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Moths are a group of insects that includes all members of the order Lepidoptera that are not butterflies. They were previously classified as suborder Heterocera, but the group is paraphyletic with respect to butterflies (suborder Rhopalocera) and neither subordinate taxon is used in modern classifications. Moths make up the vast majority of the order. There are approximately 160,000
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of moth, many of which have yet to be described. Most species of moth are
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 ...
, although there are also
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...
and diurnal species.


Differences between butterflies and moths

While the butterflies form a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
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.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
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
s in the Cretaceous period.


Etymology

The
modern English Modern English, sometimes called New English (NE) or present-day English (PDE) as opposed to Middle and Old English, is the form of the English language that has been spoken since the Great Vowel Shift in England England is a Count ...
word ''moth'' comes from
Old English Old English ( or , or ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the Early Middle Ages. It developed from the languages brought to Great Britain by Anglo-S ...
( cf. Northumbrian ) from Common Germanic (compare
Old Norse Old Norse, also referred to as Old Nordic or Old Scandinavian, was 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 ...
, Dutch , and German all meaning 'moth'). Its origins are possibly related to the Old English meaning ' maggot' or from the root of '' midge'' which until the 16th century was used mostly to indicate the larva, 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 fossils 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 (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
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 pests in many parts of the world. Examples include corn borers and bollworms. The caterpillar of the spongy moth (''Lymantria dispar'') causes severe damage to forests in the northeastern United States, where it is an invasive species. In temperate climates, the codling moth 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, and sorghum. Several moths in the family Tineidae are commonly regarded as pests because their larvae eat fabric such as clothes and blankets made from natural proteinaceous fibers such as wool or silk.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, or by other natural oils; however, many consider this unlikely to prevent infestation. Naphthalene (the chemical used in mothballs) is considered more effective, but there are concerns over its effects on human health. Despite being commonly thought to be undertaken by all moths, only the larvae of several moth species eat animal fibres, creating holes in articles of clothing, in particular those made of wool. Most species do not eat fabrics, and some moth adults do not even eat at all. Some, like the Luna, Polyphemus,
Atlas An atlas is a collection of maps; it is typically a bundle of world map, maps of Earth or of a continent or region of Earth. Advances in astronomy have also resulted in atlases of the celestial sphere or of other planets. Atlases have traditio ...
, 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. Items of fabric infested by clothes moth larvae may be treated by freezing them for several days at a temperature below . Some moths are farmed 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'', from the family Saturniidae, is a significant food resource in southern Africa. Another saturniid used as food is the cavorting emperor ('' Usta terpsichore''). In one country alone, Congo, 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

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 (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s. Moths also are eaten by some species of lizards, amphibians, cats, 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 Mandible, lower jaws. About 40% of all mammal specie ...
s, and some
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. Moth larvae are vulnerable to being parasitized by Ichneumonidae. Baculoviruses are parasite double-stranded DNA
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, ...
viruses that are used mostly as biological control 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 The Erebidae are a family (biology), family of moths in the superfamily Noctuoidea. The family is among the largest families of moths by species count and contains a wide variety of well-known macromoth groups. The family includes the underwin ...
and Sphingidae, 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. 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 Saturniidae, saturniid moth endemic to the forests of Asia. The species was described by Carl Linnaeus in his 1758 10th edition of Systema Naturae, 10th edition of ''Systema Naturae''. The Atlas mo ...
''), one of the largest moths in the world *Hercules 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'') * 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''), known for being source of continuous cell cultures * Polyphemus moth (''Antheraea polyphemus'') * Bogong moth (''Agrotis infusa''), known to have been a food source for southeastern
indigenous Australians Indigenous Australians are people with familial heritage from, or recognised membership of, the various ethnic groups living within the territory of contemporary Australia prior to History of Australia (1788–1850), British colonisation. The ...
* Ornate moth (''Utetheisa ornatrix''), the subject of numerous behavioral studies regarding sexual selection * Ghost moths (''Hepialidae'' spp.), larvae mummified by the parasitic fungus '' Ophiocordyceps sinensis'' are highly valued in traditional Chinese and Tibetan medicine * Isabella tiger moth (''Pyrrharctia isabella''), known for woollybear festivals in Canada and the U.S. * Black witch moth (''Ascalapha odorata''), known for associations with souls and death in folklore and mythology within the Americas * Vampire moths (''Calyptra'' spp.) * Morgan's sphinx moth (''Xanthopan morganii''), known for orchid pollination * Comet moth (''Argema mittrei''), known for coevolutionary arms race with bats


Moth species that may cause significant economic damage

*Spongy moth ('' Lymantria dispar''), 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 (''Cydia pomonella''), a pest mostly of apple, pear and walnut trees * Light brown apple moth (''Epiphyas postvittana''), a highly polyphagous pest * Wax moths ('' 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 from Robert Hooke's '' Micrographia'' File:Kerala Leaf Insect.jpg, Leaf-shaped moth ('' Pergesa acteus'') 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'') 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 File:Giant silk moth (Adelowalkeria tristygma).jpg, Giant silk moth ('' Adelowalkeria tristygma'') File:Adult Emperor Moth.jpg, Adult emperor moth ('' Gonimbrasia belina'') File:A Moth on marble floor.jpg, A moth on a marble floor in Kolkata, 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 File:A Moth on artificial flowers.jpg, A moth on artificial flowers in Kolkata, India File:Aglaomorpha histrio on leaf.jpg, Tiger moth ('' Aglaomorpha histrio'') File:Box Tree Moth under leaf.jpg, Box tree moth ('' Cydalima perspectalis'')


See also

* Baculovirus * Clothing moth * Comparison of butterflies and moths * Iich'aa * Insect tea * List of moths * Lepidopterism * Pollination * Mothman * Mothra


External links

* Wilkes, Benjamin (1749).
The English Moths and Butterflies
'.


References

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