Moths are a group of insects that includes all members of the orderLepidoptera that are notbutterflies.[1] They were previously classified as suborderHeterocera, but the group isparaphyletic 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 160,000species of moth,[2] many of which have yet to be described. Most species of moth arenocturnal, although there are alsocrepuscular anddiurnal species.
Etymology
Themodern English wordmoth comes fromOld Englishmoððe (cf.Northumbrianmohðe) fromCommon Germanic (compareOld Norsemotti,Dutchmot, andGermanMotte all meaning 'moth'). Its origins are possibly related to the Old Englishmaða meaning 'maggot' or from the root ofmidge which until the 16th century was used mostly to indicate thelarva, usually in reference to devouring clothes.
While thebutterflies form amonophyletic 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 andMacrolepidoptera, Heterocera andRhopalocera, Jugatae and Frenatae,Monotrysia, andDitrysia.[3]
Although the rules for distinguishing moths from butterflies are not well established, one very good guiding principle is that butterflies have thinantennae and (with the exception of the familyHedylidae) 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 theCarboniferous period, but only evolved their characteristicproboscis alongside the rise ofangiosperms in theCretaceous period.[4]
Moth larvae, orcaterpillars, makecocoons 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.[5]
Evolution
Moths evolved long before butterflies; mothfossils have been found that may be 190 million years old. Both types of Lepidoptera are thought to have co-evolved withflowering 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 isArchaeolepis mane. Its fossil fragments show scaled wings that are similar tocaddisflies in their veining.[6]
Significance to humans
An adult male pine processionary moth (Thaumetopoea pityocampa). This species is a serious forest pest when in its larval state. Notice the bristle springing from the underside of the hindwing (frenulum) and running forward to be held in a small catch of the forewing, whose function is to link the wings together.
Several moths in the familyTineidae are commonly regarded as pests because their larvae eatfabric such asclothes andblankets made from naturalproteinaceous fibers such aswool orsilk.[9] 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 fromjuniper andcedar, bylavender, or by other natural oils; however, many consider this unlikely to prevent infestation.Naphthalene (the chemical used inmothballs) is considered more effective, but there are concerns over its effects on human health.
Despite beingcommonly thought to be undertaken by all moths,[10][11] only the larvae ofseveral 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 theLuna,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).[12] Many species of adult moths do however eat: for instance, many will drinknectar.[9]
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).[13]
Some moths arefarmed for their economic value. The most notable of these is thesilkworm, the larva of the domesticated mothBombyx mori. It is farmed for the silk with which it builds itscocoon. As of 2002[update], the silk industry produces more than 130 million kilograms of raw silk, worth about 250 millionU.S. dollars, each year.[14][15][16]
Not all silk is produced byBombyx mori. There are several species ofSaturniidae 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 areused as food, particularly in Africa, where they are an important source of nutrition. The mopane worm, the caterpillar ofGonimbrasia belina, from the family Saturniidae, is a significant food resource insouthern 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.[17]
Nocturnal insectivores often feed on moths; these include somebats, some species ofowls and other species ofbirds. Moths also are eaten by some species oflizards,amphibians,cats,dogs,rodents, and somebears. Moth larvae are vulnerable to being parasitized byIchneumonidae.
Baculoviruses are parasitedouble-stranded DNAinsect viruses that are used mostly asbiological control agents. They are members of theBaculoviridae, 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 thenoctuid moth that causes it to drop a few centimeters or inches in its flight to evade attack,[18] andtiger moths can emit clicks to foil bats'echolocation.[19][20]
Moths, like butterflies, bees and other more popularly recognizedpollinating 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.[22] Some studies indicate that certain species of moths, such as those belonging to the familiesErebidae andSphingidae, may be the key pollinators for some flowering plants in the Himalayan ecosystem.[23][24] 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.[25][26][27][28] Some researchers say it is likely that many plants thought to be dependent on bees forpollination also rely on moths, which have historically been less observed because they pollinate mainly at night.[29]
Attraction to light
Moths circling an electric lightbulb
Moths frequently appear to circle artificial lights. The reason for this behavior (positivephototaxis) 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.[30]
Studies have found thatlight pollution caused by increasing use of artificial lights has either led to a severe decline in moth population in some parts of the world[31][32][33] or has severely disrupted nocturnal pollination.[34][35]
Examples of moth species
Atlas moth (Attacus atlas), one of the largest moths in the world
Ornate moth (Utetheisa ornatrix), the subject of numerous behavioral studies regarding sexual selection
Ghost moths (Hepialidae spp.), larvae mummified by the parasitic fungusOphiocordyceps 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
^Darby, Gene (1958).What is a Butterfly. Chicago:Benefic Press. p. 41.
^Hoyt, Cathryn."Evolution of Moths and Butterflies".Chihuahuan Desert Nature Center. Archived fromthe original on 6 January 2014.Studying the evolution of butterflies and moths is challenging, since fossils are so rare. But the few Lepidopteran fossils that exist, captured in amber or compressed in fine-grained rocks, show an astonishing amount of detail. The earliest Lepidopteran fossils appear in rocks that are about 190 million years old. These tiny fragments of scaled wings and bodies clearly indicate that moths evolved before butterflies.
^Conlong, D.E. (February 1994). "A review and perspectives for the biological control of the African sugarcane stalkborer Eldana saccharina Walker (Lepidoptera: Pyralidae)".Agriculture, Ecosystems & Environment.48 (1):9–17.Bibcode:1994AgEE...48....9C.doi:10.1016/0167-8809(94)90070-1.
^"Table 74. Raw silk: production (including waste)". Food and Agriculture Organization of the United Nations.Archived from the original on 18 March 2008. Retrieved2 October 2008.Table lists worldwide raw silk production 132,400 metric tonnes in 2002
^"Silk Worm Farming". Vegan Society. Archived fromthe original on 19 June 2008. Retrieved2 October 2008.World Raw Silk Production in 1996 is listed as 83,670 metric tonnes
^van Langevelde, Frank; Braamburg-Annegarn, Marijke; Huigens, Martinus E.; Groendijk, Rob; Poitevin, Olivier; van Deijk, Jurriën R.; Ellis, Willem N.; van Grunsven, Roy H. A.; de Vos, Rob (4 January 2018). "Declines in moth populations stress the need for conserving dark nights".Global Change Biology.24 (3):925–932.Bibcode:2018GCBio..24..925V.doi:10.1111/gcb.14008.ISSN1354-1013.PMID29215778.S2CID205145880.
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