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zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and ...
, a crepuscular
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
is one that is active primarily during the
twilight Twilight is light produced by sunlight scattering in the upper atmosphere, when the Sun is below the horizon, which illuminates the lower atmosphere and the Earth's surface. The word twilight can also refer to the periods of time when this ...
period, being matutinal, vespertine, or both. This is distinguished from diurnal and
nocturnal Nocturnality is an ethology, animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatures generally have ...
behavior, where an animal is active during the hours of
daylight Daylight is the combination of all direct and indirect sunlight during the daytime. This includes direct sunlight, diffuse sky radiation, and (often) both of these reflected by Earth and terrestrial objects, like landforms and buildings. Sun ...
and of
darkness Darkness, the direct opposite of lightness, is defined as a lack of illumination, an absence of visible light, or a surface that absorbs light, such as black or brown. Human vision is unable to distinguish colors in conditions of very low ...
, respectively. Some crepuscular animals may also be active by
moonlight Moonlight consists of mostly sunlight (with little earthlight) reflected from the parts of the Moon's surface where the Sun's light strikes. Illumination The intensity of moonlight varies greatly depending on the lunar phase, but even the ...
or during an overcast day. Matutinal animals are active only before
sunrise Sunrise (or sunup) is the moment when the upper rim of the Sun appears on the horizon in the morning. The term can also refer to the entire process of the solar disk crossing the horizon and its accompanying atmospheric effects. Terminology ...
, and vespertine only after
sunset Sunset, also known as sundown, is the daily disappearance of the Sun below the horizon due to Earth's rotation. As viewed from everywhere on Earth (except the North and South poles), the equinox Sun sets due west at the moment of both the spr ...
. A number of factors impact the time of day an animal is active. Predators hunt when their prey is available, and prey try to avoid the times when their principal predators are at large. The temperature at midday may be too high or at night too low. Some creatures may adjust their activities depending on local competition.


Etymology and usage

The word ''crepuscular'' derives from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
'' crepusculum'' ("twilight"). Its sense accordingly differs from diurnal and
nocturnal Nocturnality is an ethology, animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatures generally have ...
behavior, which respectively peak during hours of daylight and darkness. The distinction is not absolute however, because crepuscular animals may also be active on a bright moonlit night or on a dull day. Some animals casually described as nocturnal are in fact crepuscular. Special classes of crepuscular behaviour include matutinal (or "matinal", animals active only in the dawn) and vespertine (only in the dusk). Those active during both times are said to have a bimodal activity pattern.


Adaptive relevance

The various patterns of activity are thought to be mainly
antipredator adaptation Anti-predator adaptations are mechanisms developed through evolution that assist prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by av ...
s, though some could equally well be predatory adaptations. Many predators
forage Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also us ...
most intensively at night, whereas others are active at midday and see best in full sun. Thus, the crepuscular habit may both reduce predation pressure, thereby increasing the crepuscular populations, and in consequence offer better foraging opportunities to predators that increasingly focus their attention on crepuscular prey until a new balance is struck. Such shifting states of balance are often found in ecology. Some predatory species adjust their habits in response to competition from other predators. For example, the subspecies of short-eared owl that lives on the
Galápagos Islands The Galápagos Islands ( es, Islas Galápagos) are an archipelago of volcanic islands in the Eastern Pacific, located around the Equator west of the mainland of South America. They form the Galápagos Province of the Republic of Ecuador, with ...
is normally active during the day, but on islands like Santa Cruz that are home to the Galapagos hawk, the owl is crepuscular. Apart from the relevance to predation, crepuscular activity in hot regions also may be the most effective way of avoiding heat stress while capitalizing on available light. Crepuscular flight activity is preferred by some animals, such as the walnut twig beetle, due to warmer temperatures, moderate wind speeds, and low barometric pressure. Crepuscular activity can be influenced by the lunar cycle due to the change in nocturnal light. This creates changes in animal
sleep Sleep is a sedentary state of mind and body. It is characterized by altered consciousness, relatively inhibited Perception, sensory activity, reduced muscle activity and reduced interactions with surroundings. It is distinguished from wakefuln ...
, reproduction, and foraging behaviours, often becoming less active during periods of low light.


Migration

Animal patterns of activity sometimes change during migration due to changes in environmental conditions.
Mule deer The mule deer (''Odocoileus hemionus'') is a deer indigenous to western North America; it is named for its ears, which are large like those of the mule. Two subspecies of mule deer are grouped into the black-tailed deer. Unlike the related whit ...
are crepuscular, but they are only active at
sunset Sunset, also known as sundown, is the daily disappearance of the Sun below the horizon due to Earth's rotation. As viewed from everywhere on Earth (except the North and South poles), the equinox Sun sets due west at the moment of both the spr ...
before and during migration. However, in the spring they are only active at
sunrise Sunrise (or sunup) is the moment when the upper rim of the Sun appears on the horizon in the morning. The term can also refer to the entire process of the solar disk crossing the horizon and its accompanying atmospheric effects. Terminology ...
because the
snow Snow comprises individual ice crystals that grow while suspended in the atmosphere An atmosphere () is a layer of gas or layers of gases that envelop a planet, and is held in place by the gravity of the planetary body. A planet ...
is at its hardest so it is easier for the deer to move without sinking in the snow. During migration, some types of swallow are active primarily during daytime hours with some activity during twilight hours.


Human impact on crepuscular behaviour

Crepuscular animal activity is affected by human activity. Crepuscular animals are less likely to participate in typical foraging or reproductive behaviors and deal with increased stress and mortality rates when humans are present. Animals may also change their usual activity patterns in response to the presence of humans. For example, Asian black bears may avoid areas with high human activity during the day, but go to these locations during twilight or nighttime hours. Light pollution impacts crepuscular behaviour because it mimics natural light conditions, leading crepuscular animals to behave as they would on nights with more moonlight.


Occurrence of crepuscular behaviour

Many familiar mammal species are crepuscular, including some bats, hamsters,
housecats The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
,
stray dog A free-ranging dog is a dog that is not confined to a yard or house. Free-ranging dogs include street dogs, village dogs, stray dogs, feral dogs, etc., and may be owned or unowned. The global dog population is estimated to be 900 million, of w ...
s,
rabbit Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit s ...
s, ferrets, and
rat Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include ''Neotoma'' ( pack rats), ''Bandicota'' (bandicoot ...
s. Other crepuscular mammals include jaguars, ocelots, bobcats, servals,
strepsirrhine Strepsirrhini or Strepsirhini (; ) is a Order (biology), suborder of primates that includes the Lemuriformes, lemuriform primates, which consist of the lemurs of Fauna of Madagascar, Madagascar, galagos ("bushbabies") and pottos from Fauna of A ...
s, red pandas,
bear Bears are carnivoran mammals of the family Ursidae. They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats throughout the N ...
s,
deer Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the ...
,
moose The moose (in North America) or elk (in Eurasia) (''Alces alces'') is a member of the New World deer subfamily and is the only species in the genus ''Alces''. It is the largest and heaviest extant species in the deer family. Most adult ma ...
, sitatunga, capybaras, chinchillas, the common
mouse A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
,
skunk Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or gi ...
s, squirrels,
fox Foxes are small to medium-sized, omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull, upright, triangular ears, a pointed, slightly upturned snout, and a long bushy tail (or ''brush''). Twelv ...
es,
wombat Wombats are short-legged, muscular quadrupedal marsupials that are native to Australia. They are about in length with small, stubby tails and weigh between . All three of the extant species are members of the family Vombatidae. They are ad ...
s, wallabies, quolls,
possums Possum may refer to: Animals * Phalangeriformes, or possums, any of a number of arboreal marsupial species native to Australia, New Guinea, and Sulawesi ** Common brushtail possum (''Trichosurus vulpecula''), a common possum in Australian urban ...
and marsupial gliders, tenrecs, and spotted hyenas. Snakes, lizards, and frogs, especially those in desert environments, may be crepuscular. Crepuscular birds include the
common nighthawk The common nighthawk (''Chordeiles minor'') is a medium-sized crepuscular or nocturnal bird of the Americas within the nightjar family, whose presence and identity are best revealed by its vocalization. Typically dark (gray, black and brown), d ...
, barn owl, owlet-nightjar, chimney swift,
American woodcock The American woodcock (''Scolopax minor''), sometimes colloquially referred to as the timberdoodle, the bogsucker, the hokumpoke, and the Labrador twister, is a small shorebird species found primarily in the eastern half of North America. Woodc ...
, spotted crake, white-breasted waterhen, European nightjars, and common buzzards. Many
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of ...
s,
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s,
flies Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...
, and other
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs ...
s are crepuscular and vespertine.


See also

*
Cathemeral Cathemerality, sometimes called metaturnality, is an organismal activity pattern of irregular intervals during the day or night in which food is acquired, socializing with other organisms occurs, and any other activities necessary for livelihood ar ...
* Crypsis * Diurnality *
Nocturnality Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sen ...


References

{{Light Ethology Ethology Night de:Temporale Spezialisten