Lepidoptera

The term Lepidoptera was used in 1746 by Carl Linnaeus in his Fauna Svecica. The word is derived from Greek , gen. ("scale") and ("wing"). The word "butterfly" is from Old English '''', possibly from the pale butter color of some species' wings. The word "moth" is from Old English ' from Common Germanic and German ' meaning "moth". The word "caterpillar" is from Middle English , , probably an alteration of Old North French (from Latin , "cat" + , "hairy"). == Distribution and diversity ==

The Lepidoptera are among the most successful groups of insects. They are found on all continents, except Antarctica, 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). Among the most northern dwelling species of butterflies and moths is the Siberian Apollo (Parnassius arcticus), found in the Arctic Circle in northeastern Yakutia, at an altitude of above sea level. In the Himalayas, Apollo species such as Parnassius epaphus occur up to an altitude of . Some species are commensal or parasitic, inhabiting the bodies of organisms rather than the environment. Coprophagous pyralid moths, called sloth moths, such as Bradipodicola hahneli and Cryptoses choloepi, are unusual in that they are exclusively found in the fur of sloths. Two species of Tinea moths feed and breed on the horns of cattle. The larva of Zenodochium coccivorella is an internal parasite of Kermes scale insects. Many species breed in materials such as owl pellets, bat caves, honeycombs or diseased fruit. 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, while Australia has about 400 species of butterflies and 14,000 species of moths. The diversity in each faunal region has been estimated by John Heppner in 1991 based partly on actual counts from the literature, partly on the card indices in the Natural History Museum (London) and the National Museum of Natural History (Washington), and partly on estimates: == External morphology ==

Lepidoptera are morphologically distinguished from other orders by the presence of scales on the body and appendages, especially the wings. Butterflies and moths vary in size from microlepidoptera only a few millimeters long, to conspicuous animals with a wingspan greater than , such as Queen Alexandra's birdwing and Atlas moth. Head The head houses sensing organs and mouthparts. Like the adult, the larva has a toughened, or sclerotized head capsule. Here, two compound eyes, and chaetosema, 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. The maxillary galeae are modified to form an elongated proboscis. 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. The larvae, called caterpillars, have a toughened head capsule. Caterpillars lack the proboscis and have separate chewing mouthparts. The two pairs of wings are on the middle and third segments, or mesothorax and metathorax, 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. The caterpillar has an elongated, soft body that may have hair-like scales, three pairs of true legs, with none to 11 pairs of abdominal prolegs (usually eight) with hook-like spines for gripping called crochets. 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 of gills. The females of basal moths have only one sex organ, which is used for copulation and as an ovipositor, 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. The surface of the lamella has a complex structure. It gives color through structural coloration, and may also be pigmented. Scales function in insulation, thermoregulation, producing pheromones (in males only), and aiding gliding flight, but the most important is the large diversity of patterns they create, enabling camouflage, mimicry, signalling to rivals, and potential mates. == Internal morphology ==

Reproductive system In the reproductive system of butterflies and moths, the male genitalia 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 (in Hepialoidea and Mnesarchaeoidea) there are separate places for insemination and oviposition, both on the same sterna as the monotrysian type, i.e. 9 and 10. It is produced by corpora allata and corpora cardiaca, where it is stored. Digestive system In the digestive system, the front of the foregut is modified to form a pharyngeal sucking pump for feeding on liquids such as nectar. The esophagus 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. , showing most of the major organ systems, with characteristic reduced forelegs of that family: The corpora include the corpus allatum and the corpus cardiaca. Circulatory system The circulatory system uses hemolymph, insect blood, for thermoregulation, taking heat from the muscles and transferring it to the rest of the body. 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 the trachea. Respiratory system The respiratory system takes in air through spiracles along the sides of the abdomen and thorax, supplying the tracheae 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. == Polymorphism ==

'') mating: The female is flightless. Polymorphism is the appearance of forms or morphs which differ in color and other attributes within a species. In Lepidoptera, polymorphism can be seen both between individuals and between the sexes as sexual dimorphism, between geographically separated populations, and between generations flying at different seasons of the year (seasonal polymorphism or polyphenism). Polymorphism can be limited to one sex, typically the female. Geographical isolation causes a divergence of a species into different morphs. A good example is the Indian white admiral Limenitis procris, which has five forms, each geographically separated from the other by large mountain ranges. Sexual dimorphism is the occurrence of differences between males and females. In Lepidoptera, it is widespread and almost completely set genetically. It is present in all families of the Papilionoidea and more prominent in the Lycaenidae, Pieridae, and some of the Nymphalidae. Apart from color variation, secondary sexual characteristics may be present. Different genotypes may be expressed at the same time. == Reproduction and development ==

'' (poplar hawk-moth) showing two different color variants Species of Lepidoptera undergo holometabolism or "complete metamorphosis". Their life cycle normally consists of an egg, a larva, a pupa, and an imago or adult. Adaptations include having one, two or several seasonal generations, called voltinism. Most lepidopterans in temperate 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 called diapause. 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. 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. The egg is covered by a hard-ridged protective outer layer of shell, called the chorion. It is lined with a thin coating of wax, which prevents the egg from drying out. Each egg has micropyles, tiny funnel-shaped openings at one end, to allow sperm to enter and fertilize the egg. Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate. Other temperate region butterflies lay their eggs in the spring and have them hatch in the summer (e. g. Nymphalis antiopa). Larvae The larvae or caterpillars are the first stage in the life cycle after hatching. Caterpillars are "characteristic polypod larvae with cylindrical bodies, short thoracic legs, and abdominal prolegs (pseudopods)". 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 of prolegs; there may be hairs or other projections. Most larvae are herbivores, but a few are carnivores or detritivores. 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. Some lycaenid species such as Phengaris rebeli are social parasites of Myrmica ant nests. A species of Geometridae from Hawaii has carnivorous larvae that catch and eat flies. Some pyralid caterpillars are aquatic. The larvae develop rapidly with several generations in a year; however, some species may take up to 3 years to develop, and exceptional examples like Gynaephora groenlandica take as long as seven years. Wing development 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 branching tracheae from the wing base that precede the formation of the wing veins and begin to develop patterns associated with several landmarks of the wing. Near pupation, the wings are forced outside the epidermis under pressure from the hemolymph, 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. 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. == Behavior ==

Flight 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 are mechanically 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. Some butterflies can reach fast speeds, such as the southern dart, which can go as fast as . Sphingids are some of the fastest flying insects, some are capable of flying at over , having a wingspan of . In some species, sometimes a gliding component to their flight exists. Flight occurs either as hovering, or as forward or backward motion. In species such as hawk moths, hovering is important as they need to maintain a certain stability over flowers when feeding on the nectar. Most migratory butterflies live in semiarid areas where breeding seasons are short. The life histories of their host plants also influence the strategies of the butterflies. 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. Many moths use the Earth's magnetic field to navigate. The silver Y can correct its course with changing winds, even at high altitude, and prefers flying with favourable winds. Aphrissa statira in Panama loses its navigational capacity when exposed to a magnetic field, suggesting it uses the Earth's magnetic field. Moths tend to circle artificial lights. This suggests they use a technique of celestial navigation called transverse orientation. By maintaining a constant angular relationship to a bright celestial light, such as the Moon, 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. Migration , seen in a cluster in Santa Cruz, California, where the western population migrates for the winter Lepidopteran migration is typically seasonal, 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 the mourning cloak, painted lady, American lady, red admiral, and the common buckeye. The most well-known migrations are those of the eastern population of the monarch butterfly from Mexico to northern United States and southern Canada, a distance of about . Other well-known migratory species include the painted lady and several of the danaine butterflies. Spectacular and large-scale migrations associated with the monsoons are seen in peninsular India. Migrations have been studied in more recent times using wing tags and stable hydrogen isotopes. Moths such as the uraniids migrate. Urania fulgens undergoes population explosions and massive migrations that may be not surpassed by any other insect in the Neotropics. In Costa Rica and Panama, 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. Communication , Poland 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. In many species, a gland between the eighth and ninth segments under the abdomen in the female produces the pheromones. An example is that of the polka-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. Most moths lack bright colors, as many species use coloration as camouflage, but butterflies engage in visual communication. Female cabbage 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. == Ecology ==

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. An "evolutionary arms race" can be seen between predator and prey species. Strategies for defense and protection include aposematism, mimicry, camouflage, and deimatic displays. Some species are poisonous, such as the monarch butterfly, Atrophaneura species, as well as Papilio antimachus, and the birdwings. 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. Toxic butterflies and larvae tend to develop bright colors and striking patterns to warn predators about their toxicity (aposematism). Some caterpillars, especially members of Papilionidae, contain an osmeterium, a Y-shaped protrusible gland found in the prothoracic segment of the larvae. 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. Some caterpillars look like inedible objects, such as twigs or leaves, while those of some species, such as the common Mormon (Papilio polytes) and the western tiger swallowtail look like bird droppings. For example, adult Sesiidae species look sufficiently similar to a wasp or hornet to ward off predators. Eyespots are a type of automimicry used by some species, deflecting predators to attack these wing patterns. Batesian and Müllerian 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 genus Heliconius form a well-known Müllerian complex. 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 in noctuid moths cause them to drop a few inches in flight to evade attack. 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. Pollination drinking nectar from a species of Dianthus Some species of Lepidoptera engage in some form of entomophily (more specifically psychophily and phalaenophily for butterflies and moths, respectively), or the pollination of flowers. Most adult butterflies and moths feed on the nectar 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 reproductive pollen 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 the stigma of the next flower, where the pollen germinates and fertilizes the seeds. However, most butterflies are very poor pollinators despite high rates of floral visitation; for example, monarch butterflies contribute almost nothing to seed set in milkweeds that they visit for nectar. Among the more important moth pollinator groups are the hawk moths of the family Sphingidae. Their behavior is similar to hummingbirds, i.e., using rapid wing beats to hover in front of flowers. Most hawk moths are nocturnal or crepuscular, so moth-pollinated flowers (e.g., Silene latifolia ) tend to be white, night-opening, large, and showy with tubular corollae and a strong, sweet scent produced in the evening, night, or early morning. A lot of nectar is produced to fuel the high metabolic rates needed to power their flight. 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). Mutualism '') parasitized by Braconidae wasp larvae Mutualism is a form of biological interaction wherein each individual involved benefits in some way. An example of a mutualistic relationship would be that shared by yucca 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. The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions. Parasitism larva exits from a fox moth caterpillar Only 42 species of parasitoid lepidopterans are known (1 Pyralidae; 40 Epipyropidae). Other parasitic larvae are known to prey upon cicadas and leaf hoppers. In reverse, moths and butterflies may be subject to parasitic wasps and flies, 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 of coleopteran, and many hymenopteran parasitoids. Some species may be subject to a variety of parasites, such as the spongy moth (Lymantaria dispar), which is attacked by a series of 13 species, in six different taxa throughout its life cycle. In response to a parasitoid egg or larva in the caterpillar's body, the plasmatocytes, or simply the host's cells can form a multilayered capsule that eventually causes the endoparasite to asphyxiate. The process, called encapsulation, is one of the caterpillar's only means of defense against parasitoids. == History of study ==

, Bulgaria Linnaeus in Systema Naturae (1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx and Phalaena, with seven subgroups in Phalaena. and Edward Meyrick (1895) based their classifications primarily on wing venation. Sir George Francis Hampson worked on the microlepidoptera during this period and Philipp Christoph Zeller published The Natural History of the Tineinae also on microlepidoptera (1855). Among the first entomologists to study fossil insects and their evolution was Samuel Hubbard Scudder (1837–1911), who worked on butterflies. He published a study of the Florissant deposits of Colorado, including the exceptionally preserved Prodryas persephone. Andreas V. Martynov (1879–1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny. 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. == Evolution and systematics ==

Fossil record '', a fossil lepidopteran from the Eocene 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 are hydrophobic and prevents their body from sinking when they end up on the water's surface. 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. The location and abundance of the most common moth species are indicative that mass migrations of moths occurred over the Palaeogene North Sea, which is why there is a serious lack of moth fossils. 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. The next oldest fossils are from the Triassic-Jurassic boundary (ca 201 million years ago). They were found as rare palynological elements in the sediments of the Triassic-Jurassic boundary from the cored Schandelah-1 well, drilled near Braunschweig in northern Germany. This pushes back the fossil record and origin of glossatan lepidopterans by about 70-90 million years, supporting molecular estimates of a Norian (ca 212 million years) divergence of glossatan and non-glossatan lepidopterans. The findings were reported in 2018 in the journal Science 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 their fluid balance in the hot and arid climate of the Triassic. The earliest named lepidopteran taxon is Archaeolepis mane, a primitive moth-like species from the Early Jurassic, dated back to around , and known only from three wings found in the Charmouth Mudstone of Dorset, UK. The wings show scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared with Trichoptera (caddisflies). Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from the Cretaceous, which all belong to primitive moth-like families. Well-preserved fossil lepidopterans include the Eocene Prodryas persephone from the Florissant Fossil Beds. Phylogeny s spans from their earliest forms to domination of vegetation. Lepidoptera and Trichoptera (caddisflies) are sister groups, sharing many similarities that are lacking in others; for example the females of both orders are heterogametic, meaning they have two different sex 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 manipulate silk. Willi Hennig grouped the two orders into the superorder Amphiesmenoptera; together they are sister to the extinct order Tarachoptera. Lepidoptera descend from a diurnal moth-like common ancestor that either fed on dead or living plants. The cladogram, based on molecular analysis, shows the order as a clade, sister to the Trichoptera, and more distantly related to the Diptera (true flies) and Mecoptera (scorpionflies). }} 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. }} }} Micropterigidae, Agathiphagidae and Heterobathmiidae are the oldest and most basal lineages of Lepidoptera. The adults of these families do not have the curled tongue or proboscis, that are found in most members of the order, but instead have chewing mandibles adapted for a special diet. Micropterigidae larvae feed on leaves, fungi, or liverworts (much like the Trichoptera). Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live inside kauri pines and feed on seeds. In Heterobathmiidae the larvae feed on the leaves of Nothofagus, the southern beech tree. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon after metamorphosis. ; Heterobathmiina : This clade includes the single family Heterobathmiidae, with around 10 species of day-flying, metallic moths restricted to southern South America. Adults feed on pollen from Nothofagus (southern beech) trees, while the larvae mine leaves. They lack a functional proboscis. ; Glossata : 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. == Relationship to people ==

Culture 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. 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. In many cultures the soul of a dead person is associated with the butterfly, for example in Ancient Greece, where the word for butterfly ψυχή (psyche) also means soul and breath. In Latin, as in Ancient Greece, the word for "butterfly" papilio was associated with the soul of the dead. According to Kwaidan: Stories and Studies of Strange Things, by Lafcadio Hearn, a butterfly was seen in Japan as the personification 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 the bamboo screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as a bad omen. When Taira no Masakado was secretly preparing for his famous revolt, there appeared in Kyoto so vast a swarm of butterflies that the people were frightened—thinking the apparition to be a portent of coming evil. In the ancient Mesoamerican city of Teotihuacan, the brilliantly colored image of the butterfly was carved into temples, buildings, jewelry, and incense burners. The butterfly was sometimes depicted with the maw of a jaguar; some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies to fire and warfare persisted through to the Aztec civilization; jaguar-butterfly images have been found among the Zapotec and Maya civilizations. Pests '' caterpillar damage to a broccoli (Brassicaceae) The larvae of many lepidopteran species are major pests in agriculture. Some of the major pests include Tortricidae, Noctuidae, and Pyralidae. The larvae of the Noctuidae genus Spodoptera (armyworms), Helicoverpa (corn earworm), or Pieris brassicae can cause extensive damage to certain crops. 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. 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. Another good biological pest control method is the use of pheromone traps. A pheromone trap is a type of insect trap that uses pheromones to lure insects. 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. Species of moths that are detritivores would naturally eat detritus containing keratin, such as hairs or feathers. Well known species are cloth moths (T. bisselliella, T. pellionella, and T. tapetzella), feeding on foodstuffs that people find economically important, such as cotton, linen, silk and wool fabrics as well as furs; furthermore they have been found on shed feathers and hair, bran, semolina and flour (possibly preferring wheat flour), biscuits, casein, and insect specimens in museums. Beneficial insects Even though some butterflies and moths affect the economy negatively, many species are a valuable economic resource. The most prominent example is that of the domesticated silkworm moth (Bombyx mori), the larvae of which make their cocoons out of silk, which can be spun into cloth. Silk is and has been an important economic resource throughout history. The species Bombyx mori has been domesticated to the point where it is completely dependent on mankind for survival. A number of wild moths such as Bombyx mandarina, and Antheraea species, besides others, provide commercially important silks. 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 of weeds in place of herbicides. The pyralid cactus moth was introduced from Argentina to Australia, where it successfully suppressed millions of acres of prickly pear cactus. Breeding butterflies and moths, or butterfly 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. Food , silkworm pupae steamed or boiled and seasoned for taste, for sale by a street vendor in South Korea Lepidoptera feature prominently in entomophagy as food items on almost every continent. While in most cases, adults, larvae or pupae are eaten as staples by indigenous people, beondegi or silkworm pupae are eaten as a snack in Korean cuisine while Maguey worm is considered a delicacy in Mexico. In some parts of Huasteca, the silk nests of the Madrone butterfly are maintained on the edge of roof tops of houses for consumption. In the Carnia region of Italy, children catch and eat ingluvies of the toxic Zygaena 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 the Bogong moth. Health Some larvae of both moths and butterflies have irritant toxic hairs; species from some 12 families can inflict serious human injuries (urticarial dermatitis and atopic asthma to osteochondritis, consumption coagulopathy, renal failure, and intracerebral hemorrhage). Skin rashes are the most common, but there have been fatalities. Lonomia is a frequent cause of envenomation in humans in Brazil, killing up to 20% of people affected. == See also ==