Moths are a

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

group of insects that includes all members of the order Lepidoptera that are not

butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises t ...

, with moths making up the vast majority of the order. There are thought to be approximately 160,000

species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...

of moth, many of which have yet to be described. Most species of moth are

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 ...

, but there are also

crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of dayli ...

and diurnal species.

Differences between butterflies and moths

While the

form a

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...

group, the moths, comprising the rest of the Lepidoptera, do not. Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not monophyletic: Microlepidoptera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and

Ditrysia The Ditrysia are a natural group or clade of insects in the lepidopteran order containing both butterflies and moths. They are so named because the female has two distinct sexual openings: one for mating, and the other for laying eggs (in cont ...

.Scoble, MJ 1995. The Lepidoptera: Form, function and diversity. Oxford, UK: Oxford University Press; 404 p. Although the rules for distinguishing moths from butterflies are not well established, one very good guiding principle is that butterflies have thin antennae and (with the exception of the family Hedylidae) have small balls or clubs at the end of their antennae. Moth antennae are usually feathery with no ball on the end. The divisions are named by this principle: "club-antennae" (Rhopalocera) or "varied-antennae" (Heterocera). Lepidoptera first evolved during the Carboniferous period, but only evolved their characteristic proboscis alongside the rise of angiosperms in the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...

period.

Etymology

The modern English word ''moth'' comes from Old English ( cf. Northumbrian ) from

Common Germanic Proto-Germanic (abbreviated PGmc; also called Common Germanic) is the reconstructed proto-language of the Germanic branch of the Indo-European languages. Proto-Germanic eventually developed from pre-Proto-Germanic into three Germanic bra ...

(compare

Old Norse Old Norse, Old Nordic, or Old Scandinavian, is a stage of development of North Germanic languages, North Germanic dialects before their final divergence into separate Nordic languages. Old Norse was spoken by inhabitants of Scandinavia and t ...

, Dutch , and German all meaning 'moth'). Its origins are possibly related to the Old English meaning '

maggot A maggot is the larva of a fly (order Diptera); it is applied in particular to the larvae of Brachycera flies, such as houseflies, cheese flies, and blowflies, rather than larvae of the Nematocera, such as mosquitoes and crane flies. E ...

' or from the root of ''

midge A midge is any small fly, including species in several families of non- mosquito Nematoceran Diptera. Midges are found (seasonally or otherwise) on practically every land area outside permanently arid deserts and the frigid zones. Some m ...

'' which until the 16th century was used mostly to indicate the

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. Th ...

, usually in reference to devouring clothes.

Caterpillar

Moth larvae, or caterpillars, make cocoons from which they emerge as fully grown moths with wings. Some moth caterpillars dig holes in the ground, where they live until they are ready to turn into adult moths.

History

Moths evolved long before butterflies; moth

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

s have been found that may be 190 million years old. Both types of Lepidoptera are thought to have co-evolved with

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of ...

s, mainly because most modern species, both as adults and larvae, feed on flowering plants. One of the earliest known species that is thought to be an ancestor of moths is ''Archaeolepis mane''. Its fossil fragments show scaled wings that are similar to caddisflies in their veining.

Economics

Significance to humans

Some moths, particularly their caterpillars, can be major

agricultural Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled peopl ...

pests in many parts of the world. Examples include corn borers and

bollworm Bollworm is the common term for a moth larva that attacks the fruiting bodies of certain crops, especially cotton. The most common moths known as bollworms are: * Red or Sudan bollworm, '' Diparopsis castanea'' * Rough bollworm, '' Earias perhuege ...

s. The caterpillar of the spongy moth (''Lymantria dispar'') causes severe damage to forests in the northeastern United States, where it is an

invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species adv ...

. In temperate climates, the

codling moth The codling moth (''Cydia pomonella'') is a member of the Lepidopteran family Tortricidae. They are major pests to agricultural crops, mainly fruits such as apples and pears. Because the larvae are not able to feed on leaves, they are highly ...

causes extensive damage, especially to fruit farms. In tropical and subtropical climates, the diamondback moth ('' Plutella xylostella'') is perhaps the most serious pest of brassicaceous crops. Also in sub-Saharan Africa, the African sugarcane borer is a major pest of sugarcane,

maize Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn ( North American and Australian English), is a cereal grain first domesticated by indigenous peoples in southern Mexico about 10,000 years ago. ...

, and

sorghum ''Sorghum'' () is a genus of about 25 species of flowering plants in the grass family ( Poaceae). Some of these species are grown as cereals for human consumption and some in pastures for animals. One species is grown for grain, while many ot ...

. Several moths in the family

Tineidae Tineidae is a family of moths in the order Lepidoptera described by Pierre André Latreille in 1810. Collectively, they are known as fungus moths or tineid moths. The family contains considerably more than 3,000 species in more than 300 genera. ...

are commonly regarded as pests because their larvae eat fabric such as clothes and

blanket A blanket is a swath of soft cloth large enough either to cover or to enfold most of the user's body and thick enough to keep the body warm by trapping radiant body heat that otherwise would be lost through convection. Etymology The ter ...

s made from natural proteinaceous fibers such as

wool Wool is the textile fibre obtained from sheep and other mammals, especially goats, rabbits, and camelids. The term may also refer to inorganic materials, such as mineral wool and glass wool, that have properties similar to animal wool. ...

silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the ...

.Scott, Thomas (1995)
''Concise Encyclopedia Biology''
. Walter de Gruyter. . They are less likely to eat mixed materials containing some artificial fibers. There are some reports that they may be repelled by the scent of wood from juniper and cedar, by

lavender ''Lavandula'' (common name lavender) is a genus of 47 known species of flowering plants in the mint family, Lamiaceae. It is native to the Old World and is found in Cape Verde and the Canary Islands, and from Europe across to northern and east ...

, or by other natural oils; however, many consider this unlikely to prevent infestation.

Naphthalene Naphthalene is an organic compound with formula . It is the simplest polycyclic aromatic hydrocarbon, and is a white crystalline solid with a characteristic odor that is detectable at concentrations as low as 0.08 ppm by mass. As an aromat ...

(the chemical used in mothballs) is considered more effective, but there are concerns over its effects on human health. Moth larvae may be killed by freezing the items which they infest for several days at a temperature below . While moths are notorious for eating clothing, most species do not, and some moth adults do not even eat at all. Some, like the Luna,

Polyphemus Polyphemus (; grc-gre, Πολύφημος, Polyphēmos, ; la, Polyphēmus ) is the one-eyed giant son of Poseidon and Thoosa in Greek mythology, one of the Cyclopes described in Homer's ''Odyssey''. His name means "abounding in songs and le ...

Atlas An atlas is a collection of maps; it is typically a bundle of maps of Earth or of a region of Earth. Atlases have traditionally been bound into book form, but today many atlases are in multimedia formats. In addition to presenting geograp ...

, Promethea, cecropia, and other large moths do not have mouth parts. This is possible because they live off the food stores from when they were a caterpillar, and only live a short time as an adult (roughly a week for some species). Many species of adult moths do however eat: for instance, many will drink nectar. Some moths are

farm A farm (also called an agricultural holding) is an area of land that is devoted primarily to agricultural processes with the primary objective of producing food and other crops; it is the basic facility in food production. The name is used ...

ed for their economic value. The most notable of these is the silkworm, the larva of the domesticated moth '' Bombyx mori''. It is farmed for the silk with which it builds its cocoon. , the silk industry produces more than 130 million kilograms of raw silk, worth about 250 million U.S. dollars, each year. Not all silk is produced by ''Bombyx mori''. There are several species of Saturniidae that also are farmed for their silk, such as the ailanthus moth ('' Samia cynthia'' group of species), the Chinese oak silkmoth ('' Antheraea pernyi''), the Assam silkmoth ('' Antheraea assamensis''), and the Japanese silk moth ('' Antheraea yamamai''). The larvae of many species are used as food, particularly in Africa, where they are an important source of nutrition. The mopane worm, the caterpillar of ''

Gonimbrasia belina ''Gonimbrasia belina'' is a species of emperor moth which is native to the warmer parts of southern Africa. Its large edible caterpillar, known as the mopane worm, madora, amacimbi or masontja, feeds primarily but not exclusively on mopane tree l ...

'', from the family Saturniidae, is a significant food resource in

southern Africa Southern Africa is the southernmost subregion of the African continent, south of the Congo and Tanzania. The physical location is the large part of Africa to the south of the extensive Congo River basin. Southern Africa is home to a number ...

. Another saturniid used as food is the cavorting emperor (''

Usta terpsichore ''Usta terpsichore'', the cavorting emperor, is a species of moth in the family Saturniidae. It is found in Angola, Burkina Faso, Eritrea, Ethiopia, Malawi, Mozambique, South Africa, Tanzania, Zambia and Zimbabwe. The larvae feed on '' Commiphor ...

''). In one country alone,

Congo Congo or The Congo may refer to either of two countries that border the Congo River in central Africa: * Democratic Republic of the Congo, the larger country to the southeast, capital Kinshasa, formerly known as Zaire, sometimes referred to a ...

, more than 30 species of moth larvae are harvested. Some are sold not only in the local village markets, but are shipped by the ton from one country to another.

Predators and parasites

Tomato Hornworm Parasitized by Braconid Wasp

Nocturnal insectivores often feed on moths; these include some bats, some species of owls and other species of

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...

s. Moths also are eaten by some species of

lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia al ...

s, amphibians,

cat 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 ...

s, dogs,

rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are roden ...

s, and some

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. Moth larvae are vulnerable to being parasitized by Ichneumonidae. Baculoviruses are parasite double-stranded DNA

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 ...

viruses that are used mostly as

biological control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically als ...

agents. They are members of the Baculoviridae, a family that is restricted to insects. Most baculovirus isolates have been obtained from insects, in particular from Lepidoptera. There is evidence that ultrasound in the range emitted by bats causes flying moths to make evasive maneuvers. Ultrasonic frequencies trigger a reflex action in the noctuid moth that causes it to drop a few centimeters or inches in its flight to evade attack, and tiger moths can emit clicks to foil bats' echolocation. The fungus '' Ophiocordyceps sinensis'' infects the larvae of many different species of moths.

Ecological importance

Moths, like butterflies, bees and other more popularly recognized pollinating insects, serve an essential role as pollinators for many flowering plants, including species that bees do not visit. Nocturnal moths fly from flower to flower to feed on nectar during the night much as their diurnal relatives do during the day. A study conducted in the UK found moths dusted with pollen from 47 different plant species, including seven species largely ignored by bees. Some studies indicate that certain species of moths, such as those belonging to the families Erebidae and

Sphingidae The Sphingidae are a family of moths ( Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as “hornworms”; it includes about 1,450 species. It is best represented in the tropics, ...

, may be the key pollinators for some flowering plants in the Himalayan ecosystem. The roles of moths as pollinators have been studied less frequently than those of diurnal pollinators, but recent studies have established that moths are important, but often overlooked, nocturnal pollinators of a wide range of plants. Some researchers say it is likely that many plants thought to be dependent on bees for pollination also rely on moths, which have historically been less observed because they pollinate mainly at night.

Attraction to light

Moths frequently appear to circle artificial lights, although the reason for this behavior (positive phototaxis) is currently unknown. One hypothesis is called celestial or transverse orientation. By maintaining a constant angular relationship to a bright celestial light, such as the moon, they can fly in a straight line. Celestial objects are so far away that, even after travelling great distances, the change in angle between the moth and the light source is negligible; further, the moon will always be in the upper part of the visual field, or on the horizon. When a moth encounters a much closer artificial light and uses it for navigation, the angle changes noticeably after only a short distance, in addition to being often below the horizon. The moth instinctively attempts to correct by turning toward the light, thereby causing airborne moths to come plummeting downward, and resulting in a spiral flight path that gets closer and closer to the light source. Studies have found that light pollution caused by increasing use of artificial lights has either led to a severe decline in moth population in some parts of the world or has severely disrupted nocturnal pollination.

Noteworthy moths

*Atlas moth (''

Attacus atlas ''Attacus atlas'', the Atlas moth, is a large saturniid moth endemic to the forests of Asia. The species was first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''. Description The Atlas moth is one of the largest ...

''), one of the largest moths in the world *Herculese moth ('' Coscinocera hercules''), largest moth in Australia *White witch moth ('' Thysania agrippina''), the Lepidopteran with the longest wingspan * Madagascan sunset moth (''Chrysiridia rhipheus''), considered to be one of the most impressive and beautiful Lepidoptera * Death's-head hawkmoth (''Acherontia'' spp.), is associated with the supernatural and evil and has been featured in art and movies * Peppered moth (''Biston betularia''), the subject of a well-known study in natural selection *Luna moth (''

Actias luna The Luna moth (''Actias luna'') also known as the American moon moth is a Nearctic moth in the family Saturniidae, subfamily Saturniinae, a group commonly known as giant silk moths. It has lime-green colored wings and a white body. The larvae ( ...

'') * Grease moth (''Aglossa cuprina''), known to have fed on the rendered fat of humansBrundage, Adrienne (23 March 2009), Other Arthropods of Forensic Importance, Texas A&M University, Texas A&M University Forensic Entomology Lecture * Emperor gum moth (''Opodiphthera eucalypti'') *

Polyphemus moth ''Antheraea polyphemus'', the Polyphemus moth, is a North American member of the family Saturniidae, the giant silk moths. It is a tan-colored moth, with an average wingspan of 15 cm (6 in). The most notable feature of the moth is ...

(''Antheraea polyphemus'') * Bogong moth (''Agrotis infusa''), known to have been a food source for southeastern

indigenous Australians Indigenous Australians or Australian First Nations are people with familial heritage from, and membership in, the ethnic groups that lived in Australia before British colonisation. They consist of two distinct groups: the Aboriginal peoples o ...

* Ornate moth (''Utetheisa ornatrix''), the subject of numerous behavioral studies regarding sexual selection

Moths of economic significance

*Spongy moth (''

Lymantria dispar ''Lymantria dispar'', also known as the gypsy moth or the spongy moth, is a species of moth in the family Erebidae. ''Lymantria dispar'' is subdivided into several subspecies, with subspecies such as ''L. d. dispar'' and ''L. d. japonica'' be ...

''), an invasive species pest of hardwood trees in North America * Winter moth (''Operophtera brumata''), an invasive species pest of hardwood trees, cranberry and blueberry in northeastern North America * Corn earworm or cotton bollworm (''Helicoverpa zea''), a major agricultural pest * Indianmeal moth (''Plodia interpunctella''), a major pest of grain and flour *

Codling moth The codling moth (''Cydia pomonella'') is a member of the Lepidopteran family Tortricidae. They are major pests to agricultural crops, mainly fruits such as apples and pears. Because the larvae are not able to feed on leaves, they are highly ...

(''Cydia pomonella''), a pest mostly of apple, pear and walnut trees * Light brown apple moth (''Epiphyas postvittana''), a highly polyphagous pest *Silkworm ('' Bombyx mori''), for its silk *

Wax moth Waxworms are the caterpillar larvae of wax moths, which belong to the family Pyralidae (snout moths). Two closely related species are commercially bred – the lesser wax moth (''Achroia grisella'') and the greater wax moth (''Galle ...

s ('' Galleria mellonella'', '' Achroia grisella''), pests of bee hives *'' Duponchelia fovealis'', a new invasive pest of vegetables and ornamental plants in the United States

Gallery

File:Micrographia Schem 30.jpg, Diagram of a

plume moth The Pterophoridae or plume moths are a family of Lepidoptera with unusually modified wings. Though they belong to the Apoditrysia like the larger moths and the butterflies, unlike these they are tiny and were formerly included among the assemblag ...

from Robert Hooke's '' Micrographia'' File:Kerala Leaf Insect.jpg, Leaf-shaped moth (''

Pergesa acteus ''Pergesa'' is a monotypic moth genus in the family Sphingidae first described by Francis Walker in 1856. Its only species, ''Pergesa acteus'', the green pergesa hawkmoth, was described by Pieter Cramer in 1779. Distribution It is found in S ...

'') File:Giant grey moth.png, Giant grey moth ('' Agrius convolvuli'') File:Colourful Moth.jpg, Oleander hawk-moth or army green moth ('' Daphnis nerii'') File:Red spotted moths 1 (3745889925).jpg, Six-spot burnet moths mating (''

Zygaena filipendulae The six-spot burnet (''Zygaena filipendulae'') is a day-flying moth of the family Zygaenidae. Subspecies *''Z. f. altapyrenaica'' Le Charles, 1950 *''Z. f. arctica'' Schneider, 1880 *''Z. f. balcanirosea'' Holik, 1943 *''Z. f. campaniae'' ...

'') File:Silk cocoon.jpg, Protective silk (or similar material) case (cocoon) File:Caterpillar-----02.jpg, A caterpillar of death's-head hawkmoth File:Joined moths.JPG, Mating pair of '' Laothoe populi'', or poplar hawkmoths, showing two different color variants File:White-lined sphinx moth.JPG, White-lined sphinx moth in Colorado, United States File:Мебельная моль.jpg, Closeup of a

common clothes moth Common may refer to: Places * Common, a townland in County Tyrone, Northern Ireland * Boston Common, a central public park in Boston, Massachusetts * Cambridge Common, common land area in Cambridge, Massachusetts * Clapham Common, originally com ...

File:Giant silk moth (Adelowalkeria tristygma).jpg, Giant silk moth ('' Adelowalkeria tristygma'') File:Adult Emperor Moth.jpg, Adult emperor moth (''

'') File:A Moth on marble floor.jpg, A moth on a marble floor in

Kolkata Kolkata (, or , ; also known as Calcutta , List of renamed places in India#West Bengal, the official name until 2001) is the Capital city, capital of the Indian States and union territories of India, state of West Bengal, on the eastern ba ...

, India File:Clothes moth.jpg, Clothes moth, eye File:Female rose-myrtle lappet moth.jpg, Female rose-myrtle lappet moth hanging on the wooden door

References

{{Authority control Agricultural pest insects Household pest insects * Paraphyletic groups