Lepidoptera

(invertebrate zoology) Large order of scaly-winged insects, including the butterflies, skippers, and moths; adults are characterized by two pairs of membranous wings and sucking mouthparts, featuring a prominent, coiled proboscis.

(Butterflies, skippers, and moths)

Class: Insecta

Order: Lepidoptera

Number of families: 122

Evolution and systematics

For such a large group (arguably the second-largest group of insects, with approximately 150,000 described species), the fossil record for lepidopterans is meager. An estimated 600–700 fossil specimens are known, of which the earliest is a small moth, Archeolepis mane, from Dorset, England, dating from the early Jurassic period. Other fossils include leaf miners and preserved specimens in Cretaceous amber for primitive moths. More advanced families, including butterflies and noctuid moths, are known mostly from wings or preserved impressions of the insect; one of the most famous is a probable nymphalid butterfly, Prodryas persephone, from the rich shale Florisant fossil beds of the Oligocene period in Colorado. Fossils of immature stages are rare, but among them are a probable sphingid-like larva from the Pliocene in Germany and a possible noctuid egg from the late Cretaceous in eastern North America, for example. A factor contributing to the relative scarcity of fossil lepidopterans is their more fragile nature compared with other insect orders.

Lepidoptera is one of the two major orders (the other being the Diptera), along with scorpionflies, caddisflies, and fleas, forming the "panorpoid" complex. Trichoptera (caddisflies) is the sister group of Lepidoptera. Both possess either hairy or scaly wings, caterpillar-like larvae, reduction of mandibles in the adult stage, and similar wing venation. The Lepidoptera are distinguished by no fewer than 27 uniquely derived characters, including the possession of fleshy prolegs with hooklike crochets and a silk-producing spinneret in larvae, loss of the median ocellus, coiled sucking mouthparts, wings covered with a dense layer of overlapping shingle-like deciduous scales, and lack of cerci in adults.

The vast majority of lepidopterans are moths. Most moths have drab, somber colors; are nocturnal; do not have clubbed antennae; and often rest with their wings held rooflike over the back. Skippers (Hesperiidae) are small, mothlike, dayflying butterflies that have a clubbed antenna ending in a curved tip called the apiculus, and they usually hold their wings over their backs. Butterflies, with about 19,000 described species, represent only about 13% of all Lepidoptera, though they probably are the most popular group. They differ from skippers in not having an apiculus at the end of the antenna.

Although the Lepidoptera make up an easily definable group, the higher classification below the ordinal level has undergone significant changes in light of newer phylogenetic classification methods. For many years, such terms as Rhopalocera (butterflies excluding skippers), Heterocera (all moths), and Microlepidoptera (micro-moths) were used to classify these insects. Later, all lepidopterans were classified according to the complexity of wing venation and wing-coupling systems. Most recent classifications use four suborders based on mouthpart morphological features and female reproductive systems. Within the Glossata, Monotrysian lepidopterans have genitalia with a single opening for both copulation and oviposition, while Ditrysian lepidopterans, which include about 98% of all the Lepidoptera, have genitalia with separate openings for copulation and oviposition. The most current classification recognizes about 30 superfamilies with 122 families; all except three occur within the Glossata.

Physical characteristics

Adult lepidopterans vary widely in size and structure. The smallest species are leaf-miner moths in the families Nepticulidae (forewing 0.06 in, or 1.5 mm) and Heliozelidae (forewing 0.07 in, or 1.7 mm); the largest known species is Thysania agrippina (Noctuidae) from the American tropics, with a wingspan of up to 11.2 in (280 mm). The smallest known butterfly species probably is Micropsyche ariana from Afghanistan or the Western pygmy blue of the United States. Both have a forewing length of 0.20–0.28 in (5–7 mm). The largest is Queen Alexandra's birdwing of New Guinea, with females attaining a forewing length of up to 5.16 in (129 mm).

Adults possess a coiled tongue or proboscis (absent in primitive micro-lepidopterans, which retain mandibles) derived from the galeae of the maxillary palps; moniliform, pectinate, or clubbed antennae; large compound eyes; two lateral ocelli; a pair of erect scaled labial palps. They lack cerci. Wings usually are large compared with the small, elongate bodies and frequently are densely covered with overlapping scales that assume a wide variety of patterns and combination of colors. Wing venation varies. In small micro-moths, it can be reduced to a few veins. In many of the larger macro-moths and butterflies, however, venation consists of an oblong discal cell formed by reduction of the main basal veins (medius and part of the radius) and a series of mostly unbranched veins radiating from the discal cell. Wing venation is similar in both wings, but the hind wing usually is smaller. The leading margin of the forewing often is crowded with veins providing strength to the leading edge.

Wing coupling is made possible by an extension of a jugal lobe (in primitive ghost moths) on the rear of the forewing, which overlaps and couples with the hind wing base during wing flexing. The most common type of wing coupling involves a small, strong cluster of hairs on the hind-wing base, called the frenulum, which is retained by a retinaculum at the base of the forewing. This mechanism allows for the hind wing to be extended in unison with the forewing in the course of flexing. A loss of the frenulum/retinaculum mechanism occurs in butterflies, skippers (except for one species), and certain genera of moths, resulting in an amplexiform wing-coupling device, where the strong anterior basal lobe of the hind wing overlies the base of the forewing.

Larvae generally are fleshy, soft, elongate animals with a chitinized semicircular head capsule. Antennae are small and inconspicuous. Mouthparts comprise a set of opposable mandibles. A small, erect, silk-producing organ, the spinneret, is present on the labium. The five-segmented legs on the thorax are small and end in a simple tooth, but they are lost in some families. The ventral part of the abdomen possesses differentiating number of fleshy prolegs on segments three through six and the tenth segment. The fleshy tips often bear various series of retractable hooks, called crochets, which allow larvae to grab on to a substrate. The surface of the body possesses setae, the placement of which is important in classifying various families. The body surface can be smooth or possess clusters of hairs, fleshy tubercles, or urticating hairs. Some small micro-moths are specialized leaf miners that have secondarily lost their legs and have a forward-projecting head with rasping mouthparts. Pupae generally are elongate with appendages fastened to the body. Many are encased in a silk cocoon, others are attached to a silk substrate by a small cluster of hooks called cremaster, and still others are accompanied by a single strong silk girdle. Some moth pupae are naked and occur in the ground.

Distribution

Lepidopterans occur on all land masses except Antarctica. The most northerly species may be the lymantriid moth, Gynaephora groenlandica, which has been taken on Ward Hunt Island (83°5′ north latitude) in the Canadian Arctic. Lepidopterans are most diverse in the humid tropical zones.

Habitat

Eggs, larvae, and pupae occur in nearly all terrestrial habitats, where they often are found on or near their food plants. Larvae of a few species of moths are associated with aquatic plants in freshwater streams and ponds, and a few others (some species of Lycaenidae) live in ant nests. Adults visit nectar sources, and some species are attracted to carrion, oozing tree sap, or excrement. Adults often rest on foliage, tree trunks or any other substrate. Some species aggregate on shrubs, trees, or cave entrances.

Behavior

Butterflies and moths require a certain body temperature (usually between 77 and 79°F, or 25–26°C) to be able to fly, and they regulate internal temperature according to the environmental temperature. Butterflies of temperate areas increase heat absorption by spreading the wings, angling the exposed surface, and making direct contact with the substrate (dorsal basking) or by folding the wings above the body so that they are perpendicular to the sun's rays (lateral basking). In the tropics, butterflies fly early in the morning and at dusk, seeking refuge in the shade when the temperature climbs too high. Most moths regulate their thoracic temperature through actively preheating the thorax by vibrating their wings and simultaneously contracting antagonistic flight muscles. To diminish heat dissipation, sphingids and other large moths have insulating scales and hairs on the thorax and air sacs and diaphragms that separate the thorax from the abdomen so that the abdomen remains cooler.

Most adult butterflies and moths are solitary, but there are cases of gregarious behavior. Certain migratory species crowd together in wintering quarters, others form nocturnal roosts, and still others cluster on damp ground. Lepidopterans display a wide range of types of defensive behavior. Several larvae build protective cases, where they spend all or part of their time. For example, the bagworms of the family Psychidae form a case of silk covered with twigs, leaf fragments, and sand. Other larvae, feeding in exposed situations, show cryptic concealment, blending in by means of their green or brownish color with the leaves or bark where they feed or imitating part of a plant. When discovered, various species display startle or escape responses, such as exposing brightly colored and sharp, acrid osmeteria; regurgitating a brightly colored liquid; or faking sudden death by dropping to the ground.

Several aposematic larvae (larvae with warning coloration) in the families Lymantriidae, Limacodidae, Anthelidae, and others accompany their bright colors with urticating setae, or spines, which are physically or chemically irritating or cause allergic reactions, making the larvae inedible to predators. Certain larvae even mimic snakes. The same defensive behaviors occur in adults; for example, several species pretend to be dead when they are handled, some arctiids discharge a foul-smelling yellow fluid from specialized thoracic glands, and pyralid and noctuid moths emit ultrasonic sounds to warn bats about their bad taste. Several diurnal lepidopterans exhibit aposematic coloration, announcing their distastefulness or mimicking truly inedible species. This is the case with several species that mimic wasps and bees with their orange-and-black or yellow-and-black abdomens and translucent wings and a few species complexes in which there is Batesian mimetism (e.g., monarch butterflies) and Müllerian mimetism (clusters of similarly marked distasteful species, for example monarchs and viceroys).

Most butterflies and moths disperse, but only about 200 species regularly migrate long distances, returning to the areas where they breed. Migratory butterflies are found among the pierids, nymphalids, lycaenids, and hesperids and moths in the sphingids and noctuids (night flight) and uranids (day flight). In northern parts of North America and Europe, many species display seasonal southward movements: certain butterflies fly south in late summer or autumn and then north from Central or South America or South Africa in spring. A particular individual of a migrating species is not able to undertake a complete round trip; the return trip is accomplished by the offspring. The best-known example of a migrating butterfly is the monarch butterfly, Danaus plexippus, distributed from southern Canada to Paraguay in the New World. In North America three to five generations feed on milkweeds in the summer. In the fall they migrate south to overwintering places in California, Mexico, and Florida, where adults congregate in great numbers on certain kinds of trees. In the spring they begin their journey back north, laying eggs along the way before dying. The subsequent generation completes the return flight the following fall.

Feeding ecology and diet

Lepidopterans are predominantly phytophagous (plantfeeders), and 99% of them exploit higher plants (angiosperms); larvae feed on plant tissues, while adults feed mostly on nectar. The vast majority of these insects specialize in one (monophagy) or a few (oligophagy) food plants; only a small percentage feed on a wide range of food plants (polyphagy). Mouthparts of most adult butterflies and moths are designed for sucking, and almost all of them feed on nectar. Some lepidopterans complement their nectar-based diet with pollen, and others are saprophagous, sucking fluids on rotting fruits, carrion, dung, or droppings. Among the noctuids, for example, there are a few that feed on fruits that have been opened by other animals and others that have a strong proboscis and are able to pierce fruit skin. In the tropics a few species are able to suck the lachrymal fluids of cattle and other mammals, including humans, while others prefer urine, excreta, and cutaneous secretions. Calpe eustrigiata, an oriental noctuid moth, has developed hematophagous habits, piercing the skin of mammals to feed on their blood. Lepidopterans also imbibe water and salts from the substrate.

Herbivorous larvae usually specialize in a certain tissue of the plant: leaves, flowers, seeds, buds, or wood. There are several endophytic species, the larvae of which spend their life digging galleries inside plant tissues. Most lepidopteran larvae are exophytic, however, feeding on the outside of plants. Some larvae have unusual diets. Micropterigid larvae feed on mosses, whereas adults, which have chewing mouthparts, feed on pollen. Certain larvae feed on lichens (some arctiids, noctuids, and lycaenids), fungi (some tineids), algae and diatoms on the surface of submerged rocks (aquatic pyralids), or ferns. There are some carnivorous caterpillars that feed on other insects. For example, certain Hawaiian geometrids prey on Diptera, and Asian noctuids eat scale insects. A few lycaenids feed on aphids, others on homopterans, and still others on ant eggs, larvae, and pupae. Some noctuid larvae live in the pitcher of the carnivorous plant Nephentes, where they eat the plant's prey, and certain tineids and gelechiids steal prey from spider webs. Several microlepidopterans eat plant and animal derivatives, such as cloth, wool, fur, feathers, grains, dried fungi, paper, rotting wood, and droppings of birds and mammals.

Reproductive biology

Most species reproduce bisexually, but some European species of psychids are parthenogenetic. Two mate-location strategies are known among butterflies: territoriality and patrolling. Territorial males perch on a post, from which they defend an area and attack and pursue intruders. Patrolling males fly through proper habitats in search of receptive females. Butterflies rely on visual stimuli to locate their mates and use pheromones produced by the male only secondarily, whereas moths generally locate their mates by pheromones liberated by the female to attract the male. Among butterflies, when the male spots a female of a specific shape or color, he pursues her. The female then drops to the ground, and the male may perform a courtship ritual, moving his antennae and wings around her. He also may liberate an aphrodisiac pheromone from the glands on his wings, legs, thorax, or abdomen (modified scales or brushlike tufts of hairs called androconia), which acts over a short distance and a short period of time. If the female is receptive, mating takes place. In moths the female produces pheromones from the abdominal glands (usually located between segments eight and nine) that act over long distances to attract males. Courtship typically is simple, and mating takes place shortly after the male reaches the female. Male receptors for the pheromone are in his bipectinate antennae and allow him to detect a single molecule of the pheromone.

Females insert the eggs into plant tissues with an ovipositor or wedge-shaped papillae anales, glue them to a substrate, or simply drop them during flight. In some Monopis species (tineids) the eggs are retained in the enlarged vagina until they are ready to hatch, and the larvae emerge immediately after eggs are laid. Eggs can be laid singly or in batches; the number of eggs and their size depend on the size of the adult and the degree of alimentary specialization of the larva. Species with specialized monophagus diets produce a larger number of eggs of smaller size, which typically are dropped in flight; species with generalist polyphagus diets, on the other hand, tend to choose oviposition sites carefully. Eggs may pass through a diapause period in some species, during which they remain in a latent state; in temperate regions diapause may allow eggs to survive the winter or, in tropical regions, drought periods.

The duration of the larval stage varies with the species, its feeding ecology, the temperature, and the availability of food; it can range from 15 to 30 days to two years. When mature, the larva searches for a suitable place to pupate: in soil or litter, beneath a stone or bark (i.e., sphingids, notodontids, noctuids, and saturnids), or by rolling up a leaf and sewing the margins together with silk (i.e., tortricids and pyralids). Some larvae spin silk to attach themselves head downward to a support or to construct a protective web (arctiids), silken tube shelters (some pyralids and tineids), or cocoons (silkworm and gypsy moths). Pupae or chrysalids suspend themselves through a thread of silk from the cremaster. Emergence from cocoons takes place through caplike flaps with a secretion of fluid that dissolves the cocoon wall, or the insects may cut or force their way through the wall with sharp structures on the head.

In temperate areas most species spend short and low-temperature winter months as eggs, as larvae in diapause, or as pupae and summer months as adults. In the tropics, where seasonal temperature differences are small, some species are present as adults all year. Seasonal variations for species occur in deciduous forests and savannas but are related to rainy and dry seasons. The average life span of an adult is only a few days or weeks, although some danaines and heliconines may live for as long as six months. The timing of flight is correlated with the life cycle of the food plant and whether the species overwinter. Species that overwinter as adults, such as the mourning cloak (Nymphalis antiopa) and several Polygonia anglewings in North America, fly in early February and March. Species that overwinter as pupae fly next, followed by species that spend the winter as larvae and, finally, those that do so as eggs.

Conservation status

More than any other order of insects, lepidopterans have attracted the attention of conservationists. In all, 284 lepidopterans of 747 total insects are listed on the Red List of the World Conservation Union (IUCN); 25 are on the U.S. Fish and Wildlife Service list of endangered species; and several others, including the giant birdwing butterflies of the Indo-Australian region, are listed by the Convention on International Trade in Endangered Species (CITES). Butterflies, in particular, have been accorded conservation status because they are easily seen, colorful, day-flying insects and favorites with collectors. Collecting and commercial trafficking of butterflies often have been considered to pose a principal threat to their existence, but, as with many other insects, it is habitat loss that represents the primary threat. Although some subspecies of butterflies (e.g., Speyeria adiaste atossa from southern California and Cercyonis sthenele sthenele and Glaucopsyche lygdamus xerces from the San Francisco Bay Area) have become extinct in recent times, very little is known about the status of moths, because so few people study them. Many endemic species of micro-moths (e.g., Hyposmocoma species) from the Hawaiian Islands, for example, are known only from the original specimens collected about a century ago, and it has been postulated that many may be extinct owing to habitat loss and introduction of lepidopteran parasitoids.

Positive steps have been taken to preserve some species considered aesthetically beautiful or rare. One of the best success stories is that of the magnificently colorful birdwing butterflies (Ornithoptera and Troides) of Papua New Guinea. They represent a prime example of sustainable wildlife management by encouraging butterfly farming, breeding highly desirable species. Survival of the largest and most restricted butterfly in the world, Ornithoptera alexandrae, may depend on such a program. Natives have ranched this species and others. The practice advocates maintenance of habitat by encouraging growth of the butterfly's food plant. Some of the birdwings are offered for sale, and others are released into the environment. Butterfly-breeding farms also have cropped up in many global regions, and they annually ship thousands of living pupae to butterfly zoos throughout the world.

Significance to humans

Butterflies have always been a favorite insect motif in art; they are represented in Egyptian temples, Chinese amulets, Aztec ceramics, and an endless number of paintings, sculptures, gem carvings, textiles, glass, drawings, and poetry, symbolizing joy or sorrow and eternal life or the transience of life. In some cultures, butterflies and moths have a symbolic connection to the soul: the word for butterflies and moths in Russian means "little soul," and in Greek it means simply "soul." Some European traditions maintain that witches and fairies turn themselves into moths or butterflies to go inside houses. In the pre-Columbian cultures of Central America they were respected in religious and mythical traditions, representing souls of the dead, new plant growth, the heat of fire, sunlight, and various transformations of nature.

Many butterflies and moths fly from one flower to another and are major factors in pollination, and hence reproduction, of angiosperms. Some species are directly beneficial in that they have predatory larvae that feed on aphids and scales (e.g., several pyralids, lycaenids, noctuids, and blastobasids) or are parasites of plant-sucking leafhoppers (epipyropids). To this day, the silkworm is used in the production of textiles; the eggs of a gelechiid moth are employed commercially to raise the parasitoid wasp Trichogramma, which is released to control noctuid moths; and the larvae of a pyralid from South America are used in South Africa and Australia to keep in check invasive Opuntia cacti. In tropical areas of South America, New Guinea, Australia, Madagascar, and Africa, caterpillars are a complement of the human diet.

Because of their phytophagous nature, many caterpillars are major pests. Examples include leaf-rollers (Tortricidae), leaf-tires and webworms (Pyralidae), leaf miners (Incurvariidae and Gracillariidae), cutworms and armyworms (Noctuidae), underground grass grubs (Hepialidae), borers (Hepialidae, Cossidae, and Sesiidae), forest defoliators (Limacodidae and Geometriidae), stored fibers and foods pests (Tineidae, Gelechiidae, and Pyralidae), and crop pests (Tortricidae, Plutellidae, Gelechiidae, Noctuidae, and Pieridae).

Species accounts

Resources

Coville Jr., Charles. A Field Guide to the Moths of Eastern North America. Boston: Houghton Mifflin, 1984.

Kristensen, N. P. Lepidoptera: Moths and Butterflies. Volume 1, Evolution, Systematics, and Biogeography. Berlin and Hawthorne, NY: Walter de Gruyter, 1999.

Leverton, Roy. Enjoying Moths. London: T. and A. D. Poyser, 2001.

Nielsen, E. S., and I. F. B. Common. "Lepidoptera (Moths and Butterflies)." In: The Insects of Australia: A Textbook for Students and Research Workers. Vol. 2. 2nd edition. Carlton, Australia: Melbourne University Press, 1991.

Parsons, Michael. The Butterflies of Papua New Guinea: Their Systematics and Biology. London: Academic Press, 1998.

Sbordoni, Valerio, and Saverio Forestiero. Butterflies of the World. Westport, CT: Firefly Books, 1998.

Stehr, Frederick W., ed. "Order Lepidoptera." In Immature Insects. Vol. 1. Dubuque, IA: Kendall/Hunt Publishing, 1987.

Tyler, H., K. S. Brown Jr., and K. Wilson. Swallowtail Butterflies of the Americas: A Study in Biological Dynamics, Ecological Diversity, Biosystematics and Conservation. Gainesville, FL: Scientific Publishers, 1994.

Van-Right, R. I., P. R. Ackery. The Biology of Butterflies. Symposium of the Royal Entomological Society of London 11. London: Academic Press, 1984.

Heppner, J. B. "Classification of Lepidoptera." Part 1, "Introduction." Holarctic Lepidoptera 5, suppl. 1 (1998): 1–148.

Association for Tropical Lepidoptera. P. O. Box 141210, Gainesville, FL 32614-1210 United States. Phone: (352) 392-5894. Fax: (352) 373-3249. E-mail: jbhatl@aol.com Web site: .

Idalia Society of Mid-American Lepidopterists. 219 West 68th Street, Kansas City, MO 64113 United States. Phone: (816) 523-2948.

Lepidoptera Research Foundation, Inc.. 9620 Heather Road, Beverley Hills, CA 90210 United States.

Lepidopterists' Society. 1900 John Street, Manhattan Beach, CA 90266-2608 United States.

Lepidopterological Society of Japan. c/o Ogata Hospital, 3-2-17 Imabashi 3, Chuo-ku, Osaka, 541 Japan. E-mail: vem 15452@niftyserve.or.jp.

Societas Europaea Lepidopterologica. c/o Zoological Institute, University of Bern, Baltzerstrasse 3, Bern, CH-3012 Switzerland.

[Article by: Natalia von Ellenrieder, PhD; Rosser W. Garrison, PhD]

The order of scaly-winged insects, including the butterflies, skippers, and moths. This is one of the largest orders in the class Insecta. It has more than 100,000 species, about 10,000 in North America, and between 125 and 175 families. The adults have a covering of hairs and scales on the wings, legs, and body, and are often beautifully colored. With minor exceptions, the adults are also characterized by two pairs of membranous wings and sucking mouthparts, featuring a prominent, coiled proboscis. Butterflies and skippers usually fly in the daytime, and most moths are nocturnal. The adults usually take liquid food, such as nectar and juices of fruits. The caterpillars are almost always herbivorous.

Morphology

The most unusual feature of the head of the adult animal is the form of the mouthparts. The proboscis is extended by blood pressure created by the retraction of the stipites of the maxillae. Diagonal muscles within each proboscis unit cause the proboscis to coil. The liquid food issucked up by means of a muscular pump, formed from the pharynx, buccal cavity, and cibarium, a food pocket of the mouth cavity. Ocelli are absent in many groups. Antennae are quite variable in form.

The prothorax is well developed in some lower groups but it is considerably smaller than thepterothoracic, or wing-bearing, segments, and is largely membranous in most Lepidoptera. The most prominent feature of the dorsum of the prothorax, in most groups, is a pair of protuberant sclerotized lobes.

The scales are very variable in form. Generally, they are flat, thin, sclerotized sacks, with striated outer surfaces. The vast spectrum of colors seen in the Lepidoptera can be grouped into two categories, pigmentary and structural colors. Pigmentary colors result from pigments which are present in the scales. Structural colors are the result of either fine surface ridges on the scales or layers within the cuticle, which interfere with or diffract the light. The structural colors are generally metallic or iridescent.

In most moths, the wings are coupled by a single spine formed by fused setae which project forward from the base of the hindwing and are held by a clasp on the forewing. The spine is known as the frenulum and the clasp is the retinaculum. In the Homoneura, there is a lobe, the jugum, at the base of the forewing, which engages the hindwing, or the frenulum, when it is present.

The form of the external genitalia, especially that of the male, has been widely used in the separation of species.

Developmental stages

Metamorphosis is complete. The larvae, commonly called caterpillars, are mandibulate and cylindrical, with short thoracic legs and a variable number of abdominal prolegs. They have one pair of thoracic and eight pairs of abdominal spiracles. Pupae are variable in form and have appendages that are usually firmly cemented down (obtect), though they are sometimes partly or completely free (exarate). Pupae often are enclosed in a silken cocoon.

Classification

Unfortunately, the classification of the Lepidoptera is the subject of considerable controversy; much will doubtless be resolved when more is known of the anatomy and life history of manygroups.

A rather conservative classification has been used here. In many superfamilies only the more important families are mentioned. The table lists the important families.

*The first figure is the number of described species in North America north of Mexico. This figure is reasonably accurate. The second figure, in parentheses, is a rough estimate of the number of described species in the world. It is difficult to postulate the actual total of Lepidoptera species from these figures since in some groups, such as the Papilionidae, probably more than 90% of the existing species have been described, while in others, such as many families of microlepidoptera, the figure may be well under 25%.

Homoneura (Jugatae)

Fore- and hindwings are similar in shape and venation. They are connected by a jugum and, sometimes, also by a frenulum. Mouthparts are mandibulate, with mandibles vestigial or absent, and the galeae forms a rudimentary proboscis. The female has a single genital opening. The pupae are free or partially free.

This small suborder, including less than 1% of the species in the order, contains a diverse group of primitive forms showing certain features in common with the Trichoptera, or caddis flies.

The superfamily Micropterygoidea includes one small family, the Micropterygidae, minute moths possessing toothed, functional mandibles and lacking even the most rudimentary proboscis. The larvae, which feed on mosses, are unusual in having eight pairs of abdominal prolegs.

Superfamily Eriocranioidea is a group of tiny moths; the mandibles are greatly reduced and untoothed. Three families, the Eriocraniidae, Neopseustidae, and Mnesarchaeidea, have been recognized within the superfamily. The leaf-mining larva lacks legs. The adults reportedly do not feed. The females have a piercing ovipositor. Superfamily Hepialoidea contains medium- to large-sized moths which possess rudimentary mouth-parts. The larvae are borers. The rapid flying adults are mostly crepuscular, thus the common name swift, or ghost, moths. The only family of importance is the Hepialidae.

Heteroneura (Frenatae)

Fore- and hindwings are markedly different in shape and venation. Usually they are connected by a frenulum and retinaculum. Mouthparts are formed for sucking or, rarely, are vestigial, Adults with functional mouthparts feed on nectar of flowers, juices of rotten fruits, and other liquids. The female usually has two genital openings. Pupae are usually obtect.

One family, the Incurvariidae, comprises the superfamily Incurvarioidea. The wings are covered with microscopic spines and the females have a single genital opening. The venation is almost complete.

Superfamily Nepticuloidea includes one family, the Nepticulidae. These tiny moths have wing spines and the females have a single genital opening, but they differ from the Incurvarioidea in having a reduced venation. The larvae are generally miners in leaves, bark, and rarely, in fruits.

Superfamily Cossoidea includes one family, the Cossidae, commonly called the carpenter or goat moths. These are heavy-bodied moths, with the abdomen extending well beyond the hindwings. Mouthparts are rudimentary except for labial palpi.

Superfamily Tineoidea contains 16–39 families; the number varies with the author. This is a very large group, of uncertain composition. These moths are of small size, usually with well-developed maxillary palpi. The labial palpi have a slender, pointed third segment. Venation may be reduced, and the wings may be divided into plumes.

The small, wide-winged moths of the superfamily Tortiicoidea belong to two families, the Olethreutidae and the Tortricidae. The maxillary palpi are vestigial or absent, and the third segment of the labial palpus is short and usually obtuse. The hair fringes of the wings are always shorter than the width of the wing.

The family contains a number of agriculturally undesirable species. Paramount among these isthe codling moth (Carpocapsa pomonella L.), which is a very serious pest of apples and other fruits. The large genus Laspeyresia contains the interesting Mexican jumping bean moth (L. saltitans Westwood). The violent movements of the larvae of this moth are responsible for the action of the beans which they inhabit.

Tortricidae is a family which generally lacks the fringe of long hairs along the cubitus, characteristic of the Olethreutidae. The spruce budworm (Choristoneura fumiferana Clemens) is probably the most important injurious tortricid. In many places, especially in eastern Canada, it has defoliated vast areas of coniferous forest.

The superfamily Pyralidoidea is comprised of moths that are moderately small to medium-sized, long-legged, and slender-bodied. The maxillary palpi are usually well developed. Pyralididae is the second largest family of moths. They are small and medium-sized, and a wing expanse of 20–35 mm is not uncommon. The legs are usually long and slender. Pterophoridae is the family known as the plume moths. The wings are divided into featherlike plumes, of which there are usually two in the forewing and three in the hindwing. The moths lack maxillary palpi and have slender bodies and long legs. The larvae feed exposed or are borers.

Superfamily Zygaenoidea includes moderately small- to medium-sized moths that have complete venation, rudimentary palpi and, usually, a rudimentary proboscis. The wings are broad with short fringes. The larvae are short and more or less sluglike and are exposed feeders.

Superfamily Castnioidea includes one family, the Castniidae. They are large, diurnal, butterflylike moths with clubbed antennae, upright eggs, and boring larvae. A proboscis may be either present or absent. These moths are considered by some to be distantly related to the butterflies, but the resemblances may very well be due to convergence.

Members of the superfamily Geometroidea are small to large moths with reduced maxillary palpi and tympanal organs at the base of the abdomen. The frenulum may be present or absent.

Geometridae includes the measuring worms, loopers, and cankerworms, which make up a very large family of small- and medium-sized moths with slender bodies and relatively broad wings. The females are occasionally apterous. The larvae have the anterior prolegs reduced or absent; usually only those on segments 6 and 10 are well developed. They proceed with a characteristic looping motion, which is the basis for the scientific name. The larvae ordinarily are exposed feeders.

Sphinx, hawk, or hummingbird moths constitute the one family Sphingidae of the superfamily Sphingoidea. These medium-sized to very large, heavy-bodied moths have extremely rapid flight. The adults are mostly crepuscular or nocturnal, but a few genera are diurnal. The antennae are thickened and have a pointed apex. The proboscis is well developed and often extremely long. The wings are narrow, with the hindwing much shorter than the forewing. The larvae are external feeders and usually have a characteristic caudal horn. The pupa is in a cell in the ground or in a loose cocoon at the surface, and its long proboscis is often in a projecting case resembling a pitcher's, handle.

Moths of the superfamily Saturnioidea are medium-sized to very large moths with the frenulum almost always reduced or absent. There is no tympanum and the mouthparts are usually reduced. The antennae are ordinarily pectinate, especially in the males.

Noctuoidea is a large, rather uniform superfamily of more than 20,000 species. Most of them are moderately large moths with reduced maxillary palpi. Tympanal organs are present in the metathorax.

The superfamily Hesperioidea includes one rather large family, the Hesperiidae. The skippers are small to moderately large, heavy-bodied, mostly diurnal insects with a clubbed antenna, which is bent, curved, or reflexed at the tip. The larvae have a prominent constriction, or neck, behind the head, and often live in leaves drawn together by silk. Those of the giant skippers are borers in yucca and agave. The pupa is usually enclosed in a slight cocoon.

Butterflies of the superfamily Papilionoidea are small to large diurnal insects with clubbed antennae, which are rounded at the tip and not bent or reflexed. The forewings always have two or more veins, which are stalked.

Biological aspects

The Lepidoptera are a group of insects on which much biological research remains to be done. A great deal is still unknown about the genetics, physiology, and ecology of this group. Moreover, butterflies and moths have proved useful as experimental animals in genetical research. The larvae of many species are injurious to certain crops, causing severe economic losses. Lepidoptera of all stages are subject to the attacks of a large number of predators, including birds, mammals, lizards, frogs, spiders, and certain other insects.

The Lepidoptera penetrate almost every section of the globe, with the major exception of Antarctica. Arctic and alpine tundra areas normally support a lepidopteran fauna which, although relatively poor in species, is rich in numbers. After rains, deserts are often alive with butterflies and moths. Tropical areas are by far the richest in species. One of the strangest habitats occupied by a lepidopteran is the hair of the neotropical three-toed sloth, where the sloth moth (Bradypodicola hahneli), a pyralid, passes its entire life cycle presumably feeding on algae which grow in the hair.

Migration is the most spectacular behavior occurring in the order. It is most frequent in the butterflies Phoebis, Danaus, Libytheana, and others, but is also known in moths such as Chrysiridia. Huge migratory swarms are frequently reported in many parts of the world, but this phenomenon, as well as the related communal roosting of adults, daily use of flyways, hill-topping, and so forth, is poorly understood.

Aggregations of butterflies at a mud puddle are a frequent sight, and moths have been observed “pumping,” an act which consists of sipping water steadily from a puddle and ejecting it as a stream of drops or fine spray from the anus.

The phenomena of mimicry and protective resemblance are widespread in the Lepidoptera. Eyespot patterns in certain species elicit escape responses in passerine birds. Thus, they are an effective protective device. The appearance of dark forms of various moths in heavily industrialized, and thus heavily sooted, areas is a widespread phenomenon. This “industrial melanism” is doubtless due to shifting selection pressures and is one of the best-known examples of evolution in action. See also Insecta; Protective coloration.

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Moths, butterflies and allies Fossil range: 199–0 Ma PreЄ Є O S D C P T J K Pg N Jurassic – Recent
A Giant Leopard Moth (Hypercompe scribonia)
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Subclass:	Pterygota
Infraclass:	Neoptera
Superorder:	Endopterygota
Order:	Lepidoptera Linnaeus, 1758
Suborders
Aglossata Glossata Heterobathmiina Zeugloptera

Lepidoptera (pronounced /ˌlɛp.ɪ.ˈdɒp.tər.ə:/) is a large order of insects that includes moths and butterflies (called lepidopterans). It is one of the most speciose orders in the class Insecta, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies and found virtually everywhere. Lepidoptera contains more than 180,000 species^[1] in 128 families and 47 superfamilies. The name is derived from Ancient Greek λεπίδος (scale) and πτερόν (wing). Estimates of species suggest that the order may have more species and is among the four largest, successful orders, along with the Hymenoptera, Diptera, and the Coleoptera.^[2]

Species of the order Lepidoptera are commonly characterized as being covered in scales, having two large compound eyes, and a elongated mouthpart called a proboscis. Almost all species have membranous wings, except for a few who have crossvein wings. Thought the larva are completely different in form, having a cylindrical body with a well developed head, mandible mouthparts, and from 0–11 leg (usually 8).

The Lepidoptera have, over millions of years, evolved a wide range of wing patterns and coloration ranging from drab moths akin to the related order Trichoptera to the brightly colored and complex-patterned butterflies.^[3] Accordingly, this is the most recognized and popular of insect orders with many people involved in the observing, study, collecting, rearing and commerce of these insects. A person who collects or studies this order is referred to as a lepidopterist. Many species of the order are of economic interest by virtue of the silk they produce, or being pests, or due to the pollination they carry out.

Contents 1 Etymology 2 Distribution and diversity 3 Morphology and physiology 3.1 Head 3.2 Thorax 3.3 Abdomen 3.4 Internal physiology 4 Polymorphism 5 Reproduction and development 5.1 Mating 5.2 Life cycle 6 Behavior and ecology 6.1 Flight 6.1.1 Navigation 6.1.2 Migration 6.2 Communication 6.3 Defense and predation 7 Evolution 7.1 Phylogeny 8 Taxonomy and systematics 8.1 Distinguishing characteristics 8.2 History of study 9 Relationship to people 9.1 In culture 9.2 As pest 9.3 As beneficial 9.4 As food 10 Cited references 11 Other references 12 See also 13 External links

Etymology

The word Lepidoptera comes from the Latin word for "scaly wing", from the Ancient Greek λεπίδος (Lepidos) meaning scale and πτερόν (pteron) meaning wing. Sometimes the term Rhopalocera is used to group the species that are butterflies, from the Ancient Greek ῥόπαλον (Rhopalon) and κέρας (kæras) meaning club and horn respectively; coming from the shape of the antennae of butterflies.

The origins of the common names of many species vary. The English word butterfly is from Old English buttorfleoge, with many variations in spelling. Other than that, the origin is unknown, however it could be derived from the pale yellow color of many species' wings (eg., Yellow Sulfur: Pieridae) suggests the color of butter.^{[4][5](butterfly)} The species of Heterocera are commonly called moths. The origins of the English word moth are more clear, which comes from Old English "moððe" (cf. Northumbrian "mohðe") from Common Germanic (compare Old Norse "motti", Dutch "Mot" and German "Motte" all meaning "moth"). Perhaps its origins are related to Old English "maða" meaning "maggot" or from the root of "midge" which until the 16th century was used mostly to indicate the larva, usually in reference to devouring clothes.^[5](moth)

Distribution and diversity

Out of the more than 180,000 species described to date can be found virtually everywhere. Some 11,300 species are from North America, and 10,000 from Australia. Lepidoptera are found in a large variety of habitats, but almost always associated with higher plants, especially angiosperms (flowering plants).^[6]

Morphology and physiology

Main articles: Lepidoptera morphology and Glossary of Lepidopteran terms

Butterflies have two antennae, two compound eyes, palpi and a proboscis.

As in all insects, all species of Lepidoptera have an exoskeleton and segmented into three parts (head, thorax, abdomen) with scales and three pairs of legs. They also have two large compound eyes, and a elongated mouthpart called a proboscis. Almost all species have membranous wings, except for a few who have crossvein wings. The larva are completely different in form, having a soft body with a well developed head, mandible mouthparts, and up to 11 pairs of legs (usually 8).^[6]

Head

The adult head is segmented into six segments, though controversial, and covered with hair-like scales. Lepidoptera have a long proboscis curled under their head, and have large compound eyes which cover much of their head.^[7] Lepidoptera have long, segmented antennae that act as olfactory organs.^[8] Lepidoptera antennae also have a Johnston's organ, a collection of sensory cells found in the pedicle (or second segment) of the antennae which perceives stretching between the pedicel and the rest of the antenna.^[7] In the case of Monarch butterflies, it has been shown that antennae are necessary for proper time-compensated Sun compass orientation during migration, that antennal clocks exist in monarchs, and that they likely provide the primary timing mechanism for Sun compass orientation.^[9][10]

The larvae have a very different head then the adults, with a hard and well developed head cap for protection, and mandible mouthparts designed to feed on plants.^[6]

Thorax

Parts of an adult butterfly

The thorax contains much of the insects means of locomotion, such as the legs and the wings. The thorax consists of three invisibly divided segments, namely the prothorax, metathorax and mesothorax.^[8] Adults have two pairs of membranous wings covered, usually completely, by minute scales. In some species, wings are reduced or absent (often in the female but not the male). Antennae are prominent. In moths, males frequently have more feathery antennae than females, for detecting the female pheromones at a distance. The Trichoptera (caddisflies) which are a sister group of the Lepidoptera have scales, but also possess caudal cerci on the abdomen, a feature absent in the Lepidoptera.^[2]

Abdomen

The abdomen comprises about 9 segments, from segment 5 to 13 in larvae. The eleventh segment of the larvae holds the anal clasps which are represented as the genitalia which protrudes in the case of some taxa.^[8]

The compound eyes two large eyes made of a large number of hexagonal facets, or lenses, where each is connected to a lens-like cylinder that are attached to nerves leading to the brain.^[8] Each facet subtends and forms the part of an ommatidium, or cluster of photoreceptor cells.^[7]

Internal physiology

The digestive system of Lepidoptera consists of the proboscis, leading to the esophagus or gullet, and the stomach, over which is a large, bladder-like vessel called the proventriculus, a sort of crop preceding the true stomach, which is a cylindrical tube; the intestine is a slender tube (varying in shape in different genera) divided into the small intestine, the colon, and the rectum.^[11]

Polymorphism

Balanced polymorphism
Seasonal polymorphism
Geographic polymorphism
Sexual dimorphism
Mimicry
^[6]

Reproduction and development

Mating pair of Laothoe populi, or Poplar Hawk-moths, showing two different color variants

Species of Lepidoptera undergo Holometabolism, a form of metamorphism called complete metamorphism. Their life cycle normally consist of an egg, larva, pupa, and an imago or adult.^[6] The larvae are commonly called caterpillars, and the pupa of moths called cocoons and of butterflies called chrysalis.

Mating

Mating would begin where an adult (female or male) would attract a mate normally using visual stimuli, specially in diurnal species like most butterflies. However, most nocturnal female species (eg., moths) use pheromones instead to attracting a male, even sometimes from long distances.^[6] Some species engage in a form of acoustic courtship, or attract mates using sound or vibration such as the Polka-dot wasp moth (Syntomeida epilais).^[12]

Life cycle

The life cycle of an Anise Swallowtail

Lepidopteran almost always reproduce sexually and are oviparous, they lay eggs, though some species give to live birth called ovoviviparity. There is an great variety of difference in egg-laying and the number of eggs laid, some species simply drop their eggs in flight (these species normally have polyphagous larvae, or eat a variety of plants eg., Hepialids and some Nymphalids^[13]) though most Lepidopteran will lay their eggs near or on the host plant that larva feed on, normally attracted by its odor. The amount of eggs laid may very from only a few to thousands.^[6]

The larvae, or first stage in their life cycle after hatching, look very different from the adults and come in a variety of shapes and sizes. However, they are characterized by an elongated body with 0–11 abdominal legs (usually 8) and hooklets, called apical crochets, towards a well developed head with mandibles.^[6] The larvae eat every part of the plant, and are normally considered pest to their host plant; species have been found to lay their eggs on the fruit and other species lay the eggs even on clothing or fur (eg., Clothing Moths). A species of Geometridae from Hawaii has carnivorous larvae that grab and eat flies.^[4] Some species are carnivorous and others are even parasitic. The larvae develop rapidly with several generations in a year, however some species may take up to 3 years to develop.^[6]

After about 5 to 7 instars,^[14]:26–28 or molts, regualted by certain hormones like prothoracicotropic hormone stimulates the production of ecdysone, telling the insect to start molting. Then, the larva puparium, a sclerotized or hardened cuticle of the last larval instar, develops into the pupa. The pupa may be covered in silk and attached with many different types of debris or nothing at all depending on the species. The time it takes for pupae to emerge will vary between species. The adult will emerge from the pupa either by using abdominal hooks or a projection from the head.^[6]

Behavior and ecology

Flight

Lepidopterans fly mostly by flapping their wings. In some Lepidoptera, there is sometimes a gliding component to their flight. Flight occurs either as hovering, or as forward or backward motion.^[15]

Navigation

Timelapse of flying moths, attracted to the floodlights

Navigation is important to Lepidoptera species, specially for those that migrate. Butterflies, who have more species that migrate, have been shown to navigate using time compensated sun compasses. They can see polarized light and therefore orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is suggested to be particularly important.^[16] It is suggested that most migratory butterflies are those that belong to semi-arid areas where breeding seasons are short.^[17] The life-histories of their host plants also influence the strategies of the butterflies.^[18] Other theories include the use of landscapes. Lepidoptera may use coastal lines, mountains, but also man-made roads to orient themselves. Above sea it has been observed that the flight direction is much more accurate if the landscape on the coast is still visible.^[19]

Moths also show navigation, as seen in many studies. One study showed that many moths may use Earth's magnetic field to navigate, as a study of the stray Heart and Dart suggests.^[20] Another study, this time of the migratory behavior of the Silver Y, showed that this species, even at high altitudes, can correct its course with changing winds, and prefers flying with favourable winds, which suggests a great sense of direction.^[21][22] Aphrissa statira in Panama loses its navigational capacity when exposed to a magnetic field, suggesting it uses the Earth’s magnetic field.^[23]

Moths exhibit a tendency to circle artificial lights repeatedly. This suggests that these species 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. Celestial objects are so far away, that even after traveling 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, causing airborne moths to come plummeting downwards, and - at close range - which results in a spiral flight path that gets closer and closer to the light source.^[24]

Other explanations have been suggested, such as the idea that moths may be impaired with a visual distortion called a Mach band by Henry Hsiao in 1972. He stated that they fly towards the darkest part of the sky in pursuit of safety and are thus inclined to circle ambient objects in the Mach band region.^[25]

Migration

Main article: Lepidoptera migration

Monarch butterflies cluster in Santa Cruz. Monarch butterflies migrate to Santa Cruz to spend the winter.

Lepidopteran migration is usually seasonal, moving to escape dry seasons or other disadvantageous conditions. Most lepidopteran that migrate are butterflies, varying from short to over long distances. Some butterflies that migrate include the Mourning Cloak, Painted Lady, American Lady, Red Admiral, and the Common Buckeye.^[14]:29–30 Particularly famous migrations are those of the Monarch butterfly from Mexico to northern USA and southern Canada, a distance of about 4000 to 4800 km (2500–3000 mi). 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.^[26] Migrations have been studied in more recent times using wing tags and also using stable hydrogen isotopes.^[27][28]

Moths also undergo migrations, such as the uraniids. U. 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.^[29]

Communication

sources to use: [1]

Defense and predation

Lepidopterans are soft bodied and move slowly, therefore at risk to predators including birds, wasps, and mammals. Some caterpillars, such as the zebra swallowtail butterfly larvae, are cannibalistic and may eat other larvae of the same species. Wasps and flies may lay eggs in the caterpillar which would eventually kill it as they hatch inside its body and eat its tissues. Butterflies are more fragile and almost defenseless. Lepidopterans rely on a variety of strategies for defense and protection.^[30]

Papilio machaon showing the osmeterium, which emits unpleasant smells to ward off predators.

Some species of lepidoptera are poisonous to predators, such as the monarch butterfly and pipevine swallowtail butterfly. They obtain their toxicity from the plants they eat. The brightly colored caterpillars and adults are generally the toxic ones, giving their color a reminder to predators about their toxicity. Predators that eat poisonous lepidopterans may become sick and vomit violently, learning not to eat those types of lepidopterans. A predator who has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future, thus saving many other species as well.^[30][31]

Other caterpillars emit bad smells to ward off predators.^[30] Some caterpillars, especially members of Papilionidae, contain an osmeterium, a Y-shape protrusible gland found in the prothoracic segment of the larvae. When threatened, the caterpillar emits unpleasant smells from the organ to ward off the predators.^[32][33]

Camouflage and mimicry are also important defense strategies. Some lepidopterans blend with its surroundings, making them difficult to be spotted by predators. Caterpillars can be shades of green that matches its host plant. Others look like inedible objects, such as the Western Tiger Swallowtail larvae that look like bird droppings.^[30][34] For example, adult Sesiidae species (also known as clearwing moths) have a general appearance that is sufficiently similar to a wasp or hornet to make it likely that the moths gain a reduction in predation by Batesian mimicry.^[35]

Eyespots are a type of automimicry used by some lepidopterans. In butterflies, the spots are composed of concentric rings of scales of different colors. Studies have investigated their role in defensive behavior. The proposed role of the eyespots is to deflect attention to predators. Their resemblance to eyes provokes the predator's instinct to attack these wing patterns.^[36]

Evolution

Not much is known about ancient Lepidoptera species because so few fossils have been found. The earliest known Lepidopteran fossil, Archaeolepis mane is from the Jurassic period, about 190 million years ago. The fossil consists of a pair of wings with scales that are characteristically similar to the wing venation pattern found in Trichoptera (caddisflies). 2 other sets of Jurassic Lepidopteran fossils have been found, and 13 sets from the Cretaceous period.^[37]

Phylogeny

Apoditrysia Macrolepidoptera Rhopalocera Papilionoidea (true butterflies) Hesperiidae (skippers) Hedylidae (American moth-butterflies)   Geometroidea (geometer moths) Drepanoidea (hooktip moths) Cimeliidae (gold moths) Callidulidae (old world butterfly moths) Noctuoidea (Owlet, tiger moths)   Bombycoidea (Silk moths, hawk moths) Lasiocampoidea (lappet moths) Mimallonidae (sackbearer moths) Thyrididae (picture winged leaf moths) Hyblaeidae (teak moths) Copromorphoidea (fruitworm moths) Pyraloidea (snout moths) Immidae Whalleyana Tortricoidea (leafrollers) Zygaenoidea (burnet moths) Pterophoridae (plume moths) Alucitoidea (many-plumed moths) Epermeniidae (fringe-tufted moths) Schreckensteinia (bristle legged moths) Choreutidae (metalmark moths) Urodidae (false-burnet moths)   Sesioidea (clearwing moths) Cossoidea (carpenter moths) Gelechioidea (twirler moths) Yponomeutoidea (ermine moths) Gracillarioidea (leafminers) Tineoidea (bagworm moths) A proposed phylogeny of the principal lepidopteran groups.[38]

Lepidoptera and Trichoptera (caddisflies) share many similarities that are lacking in other insect orders.

• 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.
• Adults in both orders display a particular wing venation pattern on their forewings.
• The larvae of both orders have mouth structures and gland with which they make and manipulate silk.^[37]

Willi Hennig grouped the two sister orders into the Amphiesmenoptera superorder. This group probably evolved in the Jurassic, having split from the now extinct order Necrotaulidae.^[37]

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 order, but instead have chewing mandibles adapted for a special diet. Micropterigidae larvae feed on leaves, fungi, or liverworts (much like the Trichoptera).^[7] 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.^[7]

The Eriocraniidae have a short coiled proboscis in the adult stage, and though they retain their pupal mandibles with which they escaped the cocoon, their mandibles are non-functional thereafter.^[7] Most of these non-ditrysian families, are primarily leaf miners in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.^[37]

With the evolution of the Ditrysia 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 one cloaca, or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, Gracillarioidea and Gelechioidea are mostly leaf miners, but more recent lineages feed externally. In the Tineoidea, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.^[37]

The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.^[37] They evolved about the time that flowering plants underwent an expansive adaptive radiation in the mid-Cretaceous, and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved co-evolution or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.

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.^[37] Many also have tympanal organs, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.^[37] The main lineages in the macrolepidoptera are the Noctuoidea, Bombycoidea, Lasiocampidae, Mimallonoidea, Geometroidea and Rhopalocera. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a monophyletic group.^[37] The Rhopalocera, comprising the Papilionoidea (Butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved.^[7] There is quite a good fossil record for this group, with the oldest skipper about 56 million years old.^[37]

Taxonomy and systematics

Main articles: Taxonomy of Lepidoptera and Lepidopteran diversity

Distinguishing characteristics

The characteristics which distinguish the order Lepidoptera from other insect orders are:^[39]

• Head: The Lepidopteran head has large compound eyes and mouth parts which are almost always a proboscis.
• Scales: Scales cover the external surface of the body and appendages.
• Thorax: The prothorax in the case of most species is reduced.
• Wings: Two pairs of wings present in almost the taxa. The wings have very few cross-veins.
• Abdomen: The posterior abdominal segments are modified extensively for reproduction. Cerci are absent.
• Larva: The larvae are eruciform with well developed head and mandibles. They have 0 to 10 prolegs, usually 8.
• Pupa: The pupae in most species are adecticous and obtect, while they are decticous in others.

History of study

Linnaeus in Systema Naturae (1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx, and Phalaena with seven subgroups in Phalaena.^[7] These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Denis & Ignaz Schiffermüller (1775), Fabricius (1775) and Pierre André Latreille (1796). Jacob Hübner described many genera, and the Lepidopteran genera were catalogued by Ochsenheimer and Treitschke in a series of volumes on the Lepidopteran fauna of Europe published between 1807 and 1835.^[7] G.A.W. Herrich-Schaffer (several volumes, 1843-1856), 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 Tineinae13 volumes 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.^[37] He published a study of the Florissant deposits of Colorado. Andreas V. Martynov (1879-1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.^[37] Lepidoptera tend not to be as common as some other insects in the habitats that are most conducive to fossilization, such as lakes and ponds, and their juvenile stage has only the head capsule as a hard part that might be preserved. 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.^[37] The earliest fossil is Archaeolepis mane from the Jurassic, about 190 million years ago in Dorset, UK.^[37] It consists of wings and shows scales with parallel grooves under a scanning electron microscope and the characteristic wing venation pattern shared with Trichoptera.^[37] Only 2 more sets of Jurassic Lepidopteran fossils have been found, and 13 sets in the Cretaceous.^[37] From there, many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber.

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.^[7] Willi Hennig (1913-1976) developed the cladistic methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among monotrysian families and Kristensen worked more generally on insect phylogeny and higher Lepidoptera too.^[7][37]. While it is often found that DNA-based phylogenies differ from those based on morphology, this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.^[37]

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: Microlepidotera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.^[7]

Relationship to people

In culture

Death's-head Hawkmoth (Acherontia lachesis) an old bleached specimen still showing the classical skull-shaped head

Artistic depictions of butterflies have been used in many cultures including as early as 3500 years ago, in Egyptian hieroglyphs.^[40] Today, butterflies are widely used in various objects in art and jewelry: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and in some mixed media artworks and furnishings.^[41] Butterflies have also inspired the "butterfly fairy" as an art and fictional character, including in the Barbie Mariposa film.

In many cultures the soul of a dead person is associated with the butterfly. As in Ancient Greece, where the word for butterfly ψυχή (psyche) also means soul and breathe. In Latin, as in Ancient Greece, the word for butterfly papillio was associated with the soul of the dead.^[42]

The Death's-head Hawkmoth's skull pattern on it's thorax has helped these moths, particularly A. atropos, earn a negative reputation, such as associations with the supernatural and evil. The moth has been prominently featured in art and movies such as Un Chien Andalou (by Buñuel and Dalí) and The Silence of the Lambs, and in the artwork of the Japanese metal band Sigh's Hail Horror Hail album.

As pest

A corn earworm eating a ear of corn.

Species of Lepidoptera are major pest in agriculture, particularly the larvae. Many species cause damage, some of the major include Tortricidae, Noctuidae, and Pyralidae; some particular species of concern also include those of the Noctuidae genus Spodoptera (armyworms) and Helicoverpa (corn earworm).^[7] Helicoverpa zea larvae are polyphagous, meaning they eat a verity of crops. The larvae are commonly called mattering on what they are eating, for example if they are eating corn they are called corn earworm. Other common crops they consume are tomatoes, then called tomato fruitworm, and cotton, then called a cotton bollworm.^[43]

As beneficial

Most species of Lepidoptera engage in the pollination of flowers. The adults feed on the nectar inside flowers, using their proboscis to reach the nectar hidden at the base of the petals. In the process, the adult brushes against the flower's stamen, on which the flower's reproductive pollen is made and stored. The pollen is transferred to the adult, who flies to the next flower to feed and unwittingly deposits the pollen on the stigma of the next flower, where the pollen germinates and fertilizes the seeds.

As food

The larvae of Lepidoptera are eaten by indigenous people from every continent. The Mopane worm (Gonimbrasia belina) is collected and eaten by indigenous people in South Africa, Botswana, and Zimbabwe.^[44]

Cited references

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20. ^ Baker, R. Robin (February 1987). "Integrated use of moon and magnetic compasses by the heart-and-dart moth, Agrotis exclamationis". Animal Behaviour 35 (1): 94–101. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6W9W-4JT84SP-F&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&_docanchor=&view=c&_searchStrId=1129652847&_rerunOrigin=scholar.google&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=2892ea838fdde22df54095b4505c1c2b. 
21. ^ Breen, Amanda (May 7, 2008). "Scientists make compass discovery in migrating moths". University of Greenwich at Medway. pp. 1. http://www.gre.ac.uk/pr/articles/2008news/a1537---moths. Retrieved 9 December 2009. 
22. ^ Chapman, Jason W.; Don R. Reynolds, Henrik Mouritsen, Jane K. Hill, Joe R. Riley, Duncan Sivell, Alan D. Smith, and Ian P. Woiwod (8 April 2008). "Wind selection and drift compensation optimize migratory pathways in a high-flying moth". Current Biology 18 (7): 514–518. http://www.sciencedirect.com/science?_ob=MImg&_imagekey=B6VRT-4S6GG6K-7-7&_cdi=6243&_user=10&_coverDate=04%2F08%2F2008&_sk=%23TOC%236243%232008%23999819992%23684615%23FLA%23display%23Volume_18,_Issue_7,_Pages_471-552_(8_April_2008)%23tagged%23Volume%23first%3D18%23Issue%23first%3D7%23date%23(8_April_2008)%23&view=c&_gw=y&wchp=dGLbVlz-zSkWb&md5=dfc8a24f69a6b84eef8f82cf4394d837&ie=/sdarticle.pdf. 
23. ^ Srygley, Robert B.; Evandro G. Oliveira, Andre J . Riveros (2005). "Experimental evidence for a magnetic sense in Neotropical migrating butterflies (Lepidoptera: Pieridae)". The British Journal of Animal Behaviour 71 (1): 183–191. ISSN 0003-3472. http://users.ox.ac.uk/~zool0206/AnimBeh06.pdf. 
24. ^ Elliot, Debbie; Dr. May Berenbaum (August 18, 2007). "Why are Moths Attracted to Flame? (audio)". National Public Radio. pp. 1. http://www.npr.org/templates/story/story.php?storyId=12903572. Retrieved 12 December 2009. 
25. ^ Hsiao, Henry S. (1972). Attraction of moths to light and to infrared radiation. San Francisco Press. ISBN 0911302212. 
26. ^ Williams, C. B. 1927 A study of butterfly migration in south India and Ceylon, based largely on records by Messrs. G Evershed, E.E.Green, J.C.F. Fryer and W. Ormiston. Trans. Ent. Soc. London 75:1-33
27. ^ Urquhart, F. A. & N. R. Urquhart. 1977. Overwintering areas and migratory routes of the Monarch butterfly (Danaus p. plexippus, Lepidoptera: Danaidae) in North America, with special reference to the western population. Can. Ent. 109: 1583-1589
28. ^ Wassenaar L.I., Hobson K.A. 1998. Natal origins of migratory monarch butterflies at wintering colonies in Mexico: new isotopic evidence. Proc Natl Acad Sci U S A. 95(26):15436-9. Full text
29. ^ Smith, N. G.; Janzen, D.H. (editor) (1983b). Urania fulgens (Calipato Verde, Green Urania).. Costa Rican Natural History. Chicago: University of Chicago Press. pp. 816. 
30. ^ ^{a b c d} "Caterpillar and Butterfly Defense Mechanisms". EnchantedLearning.com. http://www.enchantedlearning.com/subjects/butterfly/allabout/Defense.shtml. Retrieved 7 December 2009. 
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Other references

• Kristensen, N.P. (Ed.). 1999. Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography. Handbuch der Zoologie. Eine Naturgeschichte der Stämme des Tierreiches / Handbook of Zoology. A Natural History of the phyla of the Animal Kingdom. Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York.
• Nye, I.W.Bb & Fletcher,D.S. 1991. Generic Names of Moths of the World. Volume 6: xxix + 368 pp. Trustees of the British Museum (Natural History), London.
• O'Toole, Christopher. 2002. Firefly Encyclopedia of Insects and Spiders. ISBN 1-55297-612-2.
• Nemos, F. Europas bekannteste Schmetterlinge. Beschreibung der wichtigsten Arten und Anleitung zur Kenntnis und zum Sammeln der Schmetterlinge und Raupen. Oestergaard Verlag, Berlin, ca. 1895, http://epic.awi.de/Publications/Dem1895a.pdf (pdf, 77 MB).

See also

• McGuire Center for Lepidoptera and Biodiversity, University of Florida
• Difference between a butterfly and a moth
• Moth
• List of moths
• Butterfly
• List of butterflies in Taiwan
• List of butterflies of Great Britain
• List of butterflies of Tobago
• List of butterflies of Menorca
• List of butterflies of India
• List of butterflies of North America

External links

Wikispecies has information related to: Lepidoptera

The Wikibook Dichotomous Key has a page on the topic of Lepidoptera

Wikimedia Commons has media related to: Lepidoptera

• European Butterflies and Moths by Christopher Jonko
• Norwegian Butterflies and Moths (Huge picture archive)
• Moths and Butterflies of Europe and North Africa
• British Butterflies and Moths
• Butterflies of Bulgaria
• Photography of European Butterflies and Moths
• Butterflies and Moths in the Netherlands
• Swedish Moths and Butterflies - Lepidoptera.se
• Butterflies of Asturias - Spain
• Lepidoptera of French Antilles
• Butterflies of Asian Russia
• Butterflies from Indo_China
• Butterflies of Turkey
• Moths of Jamaica
• Historic Moth illustrations
• Lepidoptera (TSN 117232). Integrated Taxonomic Information System.
• Caught Between the Pages: Treasures from the Franclemont Collection Online virtual exhibit featuring a selection of historic entomological writings and images from the Comstock Library of Entomology at Cornell University
• Japmoth Japanese moths. Access images via the numbers on the left.

v • d • e Extant Lepidoptera families by suborder Kingdom: Animalia · Phylum: Arthropoda · Class: Insecta · Subclass: Pterygota · Infraclass: Neoptera · Superorder: Endopterygota Aglossata Agathiphagidae Glossata Heteroneura Division Ditrysia: Acrolepiidae · Acrolophidae · Agonoxenidae · Aididae · Alucitidae · Anomoeotidae · Anthelidae · Arctiidae · Arrhenophanidae · Batrachedridae · Bedelliidae · Blastobasidae · Bombycidae · Brachodidae · Brahmaeidae · Bucculatricidae · Callidulidae · Carposinidae · Carthaeidae · Castniidae · Chimabachidae · Choreutidae · Cimeliidae · Coleophoridae · Copromorphidae · Cosmopterigidae · Cossidae · Crambidae · Cyclotornidae · Dalceridae · Doidae · Douglasiidae · Drepanidae · Dudgeoneidae · Elachistidae · Endromidae · Epermeniidae · Epicopeiidae · Epipyropidae · Eriocottidae · Ethmiidae · Eupterotidae · Galacticidae · Gelechiidae · Geometridae · Glyphipterigidae · Gracillariidae · Hedylidae · Heliodinidae · Hesperiidae · Heterogynidae · Himantopteridae · Holcopogonidae · Hyblaeidae · Immidae · Lacturidae · Lasiocampidae · Lecithoceridae · Lemoniidae · Limacodidae · Lycaenidae · Lymantriidae · Lyonetiidae · Lypusidae · Megalopygidae · Metachandidae · Mimallonidae · Mirinidae · Momphidae · Noctuidae · Nolidae · Notodontidae · Nymphalidae · Oecophoridae · Oenosandridae · Pantheidae · Papilionidae · Pieridae · Plutellidae · Psychidae · Pterolonchidae · Pterophoridae · Pyralidae · Riodinidae · Roeslerstammiidae · Saturniidae · Schreckensteiniidae · Scythrididae · Sematuridae · Sesiidae · Simaethistidae · Somabrachyidae · Sphingidae · Symmocidae · Thyrididae · Tineidae · Tineodidae · Tortricidae · Uraniidae · Urodidae · Whalleyanidae · Yponomeutidae · Ypsolophidae · Zygaenidae Division Monotrysia: Adelidae · Cecidosidae · Crinopterygidae · Heliozelidae · Incurvariidae · Nepticulidae · Opostegidae · Palaephatidae · Prodoxidae · Tischeriidae Dacnonypha Acanthopteroctetidae · Eriocraniidae Lophocoronina Lophocoronidae Exoporia Anomosetidae · Hepialidae · Mnesarchaeidae · Neotheoridae · Palaeosetidae · Prototheoridae Neopseustina Neopseustidae Heterobathmiina Heterobathmiidae Zeugloptera Micropterigidae

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