Geographic Range
Commonly known as little owls,
Athene noctua
is found within 84 countries of the Palearctic region and the northeastern portion
of the Ethiopian region. The range of
Athene noctua
extends north to the 56th parallel and south to the southern border of the Palearctic
region in Asia. The southern extent includes the Middle East to southern Ethiopia
and south of western Europe to northern Niger. The eastern limit of the range borders
the Sea of Okhotsk and the western limit is found along the Atlantic Ocean.
- Biogeographic Regions
- palearctic
- ethiopian
Habitat
Athene noctua
inhabits a range that encompasses a wide variety of landscapes. This species is adapted
to mostly dry climates as rain decreases hunting success. Habitable landscapes also
vary in altitude as
Athene noctua
has been observed at elevations ranging from sea-level to 2600 m.
Athene noctua
is a cavity nesting species and may reside in tree and rock cavities, crevices in
cliffs and man-made structures, and even the nests, holes, and burrows of other animals.
It prefers habitats with open hunting grounds, many small prey, areas to perch, cavities
for nesting, and a stable climate. These habitat preferences of
Athene noctua
can be met by many landscapes and thus this species is found in a wide variety of
habitats, both natural and man-made. It resides in many natural, temperate regions
including: open fields, grasslands, open woodlands, steppes, semi-deserts, deserts,
cliffs, non-wooded mountains, and ravines, gorges, and gullies.
Athene noctua
also strongly associates with anthropogenic habitats including agricultural landscapes
(farmlands, meadows, pastures, orchards) and urban and suburban habitats (villages
and urban buildings).
- Habitat Regions
- temperate
- terrestrial
- Terrestrial Biomes
- desert or dune
- savanna or grassland
- chaparral
- mountains
- Other Habitat Features
- urban
- suburban
- agricultural
Physical Description
At hatching
Athene noctua
has pink skin, cere, and toes and a light white or gray bill and claws. The first
down is called neoptile and is short and white. After one week the second down, the
mesoptile, begins to grow and young little owls appear a mottled gray color. The mesoptile
resembles feathers but is shorter and softer. On the tenth day, the eyes open and
the iris is a light yellow color. For three or four weeks the neoptile is still attached
to the mesoptile. Overall the young little owls are grayer and show less contrast
in their coloring than adults. Even first year adults have differences in feather
shape and texture than older adults.
Little owl adults are mostly a dark brown color with cream markings over the entire
body. The cream markings include streaks on the forehead and crown, rings around the
eyes, a cream chin, spots and streaks on the breast and abdomen, round spots on the
back and tail, and a distinct V-shape mark on the back of the head. The males and
females share a similar appearance with males tending to have a lighter face. The
bill and iris of
A. noctua
are bright yellow and the eyelids are dark. The cere, a part of the bill, is grayish
to black in color. The legs are gray with a possible yellow tint. The toes are a darker
gray to brown or black color and the claws are dark brown to black. Some coloring
anomalies have been reported including partial albinism, leucistic coloration, or
a russet color.
The wing length of
A. noctua
has a reported maximum of 174 mm and minimum of 158 mm but normally ranges between
151 mm and 178 mm. The wingspan of male little owls is 557.67 mm (plus or minus 11.90
mm) and for females the wingspan was recorded as 565.64 mm (plus or minus 13.15 mm).
Females are generally larger than males and on average have a wing length that is
3 to 5 mm longer than males. Also, one year old little owls have a wing length that
is, on average, 3 mm shorter than older adults. The subspecies of
A. noctua
do differ in sizes with the largest little owls from Tibet and Kashmir mountains
and the smallest little owls from northeast Africa.
As the common name suggests, little owls are small with an average weight of 164 g.
Male little owls are 9 to 10% lighter than females. The minimum recorded weight of
an
A. noctua
is 98 g, the wild maximum is 270 to 280 g, and the maximum recorded in captivity
is 300 to 350 g.
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- female larger
Reproduction
The courtship period of
A. noctua
begins with the male defending his territory. He patrols the borders of his territory
and calls. The call serves as both a territorial defense to fend off other males and
to attract a mate. Once a potential mate makes contact, pair bonding begins. This
includes the little owls flying in pairs and sitting in the same tree. The female
begins to make a "begging" call to her partner and this prompts the male to feed her
prey he has captured. This feeding habit is important because it indicates that there
is enough food to reproduce and feed young and also to supply the female with more
food so she can gain weight for egg formation and incubation. The vocalization between
partners increases as pair bonding strengthens and the couple begins nest visiting
and/or copulation. The male selects potential nests within his territory and, around
the time of copulation, he shows the female the nest sites until she selects one.
Upon successful copulation, the female will remain in the nest and continue to "beg"
her mate for food at an increasing rate. After the young have fledged the bond between
the pair decreases and aggression increases. Often the partners are willing to separate
at this point but not all do. The little owls show a high rate of mate fidelity. The
partners will often fly together and hunt near each other. They also share body contact
while sleeping and preen and scratch each other. Scratching and copulation outside
of the breeding season has been reported and is believed to reduce aggression between
mates when not reproducing. Although
A. noctua
is monogamous, when little owl densities are high and prey is plentiful, sometimes
extra-pair copulations occur.
- Mating System
- monogamous
Athene noctua is iteroparous (produces a clutch every year). The mating season of A. noctua begins in early February and ends as late as May depending on the region. The time of egg-laying also varies with the location but can begin as early as March and end as late as August when taking into account replacement clutches. The clutch size is normally between 1 and 7, averaging 3.3, but as many as 12 has been recorded. If the first clutch is lost for some reason, a pair can lay a replacement clutch, usually of a smaller size. As soon as the eggs are laid, the incubation period begins. The incubation period is from 18 to 35 days, usually reported as 28 days. Overall hatching occurs approximately one month after laying and owlets are dependent upon the parents for two months. The average weight of newly hatched owlets is 10 to 12 g. The female remains with the nestlings for 16 days. The owlets begin to catch their own prey at 28 days and kill prey at 34 days. The offspring become fledglings as early as 28 days and as late as 35 days. The owlets become juveniles and no longer need parental care between August and the beginning of November. The juveniles settle an average of 2700 m away from their first nest. The little owls reach reproductive age at 1 year.
Hatching success of little owls is as low as 49.3%. Fledging success has been reported
as low as 46.6%. These losses can be due to infertility, adults losing or abandoning
the nest, predation, low prey availability, and other nest complications. The most
important trait linked to the production of young is food availability. The success
of young is effected by the distance of the nest to the hunting grounds and by weather
during the preceding winter, the breeding season, and after hatching. Supplementing
nests with food has been shown to greatly increase the production of young. The number
of young fledged per breeding pair ranges from 0.6 to 2.8 fledglings per pair.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- oviparous
Before copulation occurs, little owls must strengthen pair bonding. Part of this pair
bonding includes the male defending his territory and providing food to the female.
The female must develop fat reserves to help her produce eggs before the nest site
is selected. Before the female lays eggs, she remains in the nest and the male brings
prey to her. As soon as the eggs have been laid, the female begins incubating the
eggs to keep them warm. Pre-hatching parental investment also includes the male bringing
the female food as she incubates the eggs. The eggs hatch after about 28 days and
the pre-fledging parental care begins. The male still catches prey and brings it back
to the nest for the nestlings and female. The female defends, warms, and feeds her
young. After 16 days the female also leaves the nest and begins bringing prey back
for the nestlings. The offspring become fledglings as early as 28 days and as late
as 35 days. Pre-independence parental care includes the parents still feeding and
protecting the owlets one month after they reach fledgling age. Parental care ceases
at the beginning of September to the beginning of November. In general, there is a
two month post-hatching dependency period in which the parents are very active and
energetically stressed. As the brood ages, the activity rates of the parents increase.
The males are more active than the females during this period and are the main food
provider, possibly linked to their lighter weight allowing them to move further and
faster. Also the heavier weight of the females may help them care for and defend the
nest. Although the roles of the
A. noctua
parents are different, both provide provisioning and protection throughout the development
of the young.
- Parental Investment
- altricial
- male parental care
- female parental care
-
pre-fertilization
- provisioning
-
protecting
- male
- female
-
pre-hatching/birth
-
provisioning
- male
- female
-
protecting
- male
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- male
- female
-
protecting
- male
- female
-
provisioning
-
pre-independence
-
provisioning
- male
- female
-
protecting
- male
- female
-
provisioning
Lifespan/Longevity
The longest lifespan of
A. noctua
in the wild is 15 years and 7 months old, however the median lifespan of
A. noctua
is 4 years. The mortality rate of adult little owls is between 24.2% and 39% per
year. The mortality rate of juveniles is much higher, reported to be between 69% and
94%. The lifespan of little owls is limited mostly by prey and nest availability,
but predation, parasites, weather, and human influences (habitat destruction, traffic,
chemicals, electrocution, etc.) also effect longevity. Information on
A. noctua
lifespan is not well documented in both the wild and captivity.
Behavior
Athene noctua
is a sedentary species and remains close to its birthplace for life. Little owls
have been seen flying 20 to 30 m and even higher, but they usually fly low to the
ground. Their feathers are greased every two to three days. Little owls also take
dust or sand baths regularly and have even been spotted bathing in smoke rising from
chimneys. When
A. noctua
sleeps, it is able to position itself in a way that hides all light feathers and
camouflages itself extremely well in its habitat. Also, to close their eyes to sleep,
little owls raise their lower eyelids.
Athene noctua
preens often and thoroughly. Mutual preening has also been noted between mates and
even fledgling siblings.
Athene noctua
is crepuscular; it has been seen at daytime and nighttime hunting and calling, however,
the height of activity is at dusk and dawn.
- Key Behaviors
- arboreal
- flies
- crepuscular
- motile
- sedentary
- territorial
Home Range
The home ranges of little owls vary in size throughout the year from 1.8 to 95.0 ha
(.018 to .95 km^2). The average home range is 15.1 ha (.151 km^2). Through the winter
little owls must scavenge for scarce food. This contributes to the larger sizes of
home ranges and increases overlaps of territories among individuals. Home ranges decrease
and do not overlap during the breeding season. The decrease in home range is influenced
by an increase in energy expenditure during courtship, laying, hatching, and raising
of owlets. Little owls require more energy to defend and hunt a larger territory.
Also, the parents must hunt closer to the nest to reduce the amount of energy spent
on travel. The smallest home ranges were seen during the laying period, with an increase
during incubation and nestling periods. There is another decrease in home range during
the fledgling period of offspring. The second smallest home ranges were observed as
juveniles reached independence. It is believed parents decrease and defend their home
range to scare independent juveniles out of their natal territory. Owls that have
lost clutches or were unsuccessful in breeding will increase their home ranges and
often forage in the same areas as other owls. Although,
A. noctua
is territorial, there has been little or no aggression found between little owls
near each other. The average distance between little owl nests is 560 m.
Communication and Perception
Athene noctua is a very vocal bird with a repertoire of between 22 and 40 different calls. These calls are are used to contact or attract mates, defend territories against enemies and other little owls, alarm, and communicate. The vocalizations occur on average 1.87 times per hour and there is an average of 415 total seconds of calling in an hour. The vocal activity varies with time of year and day. There is usually a peak in calling before and during courtship and is lowest in the winter and summer. Also, there is an increase in call duration at dusk, called a dusk chorus phenomenon and a less pronounced increase at dawn with a lull through the night. There are more song vocalizations than calls at night. Also, little owls have been known to vocalize during daylight. The female call is generally shorter and higher pitched than the male call. The male calls are also louder and clearer than the female calls. When a predator is detected, A. noctua quiets and waits until the threat is presumably gone before resuming vocal activity. It has also been demonstrated that little owls learn to recognize the calls of neighboring little owls versus unknown little owls to reduce energy expenditure during territory defense.
Sight and hearing also allow little owls to perceive their environment. Little owls
have retinal cells more similar to diurnal birds and have the poorest visual acuity
compared to other birds of prey.
Athene noctua
can see yellow, green, blue, and (to a lesser extent) red. Little owls can also distinguish
between different shades of gray, increasing the ability to see at night. Little owls
can also hear sound frequencies of 3 to 4 kilocycles per second (kc/s). This ability
to hear allows
A. noctua
to locate rodents extremely accurately.
Little owl pairs often preen and scratch each other and sleep in contact with each
other. These tactile cues are used to increase pair bonding between mates. Mutual
preening has also been seen amongst fledgling siblings.
- Other Communication Modes
- choruses
Food Habits
Little owls hunt in various ways, even mixing the styles of hunting to suit their
prey. They can be found perching and watching the ground or the air. To catch prey
items on the ground,
A. noctua
will run and hop along the ground. They can also catch prey on the ground while flying
low.
Athene noctua
can catch prey out of the air while flying as well.
Athene noctua
carries its prey with either its beak or claws. Males and females hunt and carry
prey similarly, but females seem to prefer to carry prey by the neck, whereas males
carry prey more often by the head. Little owls are known to catch prey as heavy as
themselves. Little owls also create holes to hold surplus food. Hunting occurs mostly
at night or dusk, but they have been seen hunting in daytime.
Athene noctua
is an opportunistic carnivore that is known to feed on a wide variety of prey including
mammals
,
birds
,
reptiles
,
amphibians
, fish,
insects
,
molluscs
,
crustaceans
, and other invertebrate species. The diet mainly consists of small rodents and large
invertebrates (earthworms and insects). In the palearctic region,
A. noctua
is recorded to prey on 544 different species. The portions of the prey that little
owls cannot digest are compacted internally and regurgitated as a pellet.
- Primary Diet
-
carnivore
- eats terrestrial vertebrates
- Animal Foods
- birds
- mammals
- amphibians
- reptiles
- fish
- insects
- terrestrial non-insect arthropods
- mollusks
- terrestrial worms
- aquatic crustaceans
- Foraging Behavior
- stores or caches food
Predation
Little owls have many anti-predator adaptations.
Athene noctua
is known to call in response to predators as an alarm and defense. When other little
owls hear the defense call they will hide. When
A. noctua
sleeps, the white markings are hidden which allows the owl to camouflage. Also, the
V-shaped marking on the back of the head mimics eyes to prevent predation from behind.
Further, it has been noted that in the presence of barn owls (
Tyto alba
), little owls will hide or become silent until the threat is perceived to be gone.
Although not a normal predator of
A. noctua
, sometimes intraguild predation occurs between little owls and
Tyto alba
. The predators of little owls are larger birds including
tawny owls
,
eagle owls
,
long-eared owls
,
peregrine falcons
,
common buzzards
,
goshawks
,
sparrowhawks
,
red kites
,
tawny eagles
,
rough-legged buzzards
,
booted eagles
,
lanners
,
marsh harriers
,
black kites
,
imperial eagles
, and
long-legged buzzards
. Other predators include
pine martens
,
feral cats
,
fox
,
domestic dogs
, and
corvids
. The main predators of the nests are mammals, including
stone martens
and
common genets
. The eggs and chicks of little owls are predated upon by
stoats
,
hedgehogs
,
brown rats
,
corvids
, and
magpies
.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Little owls feed mainly on insects and small mammals and can keep these populations
under control. Also,
A. noctua
serves as the prey for other species. Although
A. noctua
interacts with many species, there are no mutualistic relationships documented. Two
commensal species of little owls are
Trichophagata petzella
and
Monopis laevigell
. There are at least 67 parasite species of little owls. The main types of parasites
are from the groups
Protozoa
,
Sporozoa
,
Metazoa
,
Nemathelminthes
,
Arachnida
, and
Insecta
. The blood parasite
Leucocytozoon ziemanni
and the ectoparasite dipteran
Carnus hemapterus
have been reported to use little owls as hosts. It could be stated that
A. noctua
is a commensal species of
humans
. The little owls use agricultural landscapes and urban and suburban areas as habitats.
Also, little owls have been seen following farming equipment that stirs up insects
and taking "smoke" baths in the rising smoke of chimneys.
Economic Importance for Humans: Positive
Little owls eat many insects and small mammals, controlling populations that are pests
to humans. There has been a dramatic decline in some owl populations throughout Europe,
including
A. noctua
.
Athene noctua
is perfect for studying owl declines in rural habitats because the species is well
known to the public, present in significant numbers, easy to research, and quickly
reacts to conservation actions. The presence of
A. noctua
attracts predators that were once desirable to humans. Due to this, little owls used
to be traded and carried on staffs to attract and catch their predators. Little owls
were also once traded as pets.
- Positive Impacts
- controls pest population
Economic Importance for Humans: Negative
Athene noctua
can be found nesting in and on the faces of buildings. This is considered unattractive
and measures have been taken in some places to avoid this (including nets hung over
signs). Other than this superficial damage,
Athene noctua
has no known adverse affects on humans.
Conservation Status
Athene noctua
is listed by the International Union for Conservation of Nature and Natural Resources
(IUCN) Red List as least concern. However, there is a documented decline seen throughout
Europe of not only little owls, but also other owls. It is important to research little
owls to gather information about the general decline of owls. The main contributors
to population decline are prey and nest availability and human habitat destruction.
The development of nest boxes appropriate for little owls has boosted numbers, while
current conservation efforts focus on increasing food availability during the breeding
season and gathering more information about the causes of decline.
Other Comments
Little owls have been associated with humanity for centuries.
Athene noctua
is seen as an omen of death, a sign of wisdom, and even a symbol of creators across
time and civilizations. Little owls can be found on coins, medallions, in literature,
and other works of art dating as far back as 8000 to 7500 BCE. The little owls were
traded to hunt small birds and even used as domesticated pets.
Athene noctua
was the bird of Athena, the Greek Goddess of Wisdom, and the origin of the genus
name
Athene
. Little owls can still be found in modern art today. Currently there are 12 recognized
subspecies of
Athene noctua
:
A. n. noctua
,
A. n. vidalii
,
A. n. indigena
,
A. n. glaux
,
A. n. saharae
,
A. n. lilith
,
A. n. bactriana
,
A. n. ludlowi
,
A. n. orientalis
,
A. n. plumipes
,
A. n. impasta
, and
A. n. spilogastra
.
Additional Links
Contributors
Kevin Hunt (author), Indiana University-Purdue University Fort Wayne, Mark Jordan (editor), Indiana University-Purdue University Fort Wayne, Rachelle Sterling (editor), Special Projects.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Ethiopian
-
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- terrestrial
-
Living on the ground.
- desert or dunes
-
in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- chaparral
-
Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- urban
-
living in cities and large towns, landscapes dominated by human structures and activity.
- suburban
-
living in residential areas on the outskirts of large cities or towns.
- agricultural
-
living in landscapes dominated by human agriculture.
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- monogamous
-
Having one mate at a time.
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- altricial
-
young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.
- male parental care
-
parental care is carried out by males
- female parental care
-
parental care is carried out by females
- arboreal
-
Referring to an animal that lives in trees; tree-climbing.
- crepuscular
-
active at dawn and dusk
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- choruses
-
to jointly display, usually with sounds, at the same time as two or more other individuals of the same or different species
- stores or caches food
-
places a food item in a special place to be eaten later. Also called "hoarding"
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- carnivore
-
an animal that mainly eats meat
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
References
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Hardouin, L., D. Robert, V. Bretagnolle. 2008. A dusk chorus effect in a nocturnal bird: support for mate and rival assessment functions. Behavioral Ecology and Sociobiology , 62: 1909-1918.
HolsegÄrd-Rasmussen, M., P. Sunde, K. Thorup, L. Jacobsen, N. Ottesen, S. Svenne, C. Rahbek. 2009. Variation in working effort in Danish little owls Athene noctua . Ardea , 97: 547-554.
Johnson, D., D. Van Nieuwenhuyse, J. Génot. 2009. Survey protocol for the little owl Athene noctua . Ardea , 97/4: 403-412.
Le Gouar, P., H. Schekkerman, H. van der Jeugd, A. Boele, R. van Harxen, P. Fuchs, P. Stroeken, A. van Noordwijk. 2010. Long-term trends in survival of a declining population: the case of the little owl ( Athene noctua ) in the Netherlands. Oecologia , 166(2): 369-379.
MĂLLER, A. 2006. Sociality, age at first reproduction and senscence: comparative analyses of birds. Jornal of Evolutionary Biology , 19: 682-689.
Thorup, K., P. Sunde, L. Jacobsen, C. Rahbek. 2010. Breeding season food limitation drives population decline of the little owl Athene noctua in Denmark. Ibis: The International Journal of Avian Science , 152(4): 803-814.
Tomé, R., P. Catry, C. Bloise, E. KorpimÀki. 2008. Breeding density and success, and diet composition of little owls Athene noctua in steppe-like habitats in Portugal. Ornis Fennica , 85: 22-32.
Van Nieuwenhuyse, D., J. Genot, D. Johnson. 2008. The Little Owl: Conservation, Ecology and Behavior of Athene noctua . Cambridge, New York: Cambridge University Press.
Zuberogoitia, I., J. MartĂnez, J. Zabala, J. MartĂnez, A. Azkona, I. Castillo, S. Hidalgo. 2008. Social interactions between two owl species sometimes associated with intraguild predation. Ardea , 96: 109-113.
Zuberogoitia, I., J. Zabala, J. MartĂnez, S. Hidalgo, J. MartĂnez, A. Azkona, I. Castillo. 2007. Seasonal dynamics in social behaviour and spacing patterns of the little owl Athene noctua . Ornis Fennica , 84: 173-180.
ƻmihorski, M., J. Romanowski, G. Osojca. 2009. Habitat preferences of a declining population of the little owl, Athene noctua in Central Poland. Folia Zoologica , 58: 207-215.