Barn owls

(Tytonidae)

Class Aves

Order Strigiformes

Suborder Strigi

Family Tytonidae

Thumbnail description
Medium-sized owls with large head, dark eyes, heart-shaped facial disc, and with finely patterned plumage much like the wings of some moths

Size
9–22 in (23–57 cm)

Number of genera, species
2 genera; 17 species

Habitat
Most terrains, from forest to desert to urban areas, in cool temperate to tropical zones

Conservation status
Endangered: 3 species; Vulnerable: 1 species

Distribution
Almost worldwide

Evolution and systematics

Barn owls are represented in the fossil record back to the Paleocene (65 million years ago), along with at least four now-extinct owl families. The Tytonidae was most diverse through the Eocene, Oligocene, and Lower Miocene, with nine species in five genera, including two now-extinct subfamilies that shared characters of the extant genera Tyto and Phodilus. The highly diverse hawk owls (family Strigidae) arose by the Lower Miocene, and superseded the Tytonidae. The rise and diversification of owls have been linked to the tertiary radiation of small mammals.

Tyto owls appeared by the middle Miocene, and diversified through the Pliocene and Pleistocene. The now-extinct species were larger than modern Tyto, with some gigantic forms on islands. Some of these forms persisted until recent times (the Holocene Epoch). The common barn owl (Tyto alba) is known from the Pleistocene. The present center of diversity in the Australian region, with relict species in Africa, Madagascar, and Southeast Asia, suggests that the family originated in Gondwanaland.

Owls were formerly thought to be related to the diurnal birds of prey, order Falconiformes, but DNA comparisons have shown that they are instead related to the nightjars (Caprimulgiformes). Studies of anatomy, behavior, biochemistry, and genetics have shown that the barn owls, although closely related to the hawk owls (Strigidae), are distinct at the family level.

Barn owls, genus Tyto, and bay owls, genus Phodilus, are sufficiently different for separation into the subfamilies Tytoninae and Phodilinae. As of 2001, science recognizes 16 species in the genus Tyto, and one species in Phodilus, with a total of about 65 subspecies. Almost half of these are subspecies of the widespread common barn owl, which has many isolated populations on islands.

Physical characteristics

As noted by Iain Taylor in his book Barn Owls, published in 1994, these owls "when hunting over a meadow … have an ethereal quality that can be matched by no other bird." Barn owls are large-headed, short-tailed owls with a pale bill, and dark, frontally-set eyes in a flat, heart-shaped, prominently-rimmed facial disc. They range in size from 9–22 in (23–57 cm) and 0.4–2.8 lb (187–1,260 g). They are colored in browns and grays, often with a pale underside, and the dorsal plumage is intricately patterned in moth-like marbling, mottling, and flecking. The plumage is soft and fluffy, with flexible flight feathers in the large wings. Barn owls have a comb-like serration on the inner edge of the middle claw, absent in the Strigidae, but present in the nightjars. A nestling barn owl is characterized by its "long" face and bill, and emerging facial disc, and it develops extensive down that hides growing feathers. Other characteristics of barn owls which separate them from hawk owls include a tufted preen gland, primary feathers that are not narrowed at the tips or emarginated, an inner toe as long as the middle toe, and ear openings that are long slits covered by a flap of skin.

Distribution

Barn owls occur almost worldwide, being absent only from polar regions, the coldest parts of Eurasia and North America, and the driest Saharan and Middle Eastern deserts. The common barn owl is one of the most widely distributed land birds, being found on all continents except Antarctica as well

as on many islands, although only vagrants have reached New Zealand. One species occurs in Europe, the Mediterranean, the Middle East, North America, and South America; three occur in Africa; two in Madagascar and the Caribbean; four in Southeast Asia; and 11 in Australasia. There are two endemic species in Africa, one each in Madagascar and the Caribbean, two in Southeast Asia, and nine in Australasia. Their current range is little changed from their historical range, except that the common barn owl has disappeared from southern Scandinavia, Malta, and Aldabra, and the eastern grass owl (Tyto longimembris) may be extinct in Fiji. The Australian masked owl (Tyto novaehollandiae) has become locally extinct in some agricultural parts of southern Australia.

Habitat

Barn owls inhabit forested, wooded, and open habitats from the tropics to the cool temperate zones. They reach their greatest diversity in the tropics and subtropics, where several species coexist by partitioning the habitat. In Australia the sooty owl (Tyto tenebricosa) and lesser sooty owl (T. multi-punctata) inhabit rainforest or moist hardwood forest with a rainforest understory, and take prey in trees and shrubs as well as on the ground. The Australian masked owl inhabits drier, more open and grassy forest or woodland, or limestone cave systems in treeless areas, and takes mainly terrestrial prey. The common barn owl inhabits open woodland, grassland, and urban areas, and takes small terrestrial prey. The eastern grass owl specializes in aerial foraging for terrestrial prey in rank grassland. In Southeast Asia the Oriental bay owl (Phodilus badius) inhabits rainforest. Similarly, the Itombwe owl (Tyto prigoginei) and the Madagascar red owl (T. soumagnei) inhabit rainforest, alongside the common barn owl and African grass owl (T. capensis) of open habitats. Barn owls roost in cavities such as tree hollows and caves, or in dense foliage, and nest in hollows or in caves, except for the grass owls which roost and nest on the ground in thick cover.

Behavior

Barn owls occur singly, in pairs, or in family groups consisting of the pair and recently fledged, dependent young. In times of abundant prey, the smaller species of open habitats may hunt and roost in loose aggregations. Barn owls roost alone, except during the courtship phase of the breeding cycle, when the male may roost with the female in a cavity that becomes the nest. On rousing at dusk, and through the night, they advertise or maintain contact with screeching, screaming, or whistling calls, unlike the hooting of hawk owls. At close range, they communicate with quieter trilling or chattering calls.

If cornered by a perceived enemy, barn owls bluff in a threat display of puffed-up plumage, outspread wings, swaying on their perch, loud snapping of the bill, and hissing or screeching. If pressed, they lunge with the bill, strike with a foot, or eject malodorous feces. The larger barn owls of forest and woodland, such as the sooty owl, are strongly territorial and defend large, exclusive home ranges. The smaller species, such as the common barn owl, are less territorial and defend only a small area immediately about the nest. Most territorial defense is vocal, but defenders will approach an intruder with the threat display, or even chase, grapple, and fight.

Barn owls usually sleep during the day, upright on one leg with the facial disc pinched into a triangular shape, and the closed wings hunched forward to hide most of the pale underside. They hunt through the night, often from a series of perches, but smaller species also hunt on the wing. During times of food shortage, the small species of open country will hunt on dull afternoons. They return to their roosts at dawn, sometimes calling from or near the roost.

Breeding adult barn owls behave as sedentary pairs occupying permanent home ranges. Newly independent juveniles disperse, moving in search of food or vacancies in the breeding population. After prolific breeding in good times, dispersing individuals of the smaller, open-country species are irruptive, occupying marginal habitats in numbers while conditions are favorable, and even breeding there. There may be mass starvation and death when conditions deteriorate, followed eventually by a new boom-and-bust cycle. Most juveniles disperse to within 30–60 mi (50–100 km) of their birth place, although some common barn owls and grass owls disperse hundreds of miles (kilometers).

Feeding ecology and diet

Barn owls prey on a variety of vertebrates and invertebrates, mainly small mammals supplemented by birds, reptiles, amphibians, and large insects. They specialize on rodents and rodent-like mammals such as shrews or small marsupials. They forage by watching and listening from a series of perches or, in the smaller species of open country, by aerial searching and hovering. They swoop to seize prey in the claws and kill it with a bite to the neck before plucking and dismembering it or, if the prey is small, swallowing it whole. Later, at the roost, they disgorge pellets of fur or feathers that contain the skulls and other bones of their prey. When fresh, the dried pellets of barn owls have a characteristically dark, "glazed" mucous coating.

Barn owls are highly specialized for detecting prey by sound, enabling them to make captures in total darkness even when they cannot see their targets. Acute directional hearing enables them to pinpoint the sounds of mammals rustling and squeaking, to within 1–2° in the horizontal and vertical planes. The facial disc acts as a parabolic dish for focusing and amplifying sounds, and channeling them to the asymmetrical ear openings that locate the source of sounds by parallaxis. Muscles behind the ears move the facial ruff, as if the intently listening owl is cupping its ears. Owls also have frayed edges to the flight feathers for silent flight, thus avoiding detection by their prey.

Reproductive biology

Courtship among barn owls is vocal and sometimes aerial. Male and female sooty owls engage in prolonged duets of trilling calls. Male Australian masked owls and eastern grass owls perform a prolonged circling and chattering or trilling flight, respectively. Male common barn owls screech incessantly and perform a hovering flight near the nest, and pursue the female about the site with chattering calls. Males of the well-known species also bring food to the female, which begs like a juvenile. Mating, preceded by twittering calls, courtship feeding, and mutual preening, takes place at or near the nest, and culminates with a squealing call. In the better-known species, the mating system is permanent monogamy for the life of either partner.

Barn owl eggs are white and rounded oval, and take around five weeks to hatch. The clutch size is from one or two in the sooty owls to seven or eight in the common barn owl and

grass owls in good seasons. For most species the nest site is a natural or artificial cavity, such as a tree hole, but for the grass owls it is a tunnel and chamber within rank vegetation on the ground. Chicks hatch in sparse, pale down which is soon replaced by a thicker down. The growing feathers eventually show through as the down is shed, so that when fully fledged at around two months of age the owlets still have tufts of down on the head and thighs. By the time they leave the nest and fly, owlets are adult-sized but have shorter wings and tails because their flight feathers have not yet fully emerged from sheaths.

Incubation and brooding are undertaken solely by the female, which is fed on the nest by the male. The clutch is laid at two-day intervals and incubated from the start, so that eggs hatch sequentially and in a brood there is a range of ages. In times of food shortage, the youngest chicks may starve and be cannibalized. When the chicks are half-grown, feathering, and able to keep themselves warm, the female joins in providing for the family. Prey is at first dismembered and fed piecemeal to the chicks, but growing owlets are soon able to swallow whole surprisingly large items, such as rats. Fledged barn owls remain dependent on their parents for several weeks in small species to several months in large forest species, and can breed at one year of age.

In temperate regions barn owls lay in spring, and rarely manage more than one brood in a season. In warmer regions the laying season extends from autumn to spring, the onset of laying determined by food supply. In good years the smaller species of open habitats may raise two or three broods in succession, or even breed continuously during plagues of rodents.

Conservation status

Although barn owls are cryptic and under-recorded, several species such as the common barn owl, grass owls, sooty owl, and Australian masked owl are widespread and uncommon to locally common, or are common in a limited area (lesser sooty owl). Most are rarely encountered, endemic species with restricted ranges on small, tropical forested islands, particularly in Indonesia and Melanesia. Several are on the IUCN Red List. The Minahasa masked-owl (T. inexspectata) is listed as Vulnerable. The Madagascar red owl, the African bay owl (P. prigoginei), and the Taliabu masked owl (T. nigrobrunnea) are listed as Endangered. The lesser masked owl (T. sororcula) of the Moluccas, Sulawesi barn owl (T. rosenbergii), Andaman barn owl (T. deroepstorffi), and the ashy-faced owl (T. glaucops) of Caribbean islands, are also endemic species with highly restricted ranges. Two subspecies of the Australian masked owl were listed as Vulnerable on the Australian Environment Protection and Biodiversity Conservation Act 1999, but this situation was revised in 2000 to two subspecies classified as Near Threatened and two as Endangered.

Many members of the barn owl family have suffered population declines. The main reason is habitat loss: deforestation and logging in Australia and the tropics; conversion of natural grasslands to cultivation in the tropics and subtropics; and intensified agriculture in Western countries, with loss of foraging habitat and nest or roost sites. Rodenticides can also cause owl deaths. As of 2000, populations were estimated at 10,000 breeding individuals for the Australian subspecies of the sooty owl, a total of 21,300 for the Australian masked owl (all subspecies combined), and 10,000 for Australian populations of the eastern grass owl. In 1999, the European population of the common barn owl was estimated at 110,000–230,000 breeding pairs, meaning that its global population could be up to ten times that figure.

Significance to humans

Barn owls, by virtue of their nocturnal habits, ghostly appearance, eerie calls, and association with cemeteries and desolate places, have been featured in the mythology, literature, and art of all cultures since ancient times. As noted by Murray Bruce in Vol. 5 of the Handbook of the Birds of the World, "Few birds have accumulated such a wealth of varying and contradictory beliefs about them … [they are] feared or venerated, despised or admired, considered lucky or unlucky, and wise or foolish." They feature in superstitions concerning magic and witchcraft, prophecy, weather, birth, death, and other phenomena, and in potions, medicines, recipes, motifs, and the mummified contents of tombs. Common barn owls occasionally acquire a luminescent property to their plumage, which may contribute to the origin of legends concerning the "Will o' the wisp," "Jack o'lantern," and similar phenomena.

In modern times, barn owls have become popular as destroyers of rodent pests, and welcomed with the assistance of nest boxes in farm or town buildings. Common barn owls were introduced to some islands, with disastrous consequences for the endemic fauna. Barn owls are usually not dangerous to humans, although some individuals may defend owlets against human intruders at the nest by swooping and striking with their claws. The large species in Australia have become prominent in the debate over logging of old-growth forests.

Species accounts

List of Species

Sooty owl

Tyto tenebricosa

subfamily

Tytoninae

taxonomy

Strix tenebricosus John Gould, 1845, New South Wales. Two subspecies recognized (Tyto t. tenebricosa and Tyto t. arfaki). Lesser sooty owl (T. multipunctata), smaller, more spotted, and paler ventrally, is sometimes regarded as conspecific, or belonging to the same species.

other common names

English: Greater sooty or black owl, dusky barn owl; French: Effraie ombrée; German: Rußeule; Spanish: Lechuza Tenebrosa.

physical characteristics

Male 15–17 in (37–43 cm), 1.1–1.5 lb (500–700 g); female 17–20 in (44–51 cm), 1.9–2.5 lb (875–1150 g). Large, dark owl with large eyes set in rounded facial disc, massive feet. Sooty gray with fine white spots, pale belly. Female larger. Juvenile darker, with downy head and thighs when first fledged.

distribution

T. t. tenebricosa: coastal southeast Australia; T. t. arfaki: New Guinea

habitat

Tall, dense, moist forest including rainforest and eucalypt forest with a dense shrub layer of rainforest species. In montane

New Guinea extends above the tree line, into boulder fields in alpine grassland.

behavior

Nocturnal, solitary, secretive. Roosts in tree hollows, dense foliage, ravines, or caves. Sedentary and strongly territorial, advertising with a long, descending scream or wail. Defends home range of 1.5–3.9 mi² (4–10 km²).

feeding ecology and diet

Preys mostly on arboreal and terrestrial mammals such as rodents and possums, which it detects by watching and listening from perches.

reproductive biology

Monogamous. Laying recorded in most months, but usually autumn to spring. Nests in tree hollow or ledge in cave. Clutch one or two eggs, incubated for five or six weeks. Fledge at about three months.

conservation status

Not threatened. Widespread, but uncommon. Listed on Appendix II of CITES.

significance to humans

Prominent in the old-growth forest debate in Australia in the 1990s, as an indicator species for sustainable logging practices.

Australian masked owl

Tyto novaehollandiae

subfamily

Tytoninae

taxonomy

St[rix]? Novae Hollandiae Stephens, 1826, New South Wales. Five or six subspecies recognized; doubtfully valid T. n. galei usually merged with kimberli, and Tasmanian castanops sometimes regarded as full species. Forms species complex with other masked owls in Melanesia and east Indonesia: golden masked owl (T. aurantia) in Bismarck Archipelago, Manus masked owl (T. manusi) in Admiralty Island, lesser masked owl (T. sororcula) in southern Moluccan and Tanimbar islands, Taliabu masked owl (T. nigrobrunnea) in Sula Islands, and Minahasa masked owl (T. inexspectata) in northern Sulawesi.

other common names

English: Masked/tasmanian masked/chestnut-faced/cave owl; French: Effraie masquée; German: Neuhollandeule; Spanish: Lechuza Australiana.

physical characteristics

Male 13–17 in (33–42 cm), 0.9–1.8 lb (420–800 g); female 15–22 in (38–57 cm), 1.2–2.8 lb (545–1260 g). Large, pale or dark owl with large eyes set in rounded facial disc, large feet. Mottled brown and rufous (orange-brown to reddish brown) dorsally with rufous underside, or mottled gray and buff dorsally with white underside. Wings barred, underside finely spotted. Female larger. Juvenile has downy head and thighs when first fledged.

distribution

Two or three subspecies peripheral in continental north, northeast and south Australia; one subspecies in Tasmania; one subspecies on islands off northern Australia; and one subspecies in southern New Guinea.

habitat

Tall, grassy forest and woodland, often near open country, extending into treeless areas where there are caves.

behavior

Nocturnal, solitary, secretive. Roosts in tree hollows, dense foliage in gullies, or in caves. Sedentary. Territorial when breeding, advertising with a loud, harsh screech. Defends home range of 1.5–4.2 mi² (4–11 km²).

feeding ecology and diet

Preys mostly on terrestrial mammals, especially rodents, which it detects by watching and listening from perches.

reproductive biology

Monogamous. Laying recorded in most months, but usually autumn to spring. Nests in tree hollow. Clutch 1–4 eggs, usually two or three, incubated for 33–35 days. Fledge at two months.

conservation status

Widespread, but uncommon to rare. Listed on Appendix II of CITES. Two continental Australian subspecies are classified as Near Threatened and two island subspecies (Tasmania and Northern Territory) are Endangered under the Australian Environment Protection and Biodiversity Conservation Act.

significance to humans

Prominent in the old-growth forest debate in Australia in the 1990s, as an indicator species for sustainable logging practices.

Common barn owl

Tyto alba

subfamily

Tytoninae

taxonomy

Strix alba Scopoli, 1769, Italy. Twenty-seven to 30 subspecies recognized. Species complex includes ashy-faced owl (T. glaucops) of Caribbean islands, and Sulawesi barn owl (T. rosenbergii). In 1999, form on Andaman Islands elevated to full species rank as Andaman barn owl T. deroepstorffi. Some other dark, well-marked subspecies on tropical islands may also be full species.

other common names

English: Barn/cave/screech/white/ghost owl; French: Effraie des clochers; German: Schleiereule; Spanish: Lechuza Común.

physical characteristics

11–17 in (29–44 cm), 0.4–1.5 lb (187–700 g). Small, usually pale owl with small eyes set in heart-shaped facial disc, small feet. North American subspecies larger. Mottled gray and buff dorsally, with white to buff underside finely spotted. Some island forms darker, browner dorsally and rufous ventrally. Female similar. Juvenile has downy head and thighs when first fledged.

distribution

Five subspecies from Europe and Mediterranean to Middle East; one subspecies in sub-Saharan Africa and Madagascar; four subspecies on islands around Africa; one subspecies in India and Sri Lanka to Indochina and southern China; two subspecies from Malay Peninsula to eastern Indonesia; three

subspecies in Melanesia; one subspecies in Australia; one subspecies from North America to Caribbean; four subspecies on Caribbean islands; one subspecies on Galápagos; and four subspecies in Central and South America. Some disagreement on the number of subspecies, particularly in the Caribbean and South America, with some authorities also recognizing subspecies in Madagascar and Polynesia. Some disagreement over whether Caribbean forms insularis and nigrescens belong with T. alba or T. glaucops. Introduced to Seychelles and Hawaii.

habitat

Wide variety of open wooded or treeless, grassy or brushy habitats, extending to rocky areas, caves, wetland verges, and urban areas.

behavior

Nocturnal, crepuscular, and sometimes diurnal. Solitary, or in loose aggregations where prey is abundant. Roosts solitarily or communally in natural or artificial cavities or other shelter, from tree hollows, caves and dense foliage to buildings. Often resident, but also dispersive and irruptive. Breeding pairs are sedentary, occupying home ranges of 1.1–3.7 mi² (2.9–9.5km²), but defend an area of only a few yards around the nest. Call is a falsetto screech, sometimes likened to a woman screaming.

feeding ecology and diet

Preys on a wide variety of invertebrates and small vertebrates, but mostly on rodents, detected by watching and listening from perches or low quartering flight.

reproductive biology

Monogamous. Can lay in all months, and may have two or three broods in a year when prey is abundant, but at high latitudes lays in spring. Nests in a variety of natural and artificial cavities. Clutch usually 4–7 eggs; exceptionally up to 16 may involve two females laying in one nest. Incubation 29–34 days, fledging 7–10 weeks.

conservation status

Not globally threatened. Very widespread, and uncommon to locally common or even abundant during plagues of prey species. Some local declines in intensively farmed Western countries, and the status of some subspecies restricted to small islands is uncertain. Listed on Appendix II of CITES.

significance to humans

Has a long history of folklore in Europe, the Mediterranean, and among tribal peoples. Luminescing barn owls may have contributed to the origin of mythology concerning the Will o'the wisp, Jack o'lantern and similar phenomena. Has achieved popularity as a destroyer of rodents.

Eastern grass owl

Tyto longimembris

subfamily

Tytoninae

taxonomy

Strix longimembris Jerdon, 1839, India. Five or six subspecies recognized. Forms a species complex with the similar African grass owl (T. capensis), with which sometimes regarded as conspecific.

other common names

English: Grass owl; French: Effraie de prairie; German: Graseule; Spanish: Lechuza Patilarga.

physical characteristics

Male 13–14 in (32–36 cm), 0.6–0.8 lb (265–375 g); female 14–15 in (35–38 cm), 0.7–1.3 lb (320–582 g). Small, pale or dark owl with small eyes set in rather long, square facial disc, long bare legs. Mottled brown and buff dorsally, underside white or buff, finely spotted. Female larger and darker. Juvenile darker, with downy head and thighs when first fledged.

distribution

One subspecies from India through Indonesia to Australia; one subspecies in Indochina and eastern China; one subspecies in Taiwan; one subspecies in the Philippines; and one or two subspecies in New Guinea.

habitat

Rank grassland, heath, crops, and wetland verges of rushes and reeds.

behavior

Nocturnal and crepuscular, rarely diurnal. Solitary, or in loose aggregations when prey is abundant. Roosts, solitarily or communally, in dense ground cover. Resident, but also dispersive and irruptive. Territorial when breeding, defending the nesting area with trilling calls and soft screeches. Up to 40 pairs of grass owls may nest in 0.2 mi² (0.4 km², with as little as 55 yards (50 m) between neighboring nests.

feeding ecology and diet

Preys mostly on rodents, detected by watching and listening from low searching flight.

reproductive biology

Monogamous. Lays in autumn and winter. Nest is a scrape or pad on the ground, under covering vegetation. Clutch is 3–8 eggs, incubated for 31 days. Can run and leave the nest at 4–5 weeks, fully fledged and fly at two months.

conservation status

Not globally threatened. Widespread and uncommon to rare, although locally common in coastal eastern Australia during times of abundant prey. Listed on Appendix II of CITES.

significance to humans

In Australia has become a flagship species, as a destroyer of rodents in sugar cane.

Oriental bay owl

Phodilus badius

subfamily

Phodilinae

taxonomy

Strix badia Horsfield, 1821, Java. Six subspecies recognized. Formerly regarded as closely related to Itombwe owl or Congo bay owl (P. prigoginei), but the latter species more resembles Tyto and as of 1999 has been placed in that genus.

other common names

English: Bay owl, Asian bay owl; French: Phodile calong; German: Maskeneule; Spanish: Lechuza Cornuda.

physical characteristics

9–13 in (23–33 cm), 0.6–0.7 lb (255–308 g). Small chestnut owl with facial disc like barn owl, though squarer, divided down forehead and interrupted at throat, and peaked at corners (like small "ear" tufts). Barred and spotted dorsally, paler and spotted ventrally. Female and juvenile similar.

distribution

One subspecies in southwestern India; one subspecies in Sri Lanka; one subspecies from northeastern India through Indochina to southeastern China; and three subspecies in Malaysia and Indonesia. Subspecies uncertain for single record in Philippines.

habitat

Forest, and groves and plantations in farmland near forest.

behavior

Nocturnal and elusive, solitary. Roosts in tree hollows and dense foliage. Calls with a variety of mainly whistling and screeching notes.

feeding ecology and diet

Preys on small vertebrates and insects, detected by watching and listening from perches.

reproductive biology

Presumed monogamous. Lays in spring at higher latitudes, autumn-winter dry season in tropics. Nests in tree hollows, in crowns of palms, or occasionally in nest boxes. Clutch 3–5 eggs.

conservation status

Considered uncommon to rare, but probably under-recorded. Listed on Appendix II of CITES.

significance to humans

Owls are regarded as evil spirits by some tribal peoples in Southeast Asia.

Resources

Books

del Hoyo, J., A. Elliott, and J. Sargatal, eds. Barn Owls to Hummingbirds. Vol. 5, Handbook of the Birds of the World. Barcelona: Lynx Edicions, 1999.

Garnett, S., and G. Crowley. The Action Plan for Australian Birds 2000. Canberra: Environment Australia, 2000.

Higgins, P. J., ed. Parrots to Dollarbird. Vol. 4, Handbook of Australian, New Zealand and Antarctic Birds. Melbourne: Oxford University Press, 1999.

König, C., F. Weick, and J.-H. Becking. Owls, A Guide to the Owls of the World. Mountfield, United Kingdom: Pica Press, 1999.

Taylor, I. Barn Owls: Predator-Prey Relationships and Conservation. Cambridge: Cambridge University Press, 1994.

Periodicals

Bellocq, M. I. "A Review of the Trophic Ecology of the Barn Owl in Argentina." Journal of Raptor Research 34 (2000): 425–434.

Debus, S. J. S. "Surveys of the Barking Owl and Masked Owl on the North-west Slopes of New South Wales." Corella 25 (2001): 5–11.

Debus, S. J. S., L. R. Agnew, and M. Schulz. "Surveys of the Grass Owl Tyto capensis in Coastal New South Wales." Australian Bird Watcher 19 (2001): 94–102.

Durant, J. M., and Y. Handrich. "Growth and Food Requirement Flexibility in Captive Chicks of the European Barn Owl (Tyto alba)." Journal of Zoology (London) 245 (1998): 137–145.

Fajardo, I. "Monitoring Non-Natural Mortality in the Barn Owl (Tyto alba) as an Indicator of Land Use and Social Awareness." Biological Conservation 97 (2001): 143–150.

Marti, C. D. "Natal and Breeding Dispersal in Barn Owls." Journal of Raptor Research 33 (1999): 181–189.

Massemin, S., and T. Zorn. "Highway Mortality of Barn Owls in Northeastern France." Journal of Raptor Research 32 (1998): 229–232.

McCafferty, D. J., J. B. Moncrieff, and I. R. Taylor. "How Much Energy Do Barn Owls (Tyto alba) Save by Roosting?" Journal of Thermal Biology 26 (2001): 193–203.

Mlikovsky, J. "A New Barn Owl (Aves, Strigidae) from the Early Miocene of Germany, with Comments on the Fossil History of the Tytoninae." Journal für Ornithologie 139 (1998): 247–261.

Ramsden, D. J. "Effect of Barn Conversion on Local Populations of Barn Owl Tyto alba." Bird Study 45 (1998): 68–76.

Roulin, A. "Nonrandom Pairing by Male Barn Owls Tyto alba with Respect to a Female Plumage Trait." Behavioural Ecology 18 (1999): 688–695.

Roulin, A., M. Köllicker, and H. Richner. "Barn Owl (Tyto alba) Siblings Vocally Negotiate Resources." Proceedings of the Royal Society of London B267 (2000): 459–463.

Tome, R., and J. Valkama. "Seasonal Variation in the Abundance and Habitat Use of Barn Owls Tyto alba on Lowland Farmland." Ornis Fennica 78 (2001): 109–118.

Walk, J. W., T. L. Esker, and S. A. Simpson. "Continuous Nesting of Barn Owls in Illinois." Wilson Bulletin 111 (1999): 572–573.

Organizations

Australasian Raptor Association. 415 Riversdale Road, Hawthorn East, Victoria 3123 Australia. Phone: +61 3 98822622. Fax: +61 3 9882 2677. E-mail: mail@birdsaustralia.com.au Web site: <http://www.tasweb.com.au/ara/index.htm>

World Working Group on Birds of Prey and Owls. P.O. Box 52, Towcester, NN12 7ZW United Kingdom. Phone: +44 1 604 862 331. Fax: +44 1 604 862 331. E-mail: WWGBP@aol.com Web site: <http://www.Raptors-International.de>

Stephen Debus, MSc