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In
zoology Zoology ( , ) is the scientific study of animals. Its studies include the anatomy, structure, embryology, Biological classification, classification, Ethology, habits, and distribution of all animals, both living and extinction, extinct, and ...
, a crepuscular
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
is one that is active primarily during the
twilight Twilight is daylight illumination produced by diffuse sky radiation when the Sun is below the horizon as sunlight from the upper atmosphere is scattered in a way that illuminates both the Earth's lower atmosphere and also the Earth's surf ...
period, being
matutinal Matutinal, matinal (in entomology, entomological writings), and matutine are terms used in the list of life sciences, life sciences to indicate something of, relating to, or occurring in the early morning. The term may describe the morning activ ...
(active during dawn), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnal and
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
behavior, where an animal is active during the hours of
daytime Daytime or day as observed on Earth is the period of the day during which a given location experiences Daylight, natural illumination from direct sunlight. Daytime occurs when the Sun appears above the local horizon, that is, anywhere on the ...
and of
night Night, or nighttime, is the period of darkness when the Sun is below the horizon. Sunlight illuminates one side of the Earth, leaving the other in darkness. The opposite of nighttime is daytime. Earth's rotation causes the appearance of ...
, 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. History The ancient Greek philosopher Anaxagoras was aware that "''the sun provides the moon with its ...
or during an
overcast Overcast or overcast weather, as defined by the World Meteorological Organization, is the meteorological condition of clouds obscuring at least 95% of the sky. However, the total cloud cover must not be entirely due to obscuring phenomena near ...
day.
Matutinal Matutinal, matinal (in entomology, entomological writings), and matutine are terms used in the list of life sciences, life sciences to indicate something of, relating to, or occurring in the early morning. The term may describe the morning activ ...
animals are active only after
dawn Dawn is the time that marks the beginning of twilight before sunrise. It is recognized by the diffuse sky radiation, appearance of indirect sunlight being Rayleigh scattering, scattered in Earth's atmosphere, when the centre of the Sun's disc ha ...
, and vespertine only before
dusk Dusk occurs at the darkest stage of twilight, or at the very end of astronomical twilight after sunset and just before nightfall.''The Random House College Dictionary'', "dusk". At predusk, during early to intermediate stages of twilight, enoug ...
. A number of factors affect the time of day an animal is active.
Predators Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
hunt when their prey is available, and prey try to avoid the times when their principal predators are at large. The temperature may be too high at midday or too low at night. 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 languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
'' crepusculum'' ("twilight"). Its sense accordingly differs from diurnal and nocturnal behavior, which respectively peak during hours of daytime and night. The distinction is not absolute, 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 Predation, prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, na ...
s, though some could equally well be predatory
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
s. 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 used m ...
most intensively at night, whereas others are active at midday and see best in full sun. The crepuscular habit may both reduce predation pressure, increasing the crepuscular populations, and 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 The short-eared owl (''Asio flammeus'') is a widespread grassland species in the family Strigidae. Owls belonging to genus ''Asio'' are known as the eared owls, as they have tufts of feathers resembling mammalian ears. These "ear" tufts may or ...
that lives on the
Galápagos Islands The Galápagos Islands () 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 a population of sli ...
is normally active during the day, but on islands like Santa Cruz that are home to the
Galapagos hawk The Galápagos hawk (''Buteo galapagoensis'') is a large hawk endemic to most of the Galápagos Islands. Description The Galapágos hawk is similar in size to the red-tailed hawk (''Buteo jamaicensis'') and the Swainson's hawk (''Buteo swains ...
, 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 ''Pityophthorus juglandis'', also known as the walnut twig beetle, is a weevil that feeds on several different species of walnut trees (''Juglans''). It is one of only a few species in the genus ''Pityophthorus'' that is associated with hardwoods ...
, due to warmer temperatures, moderate wind speeds, and low barometric pressure. Crepuscular activity can be influenced by the
lunar cycle A lunar phase or Moon phase is the apparent shape of the Moon's directly sunlit portion as viewed from the Earth. Because the Moon is tidally locked with the Earth, the same hemisphere is always facing the Earth. In common usage, the four majo ...
due to the change in nocturnal light. This creates changes in animal
sleep Sleep is a state of reduced mental and physical activity in which consciousness is altered and certain Sensory nervous system, sensory activity is inhibited. During sleep, there is a marked decrease in muscle activity and interactions with th ...
,
reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...
, and
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
behaviours, often becoming less active during periods of low light.


Migration

Animal patterns of activity sometimes change during
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
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
dusk Dusk occurs at the darkest stage of twilight, or at the very end of astronomical twilight after sunset and just before nightfall.''The Random House College Dictionary'', "dusk". At predusk, during early to intermediate stages of twilight, enoug ...
before and during migration. In the spring they are only active at
dawn Dawn is the time that marks the beginning of twilight before sunrise. It is recognized by the diffuse sky radiation, appearance of indirect sunlight being Rayleigh scattering, scattered in Earth's atmosphere, when the centre of the Sun's disc ha ...
because the
snow Snow consists of individual ice crystals that grow while suspended in the atmosphere—usually within clouds—and then fall, accumulating on the ground where they undergo further changes. It consists of frozen crystalline water througho ...
is at its hardest, so it is easier for the deer to move without sinking in the snow. During migration, some types of
swallow The swallows, martins, and saw-wings, or Hirundinidae are a family of passerine songbirds found around the world on all continents, including occasionally in Antarctica. Highly adapted to aerial feeding, they have a distinctive appearance. The ...
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 Human behavior is the potential and expressed capacity ( mentally, physically, and socially) of human individuals or groups to respond to internal and external stimuli throughout their life. Behavior is driven by genetic and environmental fa ...
, because humans are diurnal. Crepuscular animals are less likely to participate in typical foraging or reproductive behaviors and deal with increased stress and
mortality rate Mortality rate, or death rate, is a measure of the number of deaths (in general, or due to a specific cause) in a particular Statistical population, population, scaled to the size of that population, per unit of time. Mortality rate is typically ...
s when humans are present. Animals may change their usual activity patterns in response to the presence of humans. For example,
Asian black bear The Asian black bear (''Ursus thibetanus''), also known as the Asiatic black bear, moon bear and white-chested bear, is a medium-sized bear species native to Asia that is largely adapted to an arboreal lifestyle. It is distributed from southeast ...
s may avoid areas with high human activity during the day, but go to these locations during twilight or nighttime hours.
Light pollution Light pollution is the presence of any unwanted, inappropriate, or excessive artificial Visible spectrum, lighting. In a descriptive sense, the term ''light pollution'' refers to the effects of any poorly implemented lighting sources, during the ...
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 A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
species are crepuscular, including the endangered
Amazon river dolphin The Amazon river dolphin (''Inia geoffrensis''), also known as the boto, bufeo or pink river dolphin, is a species of toothed whale Endemism, endemic to South America and is classified in the family Iniidae. Three subspecies are currently recogni ...
, some species of
bat Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...
s,
hamster Hamsters are rodents (order Rodentia) belonging to the subfamily Cricetinae, which contains 19 species classified in seven genera. They have become established as popular small pets. The best-known species of hamster is the golden or Syrian ...
s, housecats,
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 wh ...
s,
rabbit Rabbits are small mammals in the family Leporidae (which also includes the hares), which is in the order Lagomorpha (which also includes pikas). They are familiar throughout the world as a small herbivore, a prey animal, a domesticated ...
s,
ferret The ferret (''Mustela furo'') is a small, domesticated species belonging to the family Mustelidae. The ferret is most likely a domesticated form of the wild European polecat (''Mustela putorius''), as evidenced by the ferret's ability to inter ...
s,
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'' (bandicoo ...
s,
jaguar The jaguar (''Panthera onca'') is a large felidae, cat species and the only extant taxon, living member of the genus ''Panthera'' that is native to the Americas. With a body length of up to and a weight of up to , it is the biggest cat spe ...
s,
ocelot The ocelot (''Leopardus pardalis'') is a medium-sized spotted Felidae, wild cat that reaches at the shoulders and weighs between on average. It is native to the southwestern United States, Mexico, Central America, Central and South America, ...
s,
bobcat The bobcat (''Lynx rufus''), also known as the wildcat, bay lynx, or red lynx, is one of the four extant species within the medium-sized wild cat genus '' Lynx''. Native to North America, it ranges from southern Canada through most of the c ...
s,
serval The serval (''Leptailurus serval'') is a wild small cat native to Africa. It is widespread in sub-Saharan countries, where it inhabits grasslands, wetlands, moorlands and bamboo thickets. Across its range, it occurs in protected areas, and ...
s,
strepsirrhine Strepsirrhini or Strepsirhini (; ) is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and Southeast Asia. Colle ...
s,
red panda The red panda (''Ailurus fulgens''), also known as the lesser panda, is a small mammal native to the eastern Himalayas and southwestern China. It has dense reddish-brown fur with a black belly and legs, white-lined ears, a mostly white muzz ...
s,
bear Bears are carnivoran mammals of the family (biology), 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 ...
s,
deer A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) ...
,
moose The moose (: 'moose'; used in North America) or elk (: 'elk' or 'elks'; used in Eurasia) (''Alces alces'') is the world's tallest, largest and heaviest extant species of deer and the only species in the genus ''Alces''. It is also the tal ...
,
sitatunga The sitatunga (''Tragelaphus spekii'') or marshbuck is a swamp-dwelling medium-sized antelope found throughout central Africa, centering on the Democratic Republic of the Congo, the Republic of the Congo, Cameroon, parts of South Sudan, Southern ...
,
capybara The capybara or greater capybara (''Hydrochoerus hydrochaeris'') is the largest living rodent, native to South America. It is a member of the genus '' Hydrochoerus''. The only other extant member is the lesser capybara (''Hydrochoerus isthmi ...
s,
chinchilla Chinchilla refers to either of two species ('' Chinchilla chinchilla'' and '' Chinchilla lanigera'') of crepuscular rodents of the parvorder Caviomorpha, and are native to the Andes mountains in South America. They live in colonies called "her ...
s, 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,
squirrel Squirrels are members of the family Sciuridae (), a family that includes small or medium-sized rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrel ...
s,
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 ("brush"). Twelve species ...
es,
wombat Wombats are short-legged, muscular quadrupedal marsupials of the family Vombatidae that are native to Australia. Living species are about in length with small, stubby tails and weigh between . They are adaptable and habitat tolerant, and are ...
s,
wallabies A wallaby () is a small or middle-sized macropod native to Australia and New Guinea, with introduced populations in New Zealand, Hawaii, the United Kingdom and other countries. They belong to the same taxonomic family as kangaroos and som ...
,
possums Possum may refer to: Animals * Didelphimorphia, or (o)possums, an order of marsupials native to the Americas ** Didelphis, a genus of marsupials within Didelphimorphia *** Common opossum, native to Central and South America *** Virginia opossum, ...
and marsupial gliders,
tenrec A tenrec () is a mammal belonging to any species within the afrotherian family Tenrecidae, which is endemic to Madagascar. Tenrecs are a very diverse group, as a result of adaptive radiation, and exhibit convergent evolution, some resemble hed ...
s, and
spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
s.
Snakes Snakes are elongated Limbless vertebrate, limbless reptiles of the suborder Serpentes (). Cladistically Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales much like other members of ...
,
lizards Lizard is the common name used for all squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The ...
, and
frog A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order (biology), order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough ski ...
s, especially those in desert environments, may be crepuscular. Crepuscular birds include the
common nighthawk The common nighthawk or bullbat (''Chordeiles minor'') is a medium-sized crepuscular or nocturnal bird of the Americas within the nightjar (Caprimulgidae) family, whose presence and identity are best revealed by its vocalization. Typically dark ...
,
barn owl The barn owls, owls in the genus '' Tyto'', are the most widely distributed genus of owls in the world. They are medium-sized owls with large heads and characteristic heart-shaped faces. They have long, strong legs with powerful talons. The ter ...
,
owlet-nightjar Owlet-nightjars are small crepuscular birds related to the nightjars and frogmouths. Most are native to New Guinea, but some species extend to Australia, the Moluccas, and New Caledonia. A flightless species from New Zealand is extinct. There is ...
,
chimney swift The chimney swift (''Chaetura pelagica'') is a bird belonging to the swift family Apodidae. A member of the genus ''Chaetura'', it is closely related to both Vaux's swift and Chapman's swift; in the past, the three were sometimes considered to ...
,
American woodcock The American woodcock (''Scolopax minor''), sometimes colloquially referred to as the timberdoodle, mudbat, bogsucker, night partridge, or Labrador twister is a small shorebird species found primarily in the eastern half of North America. Woodco ...
, spotted crake,
white-breasted waterhen The white-breasted waterhen (''Amaurornis phoenicurus'') is a waterbird of the rail and crake family, Rallidae, that is widely distributed across South and Southeast Asia. They are dark slaty birds with a clean white face, breast and belly. The ...
,
European nightjar The European nightjar (''Caprimulgus europaeus''), common goatsucker, Eurasian nightjar or just nightjar is a crepuscular and nocturnal bird in the nightjar family that breeds across most of Europe and the Palearctic to Mongolia and Northweste ...
s, and
common buzzard The common buzzard (''Buteo buteo'') is a medium-to-large bird of prey which has a large range. It is a member of the genus '' Buteo'' in the family Accipitridae. The species lives in most of Europe and extends its breeding range across much of ...
s. Many
moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...
s,
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
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 hindwin ...
, and other
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s are crepuscular and vespertine. For example, Coprophanaeus ensifer, a South American necrophagous beetle, is most active during at dusk and dawn.


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 ...
*
Crypsis In ecology, crypsis is the ability of an animal or a plant to avoid observation or detection by other animals. It may be part of a predation strategy or an antipredator adaptation. Methods include camouflage, nocturnality, subterranean life ...
*
Diurnality Diurnality is a form of plant and ethology, animal behavior characterized by activity during daytime, with a period of sleeping or other inactivity at night. The common adjective used for daytime activity is "diurnal". The timing of activity by ...
*
Nocturnality Nocturnality is a behavior in some non-human animals 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 hi ...


References

{{Light Ethology Ethology Night