Moths are a group of winged insect of the order Lepidoptera, the other well-known members of which are butterflies, they are characterized by dark or dull patterned wings.[1] 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 of moth,[2] many of which have yet to be described. Most species of moth are nocturnal, although there are also crepuscular and diurnal species.
Moths have a multi-stage life cycle, and undergo metamorphosis. Winged adults generally lay eggs on or near plant foliage on which their larvae, known as caterpillars, will feed. Although most feed on plant matter, some species such as Eupithecia orichloris are carnivorous insectivores.[3][4] The caterpillars grow, sometimes very rapidly, and when fully developed will pupate, often after constructing a cocoon from silk. When metamorphosis is complete, the pupal skin splits, the adult insect climbs out and slowly expands its wings to dry and harden, before flying off.
Moths are commonly thought to eat clothes and fabrics, however it is only their larvae, caterpillars, which may consume natural fibres.[5]: 1–3
Moths are very widely distributed across diverse terrestrial habitats and are found on all continents except Antarctica.[6][7][8]
Etymology
The modern English word moth comes from Old English moððe (cf. Northumbrian mohðe) from Common Germanic (compare Old Norse motti, Dutch mot, and German Motte all meaning ‘moth’). Its origins are possibly related to the Old English maða 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.
Taxonomy

Among the Lepidoptera, moths comprise about 90% (remaining 10% being butterflies). There are around 160,000 described species.[9][10] There is estimated to be tens of thousands of unidentified species, especially from tropical regions. The name Heterocera is no longer used in modern classification, however the name is often used to refer to moths; as Rhopalocera is like-wise used as the name for butterflies.[11]
Among the 125 known families, moths make up 118 of the Lepidoptera families, comprising all but the seven butterfly families.[11][needs update]
While there are 160,000 species described currently, the total number is estimated to be around 500,000. On average, over 1,000 species of Lepidoptera is described annually in recent years.[12][13][needs update]
Some of the largest moth families include Erebidae, Noctuidae, Geometroidea, and Pyralidae.[14][15]
Differences between butterflies and moths
While the butterflies form a 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.[16]
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 period.[17]
Moths also tend to have far more hair-like scales than butterflies.[15]
Moths are usually nocturnal flyers in contrast to butterflies which fly by day, however there are many exceptions such as the Cinnabar moth.[18][19]
Life cycle and description
Being oviparous, moth larva hatch as eggs. Female moths will usually nurture the eggs with a layer of nutrients, sometimes providing a protective chemical layer, or covering with urticating hairs (such as in Lymantriinae). Eggs vary in shape and size, some being very elongated, and in colour, some being well camouflaged, others being brightly coloured to indicate a chemical defence (aposematism).[5]
Caterpillar

Moth larvae, or caterpillars, are the first life stage after emergence from the egg. During emergence from the egg larva will consume the egg shell. Larvae have sclerotized heads, and mouthparts adapted for chewing. They have very short antennae and stemmata, mainly used for food detection. They possess a spinneret for producing silk. The larval stage varies in duration depending on temperature and food availability, but usually takes from four to eight weeks, sometimes as short as two weeks. Species which enter diapause during the winter may take up to 10 months, some species may take years. Larvae have three pairs of thoracic legs, and multiple proleg with gripping hooks called crochets, and will walk in order to reach a suitable spot for pupation, and to reach food.[20][5]: 242 [21]
The larval stage is the main feeding stage, where the larva will in most species consume plant matter.[22]
At the end of their life stage, caterpillars will seek out a location to enter the pupal stage. Usually this will be underground, or beneath plant debris. Most species are quite specific around where they will choose to pupate. Some species will pupate without constructing a cocoon, others will construct a cocoon from silk, often also incorporating plant debris, such as Cerura vinula which may incorporate pieces of chewed bark into the silk cocoon.[23][5]: 50 [24]

Pupa
During the pupal stage, the pupa will undergo metamorphosis and will be immobile for the duration of the life stage. The outer shell of the Pupa is sclerotised, and usually shiny and smooth.[15] At the end of this life stage, the pupa will emerge usually at specific times of the day, for instance in night flying moths usually late afternoon, in day flying usually early morning. To break free from the cocoon some species may secrete potassium hydroxide from their mouth to weaken the silk.[5]: 53
Adult
The adult stage of the moth is primarily for reproduction and for dispersing from the location of pupation. Emerged moths will spend a few hours opening its wings for the first time, and allowing them to dry and harden. Usually emergence is specific to a certain time of the day, in night flying moths late afternoon, in day flying early morning. Moth wings are made of small overlapping scales. While providing insulation, the scales are also responsible for enabling the varied patterns seen on moths, each scale having a different pigment to make up the overall visible pattern. Moth scales can vary greatly in shape between families, from Filamentous and hairlike to club or paddle shaped.[5]: 66 [25]

Almost all adult moths have a distinct coiled proboscis used to pull in fluids such as plant sap or water, and in many species is used to feed on flower nectar.[26] Adult moths have two large compound eyes, which can each range from 200 to 27,000 facets. most adult moths also possess ocelli, above their compound eyes; likely for orientation purposes.[5] Moths have two sensory antennae, which take a branch like shape and have no bulge on the end but taper to a point.[27]: 559–560
Moths are able to produce sound for communication in a variety of ways, some species possessing tymbal organs, for the purpose of warning others to the presence of predators such as bats. Moths possess excellent hearing, from organs usually located on their abdomen, and can detect the sound of bats’ wings.[28][29][30]
Moths use their dark and varied colouration which may resemble bark, stones, or dead leaves, for camouflage, avoiding the detection of predators. This is often in contrast to the caterpillar stage, which are often green to resemble the leaves which they must feed from.[31]
Females utilise pheromones to attract males of same species, remaining stationary while males locate them. Male moths have extremely acute capacity to detect scent, and can reliably track a female who is releasing pheromones up to a kilometre away, due to their highly sensitive antennae.[32][33][34] Since moths are usually quite dispersed, scent is more reliable a means of mate attraction than sound as is seen in many other insects.[16]
Some of the largest known moths can be found in the family Saturniidae, among which Attacus atlas of Southeast Asia attains a wingspan of up to 300 mm.[35] The smallest moths have wing-spans as small as 2.5 mm, such as Nepticulidae. The average among all moths is around 25 mm.[15]
Evolution

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 plants, 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.[36]
Industrial melanism in modern moths
A phenomenon associated with human industrialisation has been observed in species such as the peppered moth, in which the occurrence of individuals which are darker than the typical form have increased in number.[37][31] Increases of melanism in moths was first observed in the mid nineteenth century in Britain, and was associated with industrialized areas of the country where tree trunks were blackened with soot from coal burning. The phenomenon has been widely described as a classical example of natural selection, in which the darker moths gained better camouflage against the blackened tree bark, and therefore had a higher rate of survival and subsequently reproductive success.[38][39][40][41]
Predators and parasites

Nocturnal insectivores often feed on moths; these include some bats, some species of owls and other species of birds. Moths also are eaten by some species of lizards, amphibians, cats, dogs, rodents, and some bears.[42] Moth larvae are vulnerable to being parasitised by Ichneumonidae.[43][44][45]
Baculoviruses are parasite double-stranded DNA insect 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.[citation needed]
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 centimetres or inches in its flight to evade attack,[46] and tiger moths can emit clicks to foil bats’ echolocation.[47][48]
The fungus Ophiocordyceps sinensis infects the larvae of many different species of moths.[49]
Ecology

Moths, like butterflies, bees and other more popularly recognised 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.[50] Some studies indicate that certain species of moths, such as those belonging to the families Erebidae and Sphingidae, may be the key pollinators for some flowering plants in the Himalayan ecosystem.[51][52] 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.[53][54][55][56] 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.[57]
Both as caterpillars and adult, moths are a substantial part of the insect biomass available to insectivorous vertebrates, they are also an important dietary source for many bats and passerines.[58][59]
Behaviour
Attraction to light

Moths frequently appear to circle artificial lights. The reason for this behaviour (positive phototaxis) has not been solved, however there is widespread speculation that it is a result of orientating instincts, which in the natural environment of night flying moths would cause them to fly upward towards the moon, and may be a part of dispersal behaviour.[60][61][62][63]
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.[64]
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[65][66][67] or has severely disrupted nocturnal pollination.[68][69]
Relationship to humans
Some moths, particularly their caterpillars, can be major agricultural pests in many parts of the world. Examples include corn borers and bollworms.[70] 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.[71]
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.[72] 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.[citation needed]
Despite being commonly thought to be undertaken by all moths,[73][74] 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, 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).[75] Many species of adult moths do however eat: for instance, many will drink nectar.[72]
Items of fabric infested by clothes moth larvae may be treated by freezing them for several days at a temperature below −8 °C (18 °F).[76]
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. As of 2002, the silk industry produces more than 130 million kilograms of raw silk, worth about 250 million U.S. dollars, each year.[77][78][79]
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).[citation needed]
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.[80]
Moth traps are often used by amateur collectors, entomologists, or in pest control, to capture moths.[81]
Notable species
- Atlas moth (Attacus atlas), 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[82]
- 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 humans[83]
- 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
- 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
- Magpie moth (Nyctemera annulata), pupae used in Maori children’s game
- Bamboo worm (Omphisa fuscidentalis), used as food source in Thailand and Nagaland
- Hummingbird hawk-moth (Macroglossum stellatarum), seen as good/lucky omen
- Mother Shipton moth (Callistege mi), forewing pattern represents English folklore figure Mother Shipton
- Mopone moth (Gonimbrasia belina), caterpillars are food source in southern Africa
- African wild silk worm (Gonometa postica)
- Japanese silk moth (Antheraea yamamai)
- Garden tiger moth (Arctia caja), caterpillar hairs known for causing hives and irritation
- Erasmia pulchella, known for beautiful wing patterns resembling nishiki cloth (type of Japanese brocade)
Agricultural pests
- 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
- Leuvana moth (Levuana iridescens), notorious coconut pest made extinct
Gallery
-
Leaf-shaped moth (Pergesa acteus)
-
Giant grey moth (Agrius convolvuli)
-
Oleander hawk-moth or army green moth (Daphnis nerii)
-
Six-spot burnet moths mating (Zygaena filipendulae)
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Protective silk (or similar material) case (cocoon)
-
A caterpillar of death’s-head hawkmoth
-
Mating pair of Laothoe populi, or poplar hawkmoths, showing two different color variants
-
White-lined sphinx moth in Colorado, United States
-
Closeup of a common clothes moth
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Giant silk moth (Adelowalkeria tristygma)
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Adult emperor moth (Gonimbrasia belina)
-
A moth on a marble floor in Kolkata, India
-
Clothes moth, eye
-
Female rose-myrtle lappet moth hanging on the wooden door
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A moth on artificial flowers in Kolkata, India
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Tiger moth (Aglaomorpha histrio)
-
Box tree moth (Cydalima perspectalis)
See also
References
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- ^ “Table 74. Raw silk: production (including waste)”. Food and Agriculture Organization of the United Nations. Archived from the original on 18 March 2008. Retrieved 2 October 2008.
Table lists worldwide raw silk production 132,400 metric tonnes in 2002
- ^ “Silk Exchanges of Tamil Nadu and Andhra Pradesh”. Central Silk Board of India. Archived from the original on 7 March 2007. gives silk prices in rupees. Exchange rate is about 50 RS to dollar.
- ^ “Silk Worm Farming”. Vegan Society. Archived from the original on 19 June 2008. Retrieved 2 October 2008.
World Raw Silk Production in 1996 is listed as 83,670 metric tonnes
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- ^ Waring, Paul (2001). A Guide to Moth Traps and Their Use. ISBN 978-1-8380246-1-1.
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- ^ Brundage, Adrienne (23 March 2009), Other Arthropods of Forensic Importance, Texas A&M University, Texas A&M University Forensic Entomology Lecture
External links
Quotations related to Moths at Wikiquote- Wilkes, Benjamin (1749). The English Moths and Butterflies.