Lepidopterans have several distinctive features, especially colouredscales that cover their bodies, large triangularwings, and aproboscis for drinkingnectar. Almost all species have membranous wings.Mating and egg-laying are normally near or on host plants for thelarvae. Butterflies and moths undergocomplete metamorphosis. The larva is called acaterpillar; it has a cylindrical body with a well-developed head, chewing mouthparts, three pairs of legs and up to five pairs ofprolegs. As it grows, it changes in appearance, going through a series of stages calledinstars. Once mature, it develops into apupa. A few butterflies and many moth species spin asilk casing orcocoon for protection prior to pupating, while others just go underground.[4] A butterfly pupa, called achrysalis, has a hard skin, usually with no cocoon. In the pupa, metamorphosis occurs, and a sexually mature adult emerges.
Lepidopterans first appeared in the fossil record at theTriassic-Jurassic boundary. They havecoevolved withflowering plants since theangiosperm boom in theCretaceous. They have evolved a wide range of wing patterns ranging from drab moths to the brightly colored and complex-patterned butterflies.[3] A person who studies this order is called alepidopterist.
Butterflies and moths are mostlyherbivorous as caterpillars andnectar-feeding as adults. They serve as economically usefulpollinators; conversely, caterpillars are oftenpests inagriculture, as they can quickly consume large quantities of leaves. Females often produce up to 600 eggs.
The term Lepidoptera was used in 1746 byCarl Linnaeus in hisFauna Svecica.[5][6] The word is derived fromGreekλεπίςlepís,gen.λεπίδοςlepídos ("scale") andπτερόν ("wing").[7][8] The word "butterfly" is fromOld Englishbuttorfleoge, possibly from the pale butter color of some species' wings.[9][10] The word "moth" is from Old Englishmoððe from Common Germanic andGermanMotte meaning "moth".[11]The word "caterpillar" is fromMiddle Englishcatirpel,catirpeller, probably an alteration ofOld North Frenchcatepelose (from Latincattus, "cat" +pilosus, "hairy").[12]
The Lepidoptera are among the most successful groups of insects. They are found on all continents, exceptAntarctica, and inhabit all terrestrial habitats ranging from desert to rainforest, from lowland grasslands to mountain plateaus, but almost always associated with higher plants, especially angiosperms (flowering plants).[13] Among the most northern dwelling species of butterflies and moths is the Siberian Apollo (Parnassius arcticus), found in the Arctic Circle in northeasternYakutia, at an altitude of 1,500 metres (4,900 ft) above sea level.[14] In theHimalayas, Apollo species such asParnassius epaphus occur up to an altitude of 6,000 metres (20,000 ft).[15]: 221
Some species arecommensal orparasitic, inhabiting the bodies of organisms rather than the environment.Coprophagouspyralid moths, calledsloth moths, such asBradipodicola hahneli andCryptoses choloepi, are unusual in that they are exclusively found the fur ofsloths.[16][17] Two species ofTinea moths feed and breed on the horns of cattle. The larva ofZenodochium coccivorella is an internal parasite ofKermes scale insects. Many species breed in materials such as owl pellets, bat caves, honeycombs or diseased fruit.[17]
As of 2007, there were roughly 174,250 lepidopteran species described, with butterflies andskippers estimated to comprise around 17,950, and moths making up the rest.[1][18] The vast majority live in the tropics, but substantial diversity exists on most continents. North America has over 700 species of butterflies and over 11,000 species of moths,[19][20] while Australia has about 400 species of butterflies and 14,000 species of moths.[21] The diversity in eachfaunal region has been estimated by John Heppner in 1991 based partly on actual counts from the literature, partly on the card indices in theNatural History Museum (London) and theNational Museum of Natural History (Washington), and partly on estimates:[22]
Parts of an adult butterflyA – head, B – thorax, C – abdomen, 1 – prothoracic shield, 2 – spiracle, 3 – true legs, 4 – midabdominal prolegs, 5 – anal proleg, 6 – anal plate, 7 – tentacle, a – eye, b – stemmata (ocelli), c – antenna, d – mandible, e – labrum, f – frontal triangle.
Lepidoptera are morphologically distinguished from other orders by the presence ofscales on the body and appendages, especially thewings. Butterflies and moths vary in size frommicrolepidoptera only a few millimeters long, to conspicuous animals with a wingspan greater than 25 centimetres (9.8 in), such asQueen Alexandra's birdwing andAtlas moth.[23]: 246
The head houses sensing organs and mouthparts. Like the adult, the larva has a toughened, orsclerotized head capsule.[24] Here, twocompound eyes, andchaetosema, clusters of sensory bristles unique to Lepidoptera, occur, though many taxa have lost one or both of these clusters. The antennae vary widely in form among species and between the sexes. The antennae of butterflies are usually club-shaped, those of the skippers are hooked, while those of moths have segments variously enlarged or branched.[25]: 559–560
Themaxillary galeae are modified to form an elongatedproboscis. It consists of one to five segments, kept coiled up under the head by small muscles when it is not being used to suck up nectar from flowers.[24][25]: 560 [26]
The larvae, calledcaterpillars, have a toughened head capsule. Caterpillars lack the proboscis and have separate chewingmouthparts.[24] These mouthparts, calledmandibles, are used to chew up the plant matter that the larvae eat. The lower jaw, or labium, is weak, but may carry aspinneret, an organ used to create silk. The head is made of large lateral lobes, each having a group of up to six simple eyes.[25]: 562–563
The thorax is made of three fused segments, theprothorax,mesothorax, andmetathorax, each with a pair of legs. In some males of the butterfly familyNymphalidae, the forelegs are greatly reduced and are not used for walking or perching.[25]: 586 Lepidoptera have olfactory organs on their feet, which help them to "taste" or "smell" their food.[27] The larva has 3 pairs of true legs, with up to 11 pairs of abdominal legs and hooklets.[13]
The two pairs of wings are on the middle and third segments, ormesothorax andmetathorax, respectively. In the more recent genera, the wings of the second segment are much more pronounced, while some more primitive forms have similarly sized wings of both segments. The wings are covered in scales arranged like shingles, which form an extraordinary variety of colors and patterns. The mesothorax has more powerful muscles to propel the moth or butterfly through the air, with the wing of this segment (forewing) having a stronger vein structure.[25]: 560 The largest superfamily, theNoctuoidea, has its wings modified ashearing organs.[28]
The caterpillar has an elongated, soft body that may have hair-like or other projections, three pairs of true legs, with none to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[13] The thorax usually has a pair of legs on each segment. The thorax is also lined with many spiracles on both the mesothorax and metathorax, except for a few aquatic species, which instead have a form ofgills.[25]: 563
The abdomen, less sclerotized than the thorax, consists of 10 segments, the first sometimes reduced. The last two segments form the externalgenitalia. The genitalia are highly varied, and distinguish the species.[29] The females of basal moths have only one sex organ, which is used forcopulation and as anovipositor, or egg-laying organ. About 98% of moth species have a separate organ for mating, and an external duct that carries the sperm from the male.[25]: 561
The abdomen of the caterpillar has four pairs of prolegs, normally located on the third to sixth segments of the abdomen, and a separate pair of prolegs by the anus, which have a pair of tiny hooks called crotchets. These aid in gripping and walking, especially in species that lack many prolegs (e. g. larvae ofGeometridae). In some basal moths, these prolegs may be on every segment of the body, while prolegs may be completely absent in other groups more adapted to boring (e. g.,Prodoxidae) or living in sand (e.g.Nepticulidae).[25]: 563
Wing scales form the color and pattern on wings. The scales shown here are lamellar. The pedicel can be seen attached to a few loose scales.
Much of the body is covered with minute scales. Most arelamellar, or blade-like, and attached with a pedicel.[30] The surface of the lamella has a complex structure. It gives color throughstructural coloration, and may also be pigmented.[31]Scales function in insulation, thermoregulation, producingpheromones (in males only),[32] and aiding gliding flight, but the most important is the large diversity ofpatterns they create, enablingcamouflage,mimicry, signalling to rivals, andpotential mates.[30]
In thereproductive system of butterflies and moths, the malegenitalia are complex. In females the three types of genitalia are named for the associated moth clades: 'monotrysian', 'exoporian', and 'ditrysian'. In the monotrysian type there is an opening on the fused segments of the sterna 9 and 10, which allows insemination and oviposition. In the exoporian type (inHepialoidea andMnesarchaeoidea) there are separate places for insemination and oviposition, both on the same sterna as the monotrysian type, i.e. 9 and 10.[23] The ditrysian groups have an internal duct that carries sperm, with separate openings for copulation and egg-laying.[4] In most species, the genitalia are flanked by two soft lobes, although they may be specialized and sclerotized in some species for ovipositing in areas such as crevices and inside plant tissue.[23]The glands that produce hormones drive development as the insects go through their life cycles. The first insect hormoneprothoracicotropic hormone (PTTH) operates the species life cycle anddiapause.[33] It is produced bycorpora allata andcorpora cardiaca, where it is stored.[34]: 65, 75
In thedigestive system, the front of the foregut is modified to form a pharyngeal sucking pump for feeding on liquids such as nectar. Theesophagus leads to the posterior of the pharynx and in some species forms a form of crop. The midgut is short and straight, while the hindgut is longer and coiled.[23] Ancestral lepidopterans, stemming fromHymenoptera, had midgut ceca, not present in current butterflies and moths. Instead, digestive enzymes, other than initial digestion, are immobilized at the surface of the midgut cells. In larvae, long-necked and stalkedgoblet cells are found in the anterior and posterior midgut regions, respectively. The goblet cells excrete positivepotassium ions, absorbed from leaves ingested by the larvae. Most lepidopterans display the usual digestive cycle, sometimes adapted for specialised diets.[25]: 279 Some, like theluna moth, exhibit no digestive system whatsoever; they survive as adults from stored energy consumed as larvae and live for no longer than 7–10 days.[35]
Internal morphology of adult male in the familyNymphalidae, showing most of the major organ systems, with characteristic reduced forelegs of that family: The corpora include the corpus allatum and the corpus cardiaca.
Thecirculatory system useshemolymph, insect blood, forthermoregulation, taking heat from the muscles and transferring it to the rest of the body.[36] In lepidopteran species, hemolymph is circulated through the veins in the wings by some form of pulsating organ, either by the heart or by the intake of air into thetrachea.[34]: 69
Therespiratory system takes in air throughspiracles along the sides of the abdomen and thorax, supplying thetracheae with oxygen. Three different tracheae supply and diffuse oxygen throughout the species' bodies. The dorsal tracheae supply oxygen to the dorsal musculature and vessels, while the ventral tracheae supply the ventral musculature and nerve cord, and the visceral tracheae supply the guts, fat bodies, and gonads.[34]: 71, 72
Polymorphism is the appearance of forms or morphs which differ in color and other attributes within a species.[13]: 163 [37] In Lepidoptera, polymorphism can be seen both between individuals and between the sexes assexual dimorphism, between geographically separated populations, and between generations flying at different seasons of the year (seasonal polymorphism orpolyphenism). Polymorphism can be limited to one sex, typically the female.[13]
Environmental polymorphism, in whichtraits are not inherited, is often termed polyphenism, commonly seen in seasonal morphs, especially in the butterfly families ofNymphalidae andPieridae. An Old World pierid butterfly, the common grass yellow (Eurema hecabe) has a darker summer adult morph, triggered by a day of over 13 hours, while a shorter day induces a paler morph after the monsoon.[38] Polyphenism also occurs in caterpillars, as in the peppered moth,Biston betularia.[39]
Geographical isolation causes a divergence of a species into different morphs. A good example is the Indian white admiralLimenitis procris, which has five forms, each geographically separated from the other by large mountain ranges.[40]: 26 A more dramatic showcase of geographical polymorphism is theApollo butterfly (Parnassius apollo). Because the Apollos live in small local populations, with no contact with each other, interbreeding between populations practically never occurs. They therefore form over 600 different morphs, with the size of spots on the wings varying greatly.[41]
Seasonal diphenism in the common grass yellow,Eurema hecabe
Dry-season form
Wet-season form
Sexual dimorphism is the occurrence of differences between males and females. In Lepidoptera, it is widespread and almost completely set genetically.[38] It is present in all families of the Papilionoidea and more prominent in theLycaenidae,Pieridae, and some of theNymphalidae. Apart from color variation, secondary sexual characteristics may be present.[40]: 25 Different genotypes may be expressed at the same time.[38] Polymorphic and/or mimetic females occur in some taxa in thePapilionidae to obtain a level of protection not available to the male of their species. The most distinct case of sexual dimorphism is that of adult females of manyPsychidae which have only vestigial wings, legs, and mouthparts, whereas the adult males that are strong fliers with well-developed wings and feathery antennae.[42]
Mating pair ofLaothoe populi (poplar hawk-moth) showing two different color variants
Species of Lepidoptera undergoholometabolism or "complete metamorphosis". Their life cycle normally consists of anegg, alarva, apupa, and animago or adult.[13] The polypod larvae are commonly calledcaterpillars; the pupae of moths encapsulated in silk are calledcocoons, while the uncovered pupa of butterflies is called a chrysalis (plural: chrysalides).[29]: 165–167
Males usually starteclosion (emergence) earlier than females and peak in numbers before females. Both of the sexes are sexually mature by the time of eclosion.[25]: 564 Butterflies and moths are relatively asocial, except for migrating species. Mating begins with an adult attracting a mate, normally using visual stimuli, especially indiurnal species like most butterflies. However, the females of most nocturnal species, including almost all moth species, usepheromones to attract males, sometimes from long distances.[13] Some species attract mates using sound, as in the polka-dot wasp moth,Syntomeida epilais.[43]
Adaptations include having one, two or several seasonal generations, calledvoltinism. Most lepidopterans intemperate climates are univoltine, while in tropical climates most have two seasonal broods. Others may mate at any time of year. These adaptations are controlled by hormones, with delays in reproduction calleddiapause.[25]: 567 Many species die soon after mating and laying their eggs, having only lived for a few days after eclosion. Others may still be active for several weeks, overwinter indiapause, and become active again when the weather becomes more favorable.[25]: 564
Lepidoptera usually reproduce sexually and areoviparous (egg-laying), though some give birth to live young (ovoviviparity). Some species with larvae that can eat a variety of plants simply drop their eggs in flight, as inhepialids.[44] Most lay their eggs near or on the host, the plant on which the larvae feed. The number of eggs laid varies from a few to several thousand.[13] The females select the host plant instinctively, primarily by chemical cues.[25]: 564
The eggs are derived from materials ingested as a larva and in some species, from the spermatophores received from males during mating.[45] An egg can only be 1/1000 the mass of the female, yet she may lay up to her own mass in eggs. Females lay smaller eggs as they age. Larger females lay larger eggs.[46] The egg is covered by a hard-ridged protective outer layer of shell, called thechorion. It is lined with a thin coating ofwax, which prevents the egg from drying out.[47] Each egg hasmicropyles, tiny funnel-shaped openings at one end, to allow sperm to enter and fertilize the egg.[47][48] Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate.[47]
The egg stage lasts a few weeks in most butterflies, but eggs laid prior to winter, especially intemperate regions, go throughdiapause and hatching is delayed until spring.[49] Other temperate region butterflies lay their eggs in the spring and have them hatch in the summer (e. g.Nymphalis antiopa).[50]
Lepidopteran larvae typically live and feed on plants.
The larvae or caterpillars are the first stage in the life cycle after hatching. Caterpillars are "characteristicpolypod larvae with cylindrical bodies, short thoracic legs, and abdominal prolegs (pseudopods)".[51] They have a sclerotized head capsule,mandibles (mouthparts) for chewing, and a soft tubular body. The body has three pairs of true legs, and up to five pairs ofprolegs; there may be hairs or other projections.[52] The body consists of thirteen segments, of which three are thoracic and ten are abdominal.[53] Most larvae areherbivores, but a few arecarnivores ordetritivores.[52]
Different herbivorous species have adapted to feed on every part of the plant and are normally considered pests to their host plants; some species lay their eggs on fruit, others on clothing or fur (e. g.,Tineola bisselliella, the clothes moth). Some are carnivorous, and others are parasitic. Somelycaenid species such asPhengaris rebeli aresocial parasites ofMyrmica ant nests.[54] A species ofGeometridae fromHawaii has carnivorous larvae that catch and eat flies.[55] Some pyralid caterpillars are aquatic.[56] The larvae develop rapidly with several generations in a year; however, some species may take up to 3 years to develop, and exceptional examples likeGynaephora groenlandica take as long as seven years.[13]
The larvae of some butterflies and moths use mimicry to deter predators. Some caterpillars can inflate parts of their heads to appear snake-like. Many have false eye-spots to enhance this effect. Some have structures calledosmeteria (familyPapilionidae), exposed to produce foul-smelling defensive chemicals. Host plants often contain toxic substances; caterpillars can sequester and retain these into the adult stage. This helps make them unpalatable to birds and other predators. Such unpalatability is advertised using bright red, orange, black, or whitewarning colors. The toxic chemicals in plants are often evolved specifically to protect them from being eaten by insects. Insects, in turn, develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants.[57]
No form of wing is externally visible on the larva, but when larvae are dissected, developing wings can be seen as disks, which can be found on the second and third thoracic segments, in place of the spiracles that are apparent on abdominal segments. Wing disks develop in association with a trachea that runs along the base of the wing, and are surrounded by a thin peripodial membrane, which is linked to the outer epidermis of the larva by a tiny duct. Wing disks are very small until the last larval instar, when they increase dramatically in size, are invaded by branchingtracheae from the wing base that precede the formation of the wing veins and begin to develop patterns associated with several landmarks of the wing.[58]
Near pupation, the wings are forced outside the epidermis under pressure from thehemolymph, and although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval cuticle, they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled without damage to the wings.[58]
After about five to seven instars, or molts, certain hormones, like PTTH, stimulate the production ofecdysone, which initiates insect molting. The larva starts to develop into thepupa: body parts specific to the larva, such as the abdominal prolegs, degenerate, while others such as the legs and wings undergo growth. After finding a suitable place, the animal sheds its last larval cuticle, revealing the pupal cuticle underneath.[59]: 26–28
The pupa may be covered in a silk cocoon, attached to a substrate, buried, or left uncovered. Features of the adult are externally recognizable in the pupa. All the appendages on the adult head and thorax are found cased inside the cuticle (antennae,mouthparts, etc.), with the wings wrapped around, adjacent to the antennae.[25]: 564 The pupae of some species have functional mandibles.[24]
Although the pupal cuticle is highly sclerotized, some of the lower abdominal segments are not fused, and are able to move using small muscles found in between the membrane. Moving may help the pupa, for example, escape the sun, which would otherwise kill it. The pupa of theMexican jumping bean moth (Cydia saltitans) does this. The larvae cut a trapdoor in a bean (Sebastiania) and use it as a shelter. With a sudden rise in temperature, the pupa inside twitches and jerks, pulling on the threads inside. Wiggling may help to deterparasitoid wasps from laying eggs on the pupa. Other moths make clicks to deter predators.[25]: 564, 566
The length of time before the pupa emerges varies greatly. The monarch butterfly may stay in its chrysalis for two weeks, while other may stay in diapause for more than 10 months. The adult emerges either by using abdominal hooks or from projections on the head. The mandibles of the most primitive moth families are used to escape from their cocoon (e. g.,Micropterigoidea).[13][25]: 564
Most lepidopteran species do not live long after eclosion, only needing a few days to find a mate and then lay their eggs. Others may remain active for a longer period (from one to several weeks) or go through diapause and overwintering as monarch butterflies do, or waiting out environmental stress. Some adult species of microlepidoptera go through a stage where no reproductive-related activity occurs, lasting through summer and winter, followed by mating and oviposition in the early spring.[25]: 564
Flight is used for evading predators, searching for food, and finding mates. It is the main form of locomotion in most species. The forewings and hindwings aremechanically coupled and flap in synchrony. Flight is driven primarily by action of the forewings. Although lepidopteran species reportedly can still fly when their hindwings are cut off, it reduces their linear flight and turning capabilities.[61]
Some butterflies can reach fast speeds, such as thesouthern dart, which can go as fast as 48.4 kilometres per hour (30.1 mph). Sphingids are some of the fastest flying insects, some are capable of flying at over 50 kilometres per hour (31 mph), having a wingspan of 35–150 millimetres (1.4–5.9 in).[3][62] In some species, sometimes a gliding component to their flight exists. Flight occurs either as hovering, or as forward or backward motion.[63] In species such ashawk moths, hovering is important as they need to maintain a certain stability over flowers when feeding on the nectar.[3]
Long exposure image of flying moths, attracted to the floodlights
Navigation is important to lepidopterans, especially those that migrate. Butterflies navigate using time-compensated sun compasses. They can seepolarized light, so can orient even in cloudy conditions.[64] Most migratory butterflies live in semiarid areas where breeding seasons are short.[65] The life histories of their host plants also influence the strategies of the butterflies.[66] Lepidoptera may use coastal lines, mountains, and even roads to orient themselves. Above sea, the flight direction is much more accurate if the coast is still visible.[67]
Many moths use theEarth's magnetic field to navigate.[68] Thesilver Y can correct its course with changing winds, even at high altitude, and prefers flying with favourable winds.[69][70]Aphrissa statira inPanama loses its navigational capacity when exposed to a magnetic field, suggesting it uses the Earth's magnetic field.[71]
Moths tend to circle artificial lights. This suggests they use a technique ofcelestial navigation calledtransverse orientation. By maintaining a constant angular relationship to a bright celestial light, such as theMoon, they can fly in a straight line. When a moth encounters a much closer artificial light and uses it for navigation, the angle changes rapidly; the moth attempts to correct by turning toward the light, causing it to plummet downwards, and at close range, which results in a spiral flight path.[72]
Lepidopteran migration is typicallyseasonal, as the insects moving to escape dry seasons or other disadvantageous conditions. Most lepidopterans that migrate are butterflies, and the distance travelled varies. Some butterflies that migrate include themourning cloak,painted lady,American lady,red admiral, and thecommon buckeye.[59]: 29–30 A notable species of moth that migrates long distances is thebogong moth.[73] The most well-known migrations are those of the eastern population of themonarch butterfly from Mexico to northern United States and southern Canada, a distance of about 4,000–4,800 km (2,500–3,000 mi). Other well-known migratory species include the painted lady and several of thedanaine butterflies. Spectacular and large-scale migrations associated with themonsoons are seen in peninsular India.[74] Migrations have been studied in more recent times using wing tags and stable hydrogen isotopes.[75][76]
Moths such as theuraniids migrate.Urania fulgens undergoes population explosions and massive migrations that may be not surpassed by any other insect in theNeotropics. InCosta Rica andPanama, the first population movements may begin in July and early August and depending on the year, may be very massive, continuing unabated for as long as five months.[77]
Pheromones are involved in mating rituals among species, especially moths, but they are also an important aspect of other forms of communication. Usually, the pheromones are produced by one sex and detected by members of the opposite sex with their antennae.[78] In many species, a gland between the eighth and ninth segments under the abdomen in the female produces the pheromones.[13] Communication can also occur through stridulation, or producing sounds by rubbing parts of the body together.[70]
Moths engage in acoustic communication, most often as courtship, attracting mates using sound or vibration. They pick up these sounds using tympanic membranes in the abdomen.[79] An example is that of thepolka-dot wasp moth (Syntomeida epilais), which produces sounds with a frequency above that normally detectable by humans (about 20 kHz). These sounds also function as tactile communication, or communication through touch, as they stridulate, or vibrate a substrate like leaves and stems.[43]
Most moths lack bright colors, as many species use coloration ascamouflage, but butterflies engage in visual communication. Femalecabbage butterflies, for example, use ultraviolet light to communicate, with scales colored in this range on the dorsal wing surface. When they fly, each down stroke of the wing creates a brief flash of ultraviolet light which the males apparently recognize as the flight signature of a potential mate. These flashes from the wings may attract several males that engage in aerial courtship displays.[79]
Moth and butterfly caterpillars eat flowering plants, and are eaten by birds and parasitic insects. Adults are eaten by predators in many groups, such as birds, small mammals, and reptiles.[25]: 567
Papilio machaon caterpillar showing theosmeterium, which emits unpleasant smells to ward off predatorsThis Lepidoptera larvae disposes of itsfrass that might attract predators or parasites.
Lepidopterans are soft bodied, fragile, and almost defenseless, while the immature stages move slowly or are immobile, hence all stages are exposed topredation. Adult butterflies and moths are preyed upon bybirds,bats,lizards,amphibians,dragonflies, andspiders.[80]
Some species are poisonous, such as the monarch butterfly,Atrophaneura species, as well asPapilio antimachus, and thebirdwings. They obtain their toxicity by sequestering chemicals from the plants they eat. Some Lepidoptera manufacture their own toxins. Predators that eat poisonous butterflies and moths may become sick and vomit violently, learning not to eat those species, and other species with similar markings.[81][82] Toxic butterflies and larvae tend to develop bright colors and striking patterns to warn predators about their toxicity (aposematism).[83] Some caterpillars, especially members ofPapilionidae, contain anosmeterium, a Y-shaped protrusiblegland found in theprothoracic segment of the larvae.[81] When threatened, the caterpillar emits unpleasant smells from the organ to ward off the predators.[84][85]
Camouflage is an important defense strategy. Some lepidopteran species blend with their surroundings, making them difficult to spot by predators. Caterpillars can exhibit shades of green that match its host plant. Caterpillars can detect the color of their surroundings using organs on their feet.[86] Some caterpillars look like inedible objects, such as twigs or leaves, while those of some species, such as the common Mormon (Papilio polytes) and thewestern tiger swallowtail look like bird droppings.[81][87] For example, adultSesiidae species looksufficiently similar to awasp orhornet to ward off predators.[88]Eyespots are a type ofautomimicry used by some species, deflecting predators to attack these wing patterns.[89]
Batesian andMüllerian mimicry complexes are commonly found in Lepidoptera. In Batesian mimicry, an edible species (the mimic) gains a survival advantage by resembling inedible species (the model). In Müllerian mimicry, two or more inedible species benefit by resembling each other so as to reduce the sampling rate by predators that need to learn about the insects' inedibility. Taxa from the toxic genusHeliconius form a well-known Müllerian complex.[90]
Moths can hear ultrasound emitted by bats, which causes flying moths to make evasive maneuvers because bats are a main predator of moths; a reflex action innoctuid moths cause them to drop a few inches in flight to evade attack.[91]Tiger moths in a defense emit clicks within the same range of the bats, which interfere with the bats and foil their attempts to echolocate it.[92]
Some species of Lepidoptera engage in some form ofentomophily (more specifically psychophily and phalaenophily for butterflies and moths, respectively), or thepollination of flowers.[93] Most adult butterflies and moths feed on thenectar inside flowers, using their probosces to reach the nectar hidden at the base of the petals. In the process, the adults may brush against the flowers'stamens, on which the reproductivepollen is made and stored. The pollen is transferred on appendages on the adults, which fly to the next flower to feed and unwittingly deposit the pollen on thestigma of the next flower, where the pollengerminates and fertilizes the seeds.[25]: 813–814
Flowers pollinated by butterflies tend to be large, flamboyant, and scented, as butterflies are typically day-flying. Since butterflies do notdigest pollen (except forheliconid species,[93]) more nectar is offered than pollen. The flowers have simple nectar guides, with the nectaries usually hidden in narrow tubes or spurs, reached by the long "tongue" of the butterflies. Butterflies such asThymelicus flavus have been observed to engage inflower constancy, which means they are more likely to transfer pollen to other conspecific plants, because it reduces the loss of pollen during foraging, and the stigmas receive less pollen from other flower species.[94] However, most butterflies are very poor pollinators despite high rates of floral visitation; for example,monarch butterflies contribute almost nothing to seed set inmilkweeds that they visit for nectar.[95]
Among the more important moth pollinator groups are thehawk moths of thefamily Sphingidae. Their behavior is similar tohummingbirds, i.e., using rapid wing beats to hover in front of flowers. Most hawk moths arenocturnal orcrepuscular, so moth-pollinated flowers (e.g.,Silene latifolia ) tend to be white, night-opening, large, and showy with tubularcorollae and a strong, sweet scent produced in the evening, night, or early morning. A lot of nectar is produced to fuel the highmetabolic rates needed to power their flight.[96] Other moths (e.g.,noctuids,geometrids,pyralids) fly slowly and settle on the flower. They do not require as much nectar as the fast-flying hawk moths, and the flowers tend to be small (though they may be aggregated in heads).[97]
Mutualism is a form ofbiological interaction wherein each individual involved benefits in some way. An example of a mutualistic relationship would be that shared byyucca moths (Tegeculidae) and their host,yucca (Asparagaceae). Female yucca moths enter the flowers, collect the pollen into a ball using specialized maxillary palps, then move to the apex of the pistil, where pollen is deposited on the stigma, and lay eggs into the base of the pistil where seeds will develop. The larvae develop in the fruit pod and feed on a portion of the seeds. Thus, both insect and plant benefit, forming a highly mutualistic relationship.[25]: 814 Another form of mutualism occurs between larvae of Lycaenid butterflies andants. The larvae communicate with the ants using vibrations and chemical signals.[98] The ants provide some degree of protection to these larvae and they in turn gatherhoneydew secretions.[99]
Only 42 species ofparasitoid lepidopterans are known (1Pyralidae; 40Epipyropidae).[25]: 748 The larvae of thegreater andlesser wax moths feed on thehoneycomb insidebee nests and may becomepests; they are also found inbumblebee andwasp nests, albeit to a lesser extent. In northern Europe, the wax moth is regarded as the most serious parasitoid of the bumblebee and is found only in bumblebee nests. In some areas in southern England, as much as 80% of nests can be destroyed.[100] Other parasitic larvae are known to prey uponcicadas andleaf hoppers.[101]
In reverse, moths and butterflies may be subject toparasitic wasps andflies, which may lay eggs on the caterpillars, which hatch and feed inside its body, resulting in death. Although, in a form of parasitism called idiobiont, the adult paralyzes the host, so as not to kill it but for it to live as long as possible, for the parasitic larvae to benefit the most. In another form of parasitism, koinobiont, the species live off their hosts while inside (endoparasitic). These parasites live inside the host caterpillar throughout its life cycle or may affect it later on as an adult. In other orders, koinobionts include flies, a majority ofcoleopteran, and manyhymenopteran parasitoids.[25]: 748–749 Some species may be subject to a variety of parasites, such as thespongy moth (Lymantaria dispar), which is attacked by a series of 13 species, in six different taxa throughout its life cycle.[25]: 750
In response to a parasitoid egg or larva in the caterpillar's body, theplasmatocytes, or simply the host's cells can form a multilayered capsule that eventually causes the endoparasite toasphyxiate. The process, called encapsulation, is one of the caterpillar's only means of defense against parasitoids.[25]: 748
Among the first entomologists to study fossil insects and their evolution wasSamuel Hubbard Scudder (1837–1911), who worked on butterflies. He published a study of theFlorissant deposits of Colorado, including the exceptionally preservedProdryas persephone.Andreas V. Martynov (1879–1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.[108]
Major contributions in the 20th century included the creation of the Monotrysia and Ditrysia (based on female genital structure) by Borner in 1925 and 1939.[102]Willi Hennig (1913–1976) developed thecladistic methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships amongmonotrysian families.[102][108] 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 notmonophyletic: Microlepidoptera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.[102] A 2024 genetic study found that thechromosome organisation of butterflies and moths has been largely conserved in most groups over the past 250 million years.[109]
The fossil record for Lepidoptera is lacking in comparison to other winged species and tends not to be as common as some other insects in habitats that are most conducive to fossilization, such as lakes and ponds; their juvenile stage has only the head capsule as a hard part that might be preserved. Also, the scales covering their wings arehydrophobic and prevents their body from sinking when they end up on the water's surface.[110] Lepidopteran bodies tend to come apart after death, and decompose quickly, so fossil remains are often extremely fragmentary. Of the fossils known, only an estimated 7% have been described.[111] The location and abundance of the most common moth species are indicative that mass migrations of moths occurred over the PalaeogeneNorth Sea, which is why there is a serious lack of moth fossils.[112] Yet there are fossils, some preserved in amber and some in very fine sediments.Leaf mines are also seen in fossil leaves, although the interpretation of them is tricky.[108]
Putative fossil stem group representatives ofAmphiesmenoptera (the clade comprising Trichoptera and Lepidoptera) are known from theTriassic.[25]: 567 The earliest known probable lepidopteran fossils are fossilized scales found in approximately 236 million years old fossilized herbivore dung in Argentina that are thought to have been incidentally ingested.[113] The next oldest fossils are from theTriassic-Jurassic boundary (ca 201 million years ago). They were found as rarepalynological elements in the sediments of the Triassic-Jurassic boundary from the cored Schandelah-1 well, drilled nearBraunschweig in northernGermany. This pushes back the fossil record and origin ofglossatan lepidopterans by about 70-90 million years, supporting molecular estimates of aNorian (ca 212 million years) divergence of glossatan and non-glossatan lepidopterans. The findings were reported in 2018 in the journalScience Advances. The authors of the study proposed that lepidopterans evolved a proboscis as an adaptation to drink from droplets and thin films of water for maintaining theirfluid balance in the hot and aridclimate of theTriassic.[114]
The earliest named lepidopteran taxon isArchaeolepis mane, a primitive moth-like species from theEarly Jurassic, dated back to around190 million years ago, and known only from three wings found in theCharmouth Mudstone ofDorset,UK. The wings show scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared withTrichoptera (caddisflies).[115][116] Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from theCretaceous, which all belong to primitive moth-like families.[108]
Phylogenetic hypothesis of major lepidopteran lineages superimposed on the geologic time scale. Radiation ofangiosperms spans130 to 95 million years ago from their earliest forms to domination of vegetation.
Lepidoptera andTrichoptera (caddisflies) aresister groups, sharing many similarities that are lacking in others; for example the females of both orders areheterogametic, meaning they have two differentsex chromosomes, whereas in most species the males are heterogametic and the females have two identical sex chromosomes. The adults in both orders display a particular wing venation pattern on their forewings. The larvae in the two orders have mouth structures and glands with which they make and manipulatesilk.Willi Hennig grouped the two orders into the superorderAmphiesmenoptera; together they are sister to the extinct orderTarachoptera.[119] Lepidoptera descend from a diurnal moth-like common ancestor that either fed on dead or living plants.[120]
Thecladogram, based on molecular analysis, shows the order as aclade, sister to the Trichoptera, and more distantly related to theDiptera (true flies) andMecoptera (scorpionflies).[121]
The internal phylogeny of Lepidoptera is still being resolved. While many large clades have been established, interfamilial and superfamilial relationships are poorly understood. A large scale study by Regier et al. attempts to resolve these relationships using three different analysis methods, which is shown in the following cladogram.[122]
Micropterigidae,Agathiphagidae andHeterobathmiidae are the oldest and mostbasal lineages of Lepidoptera. The adults of these families do not have the curled tongue orproboscis, that are found in most members of the order, but instead have chewingmandibles adapted for a special diet. Micropterigidae larvae feed onleaves,fungi, orliverworts (much like theTrichoptera).[102] Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live insidekauri pines and feed on seeds. In Heterobathmiidae the larvae feed on the leaves ofNothofagus, the southern beech tree. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon aftermetamorphosis.[102]
With the evolution of theDitrysia in the mid-Cretaceous, there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is onecloaca, or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia,Gracillarioidea andGelechioidea are mostly leaf miners, but more recent lineages feed externally. In theTineoidea, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.[108]
In the so-called "macrolepidoptera", which constitutes about 60% of lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant.[108] Many havetympanal organs, that allow them to hear.[108]The main lineages in the macrolepidoptera are theNoctuoidea,Bombycoidea,Lasiocampidae,Mimallonoidea,Geometroidea andRhopalocera. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be amonophyletic group.[108] The Rhopalocera, comprising thePapilionoidea (butterflies),Hesperioidea (skippers), and theHedyloidea (moth-butterflies), are the most recently evolved.[102] There is quite a good fossil record for this group, with the oldest skipper dating from56 million years ago.[108]
There are over 120 recognized lepidopteran families, in 45 to 48 superfamilies. Historically, lepidopterists classified the order into five suborders, with one encompassing the so-called "primitive" moths—groups that were long viewed as relics, retaining morphological features reminiscent of their distant ancestors. This portrayed these lineages as static holdovers from the past. In contrast, the vast majority of species fall within the dominant suborderDitrysia, characterized by "advanced" reproductive and morphological traits. Phylogenetic analyses show that these early-diverging lineages areparaphyletic relative to later lineages. Consequently, lepidopterists have largely abandoned intermediate clades such as suborders.[25]: 569
The basal clades of Lepidoptera represent early-diverging lineages that retain ancestral traits, providing key insights into the order's evolutionary origins. These groups are paraphyletic and form successive sister relationships leading to the dominant Glossata. Below is an overview of the major clades:
This clade comprises the single superfamilyMicropterigoidea, one of the earliest-diverging lineages, with adults uniquely retaining functional chewing mouthparts (mandibles). Approximately 120 species are known worldwide.[25]: 569
This clade contains a single small family,Agathiphagidae. These moths lack a functional proboscis, reflecting their early divergence, and their caterpillars feed exclusively on the wood of kauri trees (genusAgathis).[25]: 569 [123][124][125]
This clade includes the single familyHeterobathmiidae, with around 10 species of day-flying, metallic moths restricted to southern South America. Adults feed on pollen fromNothofagus (southern beech) trees, while the larvae mine leaves. They lack a functional proboscis.[25]: 569 [126]
Encompassing most lepidopteran species, this clade is distinguished by the loss of functional mandibles and the development of elongated maxillary galeae forming a proboscis for liquid feeding.[25]: 569
Butterflybrooch from the Russian court in gold, silver, diamond, and sapphire. 19th century
Artistic depictions of butterflies have been used in cultures from at least 3500 years ago, in Egyptian hieroglyphs.[127] Butterflies are widely used in various objects of art and jewelry: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and in some mixed media artworks and furnishings.[128]
In many cultures the soul of a dead person is associated with the butterfly, for example inAncient Greece, where the word for butterflyψυχή (psyche) also meanssoul andbreath. InLatin, as in Ancient Greece, the word for "butterfly"papilio was associated with the soul of the dead.[129] According toKwaidan: Stories and Studies of Strange Things, byLafcadio Hearn, a butterfly was seen in Japan as thepersonification of a person's soul; whether they were living, dying, or already dead. A Japanese superstition says that if a butterfly enters your guestroom and perches behind thebamboo screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as a badomen. WhenTaira no Masakado was secretly preparing for his famous revolt, there appeared inKyoto so vast a swarm of butterflies that the people were frightened—thinking the apparition to be a portent of coming evil.[130]
In the ancientMesoamerican city ofTeotihuacan, the brilliantly colored image of the butterfly was carved into temples, buildings, jewelry, andincense burners. The butterfly was sometimes depicted with the maw of ajaguar; some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies tofire and warfare persisted through to theAztec civilization; jaguar-butterfly images have been found among theZapotec andMaya civilizations.[131]
Thelarvae of many lepidopteran species are major pests inagriculture. Some of the major pests includeTortricidae,Noctuidae, andPyralidae. The larvae of the Noctuidae genusSpodoptera (armyworms),Helicoverpa (corn earworm), orPieris brassicae can cause extensive damage to certain crops.[102]Helicoverpa zea larvae (cotton bollworms or tomato fruitworms) arepolyphagous, meaning they eat a variety of crops, includingtomatoes andcotton.[132]Peridroma saucia (variegated cutworms) are described as one of the most damaging pests to gardens, with the ability to destroy entire gardens and fields in a matter of days.[133]
Butterflies and moths are one of the largest taxa to solely feed and be dependent on living plants, in terms of the number of species, and they are in many ecosystems, making up the largest biomass to do so. In many species, the female may produce anywhere from 200 to 600 eggs, while in some others it may go as high as 30,000 eggs in one day. This can create many problems for agriculture, where many caterpillars can affect acres of vegetation. Some reports estimate that there have been over 80,000 caterpillars of several different taxa feeding on a single oak tree. In some cases, phytophagous larvae can lead to the destruction of entire trees in relatively short periods of time.[25]: 567
Ecological ways of removing pest Lepidoptera species are becoming more economically viable, as research has shown ways like introducing parasitic wasps and flies. For example,Sarcophaga aldrichi, a fly which deposited larvae feed upon the pupae of theforest tent caterpillar moth. Pesticides can affect other species other than the species they are targeted to eliminate, damaging the natural ecosystem.[134] Another good biological pest control method is the use ofpheromone traps. A pheromone trap is a type ofinsect trap that usespheromones to lureinsects. Sex pheromones and aggregating pheromones are the most common types used. A pheromone-impregnated lure is encased in a conventional trap such as a Delta trap,water-pan trap, or funnel trap.[135]
Even though some butterflies and moths affect the economy negatively, many species are a valuable economic resource. The most prominent example is that of thedomesticated silkworm moth (Bombyx mori), the larvae of which make their cocoons out ofsilk, which can be spun into cloth. Silk is and has been an important economic resourcethroughout history. The speciesBombyx mori has been domesticated to the point where it is completely dependent on mankind for survival.[137] A number of wild moths such asBombyx mandarina, andAntheraea species, besides others, provide commercially important silks.[138]
The preference of the larvae of most lepidopteran species to feed on a single species or limited range of plants is used as a mechanism for biological control ofweeds in place of herbicides. Thepyralidcactus moth was introduced from Argentina to Australia, where it successfully suppressed millions of acres ofprickly pear cactus.[25]: 567 Another species of the Pyralidae, called thealligator weed stem borer (Arcola malloi), was used to control theaquatic plant known asalligator weed (Alternanthera philoxeroides) in conjunction with thealligator weed flea beetle; in this case, the two insects work insynergy and the weed rarely recovers.[139]
Breeding butterflies and moths, orbutterfly gardening/rearing, has become an ecologically viable process of introducing species into the ecosystem to benefit it.Butterfly ranching in Papua New Guinea permits nationals of that country to "farm" economically valuable insect species for the collectors market in an ecologically sustainable manner.[140]
Beondegi,silkworm pupae steamed or boiled and seasoned for taste, for sale by a street vendor in South Korea
Lepidoptera feature prominently inentomophagy as food items on almost every continent. While in most cases, adults, larvae or pupae are eaten as staples by indigenous people,beondegi orsilkwormpupae are eaten as a snack inKorean cuisine[141] whileMaguey worm is considered a delicacy inMexico.[142] In some parts ofHuasteca, the silk nests of theMadrone butterfly are maintained on the edge of roof tops of houses for consumption.[143] In theCarnia region of Italy, children catch and eatingluvies of the toxicZygaena moths in early summer. The ingluvies, despite having a very low cyanogenic content, serve as a convenient, supplementary source of sugar to the children who can include this resource as a seasonal delicacy at minimum risk. Outside of this instance, adult Lepidoptera are rarely consumed by humans, with the sole exception of theBogong moth.[144]
Some larvae of both moths and butterflies have irritant toxic hairs; species from some 12 families can inflict serious human injuries (urticarial dermatitis andatopicasthma toosteochondritis, consumptioncoagulopathy,renal failure, andintracerebralhemorrhage).[145] Skin rashes are the most common, but there have been fatalities.[146]Lonomia is a frequent cause of envenomation in humans in Brazil, killing up to 20% of people affected.[147]
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