakes (''Thamnophis'' ssp.) and night snakes (''Hypsiglena torquata'') to venomous species like cottonmouths (''Agkistrodon piscivorus'') and prairie rattlesnakes (''Crotalus virdis'') and formidable, large species like common king snakes (''Lampropeltis getula'') and black rat snake