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.

Moths, butterflies and allies Fossil range: 199–0 Ma PreЄ Є O S D C P T J K Pg N Jurassic - Recent
The Clipper, Parthenos sylvia (Papilionoidea: Nymphalidae: Limenitidinae)
Scientific classification
Kingdom: Animalia Phylum: Arthropoda Class: Insecta Subclass: Pterygota Infraclass: Neoptera Superorder: Endopterygota Order: Lepidoptera Linnaeus, 1758
Subdivisions
Suborder Aglossata Suborder Glossata Suborder Heterobathmiina Suborder Zeugloptera See Taxonomy of Lepidoptera and Lepidopteran diversity.

Lepidoptera is an order of insects that includes moths and butterflies. 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. Members of the order are referred to as lepidopterans. A person who collects or studies this order is referred to as a lepidopterist. This order has 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]

Contents 1 General characteristics 2 Families 2.1 History of study 2.2 Phylogeny 3 Cited references 4 Other references 5 See also 6 External links

General characteristics

Main article: Glossary of Lepidopteran terms

A labeled Papilio machaon: A- Forewing B - Antenna C - Compound eye D - Proboscis E - Thorax F - Leg G- Abdomen H - Hindwing I - «tail»

A Painted Arachnis (Arachnis picta), showing scales and hair

Lepidopterans like all holometabola, undergo complete metamorphosis, going through a four-stage life cycle: egg, larva/caterpillar, pupa/chrysalis, and imago/adult.^[2] Their lifecycle can include a inactive periods or diapauses in any of the pre-adult stages that help them overcome unsuitable environmental conditions.^[2] A species of Lymantrid moth, Gynaephora groenlandica, found in the Arctic has an exceptional life cycle that was thought to take 14 years^[3] but now estimated to take 7 years.^[4]

The larvae, caterpillars, have a toughened (sclerotized) head capsule, chewing mouthparts, and a soft body, that may have hair-like or other projections, 3 pairs of true legs, and additional prolegs (up to 5 pairs). They can be confused with the larvae of sawflies. Lepidopteran larvae can be differentiated by the presence of crochets on the prolegs which are absent in the Symphyta (sawflies). Most caterpillars are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores.^[5]

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]

Adult mouth parts prominently include the proboscis formed from maxillary galeae and are adapted for sucking nectar. Some species have reduced mouth parts (some species do not feed as adults), and others have them modified to pierce and suck blood or fruit juices (some Noctuids).^[6] Mandibles are absent in all except the Zeugaloptera which have chewing mouthparts.^[7] The maxillary palpi are reduced and consist of up to five segments. They are conspicuous in some of the more primitive families and are often folded. The labial palpi are more prominent and upward pointed.^[2](See also: difference between a butterfly and a moth)

The three thoracic segments are fused and consist of non-movable sclerites. The wings arise from the meso- and meta-thoracic segments and are similar in size in the primitive groups. In the more recent groups, the meso-thoracic wings are larger with more powerful musculature at their bases and more rigid vein structures on the costal edge. In the Noctuoidea, the metathorax is modified with a pair of tympanal organs. There are a variety of wing coupling mechanisms that connect the forewings and the hindwings. The more primitive groups have an enlarged lobe, jugum, at the base of the forewing that folds under the hindwing in flight. Other groups have a frenulum on the hindwing that hooks under a retinaculum on the forewing. In some groups such as the Psychidae, Lymantriidae, the females are flightless and have reduced wings.^[2]

The abdominal segments 7-10 or 8-10 are modified to form the external genitalia. The abdomen is connected to the thorax in the more recent families by muscles connectd to projections from the abdominal sternite 2. Paired hearing organs at the base of the abdomen occur in the Pyraloidea and Geometroidea. Males have glandular organs such as expandable hair brushes or tufts, or as thin-walled, eversible sacs (coremata), from the intersegmental membranes. The genitalia are complex and provide the basis for species discrimination in most families and also in family identification.^[2]

Primitive groups have a single genital aperture near the end of the abdomen through which both copulation and egg laying occur. This character is used to designate the Monotrysia. Hepialidae and related families have an external groove that carries sperm from the copulatory opening (gonopore) to the (ovipore) and are termed exoporian. The remaining groups have an internal duct that carry sperm and form the Ditrysia, with two distinct openings each for copulation and egg-laying.^[2]It also has antennaes.

Families

Forester Moth (Zygaenidae)

There are about 69 families in this order with variations depending on the taxonomic treatment (see the family template box at the bottom of this section).

The Lepidoptera are divided into several suborders, the largest being Glossata, the vast majority of which are Ditrysia.

Several other classifications of lepidopteran families are used in older literature. These include the Rhopalocera (club-horned) consisting of what are commonly called butterflies and the Heterocera (varied-horned) consisting of the moths. However, Rhophalocera is a natural (monophyletic) group, while Heterocera is a paraphyletic assemblage.

Another non-standard classification separates the Lepidoptera into Microlepidoptera for the smaller species (mostly moths) and Macrolepidoptera for the larger species.

History of study

Linnaeus in Systema Naturae (1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx, and Phalaena with seven subgroups in Phalaena.^[6] 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.^[6] 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.^[8] 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.^[8] 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.^[8] The earliest fossil is Archaeolepis mane from the Jurassic, about 190 million years ago in Dorset, UK.^[8] It consists of wings and shows scales with parallel grooves under a scanning electron microscope and the characteristic wing venation pattern shared with Trichoptera.^[8] Only 2 more sets of Jurassic Lepidopteran fossils have been found, and 13 sets in the Cretaceous.^[8] 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.^[6] 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.^[6][8]. 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.^[8]

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.^[6]

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.[9]

It has long been noted that the Lepidoptera and the Trichoptera (caddisflies) share many similarities that are lacking in other insect orders. Among these are:

• females, rather than males, are heterogametic (i.e. their sex chromosomes differ)
• dense setae on the wings (modified into scales in Lepidoptera)
• a particular wing venation pattern on the forewings
• larvae with mouth structures and glands to make and manipulate silk.^[8]

Thus the two sister orders are grouped into the Amphiesmenoptera. The group probably evolved in the Jurassic, diverging from the extinct Necrotaulidae.^[8] Lepidoptera differ from the Trichoptera in several features, including wing venation, form of the scales on the wings, loss of the cerci, loss of an ocellus, and changes to the legs.^[8]

The oldest, most basal lineages of Lepidoptera have as adults, not the curled tongue or proboscis characteristic of most members of the order, but chewing mandibles (Micropterigidae, Agathiphagidae and Heterobathmiidae). Micropterigidae larvae feed on decaying leaves (much like the Trichoptera), fungi, liverworts or live leaves.^[6] The adults chew pollen or spores of ferns. In the Agathiphagidae, larvae feed inside seeds of kauri pines, and in Heterobathmiidae the larvae mine leaves of Nothofagus, the southern beech. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon just before adult emergence.^[6]

The Eriocraniidae have a short coiled proboscis in the adult stage, and retain mandibles for the purpose of escaping the cocoon, but they are non-functional thereafter.^[6] They, and most of the other 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.^[8]

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.^[8]

The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.^[8] 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.^[8] 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.^[8] 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.^[8] The Rhopalocera, comprising the Papilionoidea (Butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved.^[6] There is quite a good fossil record for this group, with the oldest skipper about 56 million years old.^[8]

Cited references

1. ^ "The Lepidoptera Taxome Project Draft Proposals and Information". Centre for Ecology and Evolution, University College London. http://www.ucl.ac.uk/taxome/. Retrieved on 2007-03-05. 
2. ^ ^{a b c d e f g h} Powell, Jerry A. Lepidoptera (pp. 631-664) in Resh, V. H. & R. T. Cardé (Editors) 2003. Encyclopedia of Insects. Academic Press.
3. ^ Kevan, P. G., and O. Kukal. 1993. Corrigendum: A balanced life table for Gynaephora groenlandica (Lepidoptera: Lymantriidae), a long-lived high arctic insect, and implications for the stability of its populations. Can. J. Zool. 71:1699-1701 PDF
4. ^ W. Dean Morewood and Richard A. Ring (1998) Revision of the life history of the High Arctic moth Gynaephora groenlandica (Wocke) (Lepidoptera: Lymantriidae). Can. J. Zool. 76(7):1371–1381 doi:10.1139/cjz-76-7-1371
5. ^ Dugdale, JS, 1996. Natural history and identification of litter-feeding Lepidoptera larvae (Insecta) in beech forests, Orongorongo Valley, New Zealand, with especial reference to the diet of mice (Mus musculus). Journal of The Royal Society of New Zealand, Volume 26, Number 2, pp 251-274
6. ^ ^{a b c d e f g h i j} Scoble, MJ 1995. The Lepidoptera: form, function and diversity. Oxford, UK: The Oxford University Press; 404 p.
7. ^ Borror, D.J., Triplehorn, C.A. Johnson. ( 1989) Introduction to the Study of Insects. 6th ed. Brooks Cole.
8. ^ ^{a b c d e f g h i j k l m n o p q r} Grimaldi, D, and M S Engel, 2005. Evolution of the Insects. Cambridge University Press.
9. ^ Tree of Life Accessed January 2007

Other references

• Kristensen, NP (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, IWB & DS Fletcher, 1991. Generic Names of Moths of the World. Volume 6: xxix + 368 pp. Trustees of the British Museum (Natural History), London.
• Koçak, AO & M Kemal, 2006. Checklist of Lepidoptera of Turkey. Cent. ent. Stud., Priamus Suppl. 1: 1-196.
• Kemal, M & AO Koçak, 2007. Annotated check-list of the butterflies of the Uighur Ili (Lepidoptera). Priamus 11 (4): 61-80.
• Koçak, AO & M Kemal, 2007. Revised and annotated checklist of the Lepidoptera of Turkey. Cent. ent. Stud., Priamus Suppl. 8: 1-150, 2 Tables.
• Koçak, AO & S Seven, & M Kemal, 2007. Chapter 3: Valid species group names (species, and subspecies) recorded in the Databank of the CESA, [in] Koçak,A.Ö. & M.Kemal, Results of the International Project of the CESA on the Lepidoptera of the World-I. Cent. ent. Stud., Memoirs 3-4: 143-1116.
• Kemal,M & AO Koçak, 2008. First attempt for a list of the patronyms in the Lepidopterology based upon the Info-System of the Cesa. Cent. Ent. Stud., Priamus Suppl. 12: 1-89, fig.
• Koçak, AO & M Kemal, 2008. Synonymical and distributional list of the Lepidoptera of Turkey -I.Tortricidae. Cent. ent. Stud., Priamus Suppl. 13: 1-39.
• Koçak, AO & M Kemal, 2008. Checklist of the Diurnal Lepidoptera of Turkey. Cent. ent. Stud., Priamus Suppl. 15: 1-41.
• Koçak, AO & M Kemal, 2008. Reports on the Temporary Results of the Lepidopteran Lists of the Asian Continent based upon the Info-system of the Cesa-I. Russia. Cent. ent. Stud., Priamus Suppl. 14: 1-466.
• Koçak, AO & M Kemal, 2008. Patronymic Names in Lepidoptera I. A List of the Scientific Names Referring to the Persons based upon the Info-System of the Cesa. 212 pp. 32 figs. CD. Cesa & Yüzüncü Yıl University Publication.
• Koçak, AO & M Kemal, 2009. Report of the Project “Lepidoptera of Indo-China (LIC)” 1- Temporary index of the species group names of the Lepidoptera. Cent. ent. Stud., Priamus Suppl. 16: 1-330, 81 figs.
• Koçak, AO & M Kemal, 2009. Third Report on the Temporary Results of the Lepidopteran List of Africa Continent based upon the Info-system of the Cesa stand 1. 3. 2009. Cesa Publ. African Lepid. 25: 1-2810, figs. maps.
• Firefly Encyclopedia of Insects and Spiders, edited by Christopher O'Toole, ISBN 1-55297-612-2, 2002
• F. Nemos: Europas bekannteste Schmetterlinge. Beschreibung der wichtigsten Arten und Anleitung zur Kenntnis und zum Sammeln der Schmetterlinge und Raupen. Oestergaard Verlag, Berlin, ca. 1895, http://hdl.handle.net/10013/epic.28790.d001 (pdf, 77 MB).

See also

• McGuire Center for Lepidoptera and Biodiversity, University of Florida

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)
• Lepidoptera on GlobalTwitcher.com
• Forum Lepidoptera
• Lepidopterology.com
• Moths and Butterflies of Europe and North Africa
• Centre for Entomological Studies Ankara (CESA)
• Cesa News
• Cesa Publications on African Lepidoptera (series)
• 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
• Lepidoptera of North America
• Lepidoptera in online insect museum
• Butterflies of Asturias - Spain
• Lepidoptera of French Antilles
• Butterflies of Asian Russia
• Butterflies from Indo_China
• Butterflies of Turkey
• Moths of Jamaica
• Moths of Japan Excellent photographs.In Japanese but uses binomial names
• 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.
• Insect Life Forms Butterflies and Moths - Order Lepidoptera

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 · 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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