Lepidoptera

(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

List of Species

Silkworm

Bombyx mori

family

Bombycidae

taxonomy

Phalaena mori Linnaeus, 1758, China.

other common names

English: Silkmoth; French: Ver de la soie, bombyx du mûrier; German: Seidenspinner; Spanish: Gusano de seta; Finnish: Silkkiperhonen.

physical characteristics

Caterpillars (1.5 in, or 4 cm) are pale brown, with brown marks on the thorax and a horn on the tail. They pupate in a white to yellow cocoon, the color depending on genetics and diet. The silk forming the cocoon is a single, continuous thread (1,000–3,000 ft, or 300–900 m, long) of a protein secreted from salivary glands. Adults are heavy, rounded, furry, and whitish with pale brown lines. The forewings have a hooked tip, and the wingspan is 1.5–2.5 in (4–6 cm).

distribution

Originally from the north of China, the north of India, Japan, Taiwan, and Korea. Now also bred in Europe and North and South America as a commodity in the textile market.

habitat

On mulberry worldwide.

behavior

Adults cannot fly. Larvae are so domesticated now that they cannot survive without the assistance of humans.

feeding ecology and diet

Caterpillars feed on mulberry leaves; adults have atrophied mouthparts and do not feed.

reproductive biology

The female lays 200–500 lemon-yellow eggs that turn black and hatch in spring. In four to six weeks, larvae undergo four molts and then spin a silk cocoon (in a process taking three or more days) to pupate. Adults emerge in three weeks, reproduce, and die within five days. Univoltine (having one generation per year) under natural conditions.

conservation status

Bred in captivity for thousands of years; no wild colonies remain.

significance to humans

Used to make silk and for education and research. Originally domesticated in China. To harvest silk, cocoons are boiled in water to kill pupae and help unravel thread. Dead pupae sometimes are used as cockroach bait or fish food or to fertilize mulberry trees.

Citrus leaf miner

Phyllocnistis citrella

family

Gracillaridae

taxonomy

Phyllocnistis citrella Stainton, 1856, Calcutta, India.

other common names

German: Citrusmotte; Spanish: Minador de los cítricos, lagarta minadora dos citros; Portuguese: Lagarta minadora dos citros.

physical characteristics

Adults are small, with a wingspan of 0.16 in (4 mm) and a length of 0.08 in (2 mm) at rest. The forewing is white with silvery reflections, black and tan markings, and a small apical black spot. Full-grown larvae (0.12 in, or 3 mm) are flattened and translucent green with atrophied prolegs and legs.

distribution

Originally from India, but now associated with citrus groves worldwide.

habitat

Can occur wherever citrus is grown. Adults are seldom seen. Larvae make narrow, serpentine mines within the leaves of citrus, visible from the underside.

behavior

Adults fly at night and rest on the underside of leaves or tree trunks during the day. Hatched larvae bore into leaves, where they immediately begin to mine soft tissue between the upper and lower surfaces.

feeding ecology and diet

Larvae are sap feeders. Hosts include citrus and other Rutaceae and also Jasiminum, mistletoe on citrus, and kumquats. Adults feed on nectar.

reproductive biology

Mating takes place 14–24 hours after the emergence of the adult. Larvae undergo four molts, and pupation occurs within the mine near the leaf's edge. Total developmental time is about 5–20 days.

conservation status

Not threatened.

significance to humans

A significant agricultural pest of citrus throughout the world. Current infestations have been subjected to control measures. These measures include biological control by introducing Chalcidoid parasitic wasps from the Old World and pesticide applications as part of an integrated pest management program.

Large blue

Maculinea arion

family

Lycaenidae

taxonomy

Papilio arion Linnaeus, 1758, Nuremberg, Germany.

other common names

French: l'Argus arion; German: Schwarzfleckenbläuling, Schwarzgefleckte Bläuling; Spanish: Hormiguera de lunares;

Dutch: Tijmblauwtje; Finnish: Muurahaissinisiipi; Polish: Modraszek arion; Swedish: Svartfläckig blåvinge.

physical characteristics

Adults' wingspan is 0.64–0.8 in (16–20 mm). Upper side of the forewing is bright blue with large black spots; underside is grayish with large black spots and bluish or greenish area near the base. Larvae head is small, the head and legs are hidden, and the body is covered with short setae.

distribution

Western Europe to southern Siberia, Mongolia, and China.

habitat

Rough, dry, open grasslands where Myrmica sabuleti ants occur; in northern latitudes ants are restricted to warm, south-facing slopes with short grass.

behavior

Larvae are predators in underground nests of Myrmica sabuleti ants, which they attract through secretions from the honey gland.

feeding ecology and diet

Young larvae feed on pollen and seeds of wild thyme and oregano, older larvae eat ant eggs and larvae.

reproductive biology

Univoltine, adults fly for three to four weeks between June and August in small, isolated colonies; they are slow to disperse. Females may mate before their wings have dried and lay their eggs singly on flowers of thyme or oregano by the end of their first day as adults. After three weeks of feeding on host plants, caterpillars drop and hide on the ground, where ants mistake them for ant larvae and take them to nests. Pupation occurs in ant nests; emergence of adults takes place the following summer.

conservation status

Categorized as Lower Risk/Near Threatened by the IUCN. Threatened by changes in modern agriculture and land management. Extinct or declining in northern Europe; now reestablished in England, and still common in Siberia and the Far East.

significance to humans

Study of the relationship between these butterflies and ants applies to practical conservation and habitat management; also allows construction and testing of population models and generates new theories in evolutionary biology.

Gypsy moth

Lymantria dispar

family

Lymantridae

taxonomy

Phalaena (Bombyx) dispar Linnaeus, 1758. Type locality not given but certainly in Europe.

other common names

French: Spongieuse; German: Schwammspinner; Spanish: Bombyx disparate; Swedish: Lövskogsnunna.

physical characteristics

Adults are sexually dimorphic. Males are light to dark brown with irregular black markings, a wingspan of 1–1.5 in (25.4–38 mm), and bipectinate antennae. Females are all white with irregular black lines on the wings, a wingspan of 2.24–2.68 in

(56–67 mm), and narrower bipectinate antennae. Full-grown larva (1.48–2.40 in, or 37–60 mm) are gray with a series of hair tufts and five pairs of blue followed by six pairs of red spots. Pupa (0.76–1 in, or 19–25 mm) are brown with small circlets of hairs.

distribution

Palearctic region with the exception of the extreme north and south; introduced into northeastern North America.

habitat

Occurs in forests, fields, and even within cities and suburbs.

behavior

Males are active fliers in late afternoon or at night; females do not fly (except for Japanese race). Larvae remain on food plants while feeding.

feeding ecology and diet

Larvae are voracious feeders of more than 500 different trees and shrubs, including many pines, oaks, poplars, willows, and birches. Adults do not feed.

reproductive biology

Males fly to pheromone-emitting females for mating, which occurs in the summer. Females lay clusters of 100–1,000 eggs on any surface, including cars and picnic furniture, allowing for easy dispersal (hence their common name). Eggs overwinter, and larvae hatch in the spring and go through five (males) or six (female) instars, lasting 20–60 days. Adults emerge from the pupal stage (14–17 days) in late July or early August.

conservation status

Not threatened.

significance to humans

Important forest pest in Europe, Asia, and the northeastern United States. Quarantine laws in the United States and other countries have been enacted to halt further spread of this species.

No common name

Micropterix calthella

family

Micropterygidae

taxonomy

Phalaena calthella Linnaeus, 1761, Sweden.

other common names

German: Dotterblumen-Schabe.

physical characteristics

Tiny moths, with a wingspan of 0.32–0.4 in (8–10 mm). Yellow head with a tuft of hairs, functional mandibles, and no proboscis. Metallic bronzy forewings with a purplish patch at the base, similar in venation to the hind wings, with which they are coupled through the jugum. Legs and abdomen are golden brown. Larvae have eight pairs of nonmuscular conical abdominal prolegs ending in a single claw. Pupae have free appendages and articulated mandibles, used to open the cocoon.

distribution

Europe to central Siberia in Asia.

habitat

Adults fly in damp and shady habitats, such as trails and the margins of forests. Larvae are secretive, burrowing through thick mats of leaf litter, mosses, or lichens as deep as 4 in (10 cm) in loose soil.

behavior

Adults feed on flowers during the daytime and can be attracted to lights at night.

feeding ecology and diet

Adults feed on pollen from various plants with exposed flowers, from trees and shrubs to grasses; they have a preference for species of Ranunculaceae, a primitive group of angiosperms. Larvae are external feeders on decayed plant detritus, fresh leaves of angiosperms, liverworts, and possibly fungal hyphae.

reproductive biology

Eggs are laid singly or in clusters of two to 45 on the food plant. Larval development takes from 132 to 141 days. It is thought that there are three larval instars, with the last one overwintering. Pupation occurs in early spring inside a silken cocoon on the ground or on detritus; adults emerge in spring and fly from May to June.

conservation status

Not threatened.

significance to humans

None known.

Corn earworm

Helicoverpa zea

family

Noctuidae

taxonomy

Phalaena zea Boddie, 1850, North America.

other common names

English: Tomato fruitworm, sorghum headworm, vetchworm, cotton bollworm; Spanish: Gusano cogollero, tomatero, bellotero, gusano del fruto.

physical characteristics

Adults are medium-sized and sandy-colored, with a wingspan of 1.50 in (37.5 mm). Forewings have faint, irregular markings on the apical half; the hind wings are a dull translucent white with a narrow dark apical band. Full-grown larvae (1.28–1.60 in, or 32–40 mm) are variously colored red, pink, or green with irregular longitudinal dark stripes. Body has spinules.

distribution

Southern Canada to Argentina.

habitat

Adults are found on flowers at dusk and can be attracted to lights; larvae are found on ears of corn, tomatoes, and other crops.

behavior

Adults fly mainly at dusk. Hatched larvae tunnel into fruit often associated with frass (insect excrement) but can be found on flowers, leaves, and seedlings.

feeding ecology and diet

Larvae are serious pests of corn, tomatoes, and cotton. Occasional hosts include bean, broccoli, cabbage, chrysanthemum, eggplant, head cabbage, lettuce, okra, pepper, and more than 100 other plants.

reproductive biology

Females can lay eggs on corn silks, on terminal leaflets of tomatoes, and on the crowns of seedlings of lettuce. Larvae pupate about 2–6 in (5–15 cm) beneath the soil and overwinter.

conservation status

Not threatened.

significance to humans

A major pest of several important food crops, necessitating various forms of integrated pest management, especially in the warmer parts of its range.

Blue morpho

Morpho menelaus

family

Nymphalidae

taxonomy

Papilio menelaus Linnaeus, 1758, Guianas.

other common names

English: Blue morpho butterfly; Spanish: Mariposa azul.

physical characteristics

Iridescent blue wings. The female is duller, with a brown edge and white spots surrounding blue. The undersides are brown with bronze eyespots. The wingspan is 6 in (150 mm). Larva are red-brown with bright patches of lime green and reddish-brown and white tufts of hair on the dorsum.

distribution

South America, from the Guianas to Brazil and Bolivia.

habitat

Rainforests.

behavior

Adults soar in the canopy of the jungle, coming near the ground in clearings; the underside of the wings provides camouflage at rest. The slow beating of the wings gives rise to alternating flashing and disappearance of their iridescent blue coloring, possibly startling potential predators. Males are territorial; their vivid blue is used to intimidate other males. Larvae are active at night and, if threatened, release a strong smell from scent gland located between the forelegs.

feeding ecology and diet

Adults imbibe the juices of rotting fruit; larvae feed on the neotropical plant Erythroxylum pilchrum.

reproductive biology

Nothing is known.

conservation status

Not threatened.

significance to humans

Owing to the microscopic structure of the wing scales, the wings are iridescent, and their color changes when seen from different angles. This quality makes them useful as models to create counterfeit-proof currency and charge cards.

Queen Alexandra's birdwing

Ornithoptera alexandrae

family

Papilionidae

taxonomy

Troides alexandrae Rothschild, 1907, Biagi, southeastern New Guinea.

other common names

None known.

physical characteristics

Adults are sexually dimorphic. Males have long (3.24–4.36 in, or 81–109 mm; wingspan of 6.7–7.5 in, or 170–190 mm), iridescent blue and black, ellipsoid forewings and smaller hind wings. The thorax is crimson red on the sides, and the abdomen is brilliant yellow. Females' wings are larger (4.08–5.16 in, or 102–129 mm long; wingspan of 7.1–8.3 in, or 180–210 mm) and brown; the outer half of the hind wings has creamy white, elongate, wedge-shaped spots. The thorax is the same as in the male, and the abdomen is creamy white. Eggs are round, white, and up to 0.20 in (5.1 mm) in diameter. Full-grown larva (up to 4.72 in, or 118 mm) are dark wine red and covered with fleshy tubercles, with a pale yellow saddle mark on fourth abdominal segment. Pupa (up to 3.6 in, or 90 mm) are light brown with irregular patches of yellow.

distribution

Restricted to lowland areas of Northern Province of Papua New Guinea.

habitat

Occurs in primary and secondary rainforest; larvae live on the food plant.

behavior

Adults are diurnal and fly high during the day, visiting flowers. Larvae remain motionless on the food plant when not eating. If touched, larvae extend the osmeterium. Before pupating, larvae may ring a branch or stem of the food plant, thereby killing it. Larvae often wander for 24 or more hours in search of a proper pupation site, usually under leaves from 4 in (10 cm) off the ground to high in the canopy.

feeding ecology and diet

Adults imbibe nectar at flowers of introduced ornamentals, Caesalpinia and Zanthoxylum, often high up in forest canopy. Larvae feed on Pararistolochia alexandriana and P. meridionaliana. First-stage larvae eat their eggshells; younger larvae eat young, tender leaves; and older, full-grown larvae consume larger, older leaves.

reproductive biology

Low egg fecundity (about 240 eggs per female). Females inspect the food plant carefully before depositing eggs under-neath a leaf. Larvae normally undergo five molts, lasting about 70 days. Pupation lasts about 40 days. Marked adults have been seen up to three months after initial marking.

conservation status

Listed as Endangered by the IUCN and by CITES I and the U.S. Fish and Wildlife Service. These acts effectively ban the commercial exchange of this species. Its major threat is habitat destruction due to the ever encroaching oil palm and timber industries. The best hope for conserving O. alexandrae may be the commercial breeding of specimens, since its high demand probably will ensure its survival.

significance to humans

The species is arguably the flagship emblem of butterfly conservation, and it often is illustrated in many books relating to invertebrate species conservation.

European cabbage white

Pieris rapae

family

Pieridae

taxonomy

Papilio rapae Linnaeus, 1758, Sweden.

other common names

English: Small white; French: Petit blanc du chou; German: Kleiner Kohlweissling; Spanish: Blanquita de la col; Polish: Bielinek rzepnik.

physical characteristics

Wingspan of 1.75–2.25 in (4.5–5.8 cm). The upper side of wings is white; forewing has black tip. Two submarginal black spots in the female and one in the male. Undersides evenly yellow-green or gray-green. Eggs are pale yellow, bottle-shaped, ridged, and 0.06 in (1.5 mm) high. Larva (up to 0.75 in, or 3 cm) are green with pale yellow line on the back and a line of yellow spots on each side. Pupa sculptured and angular; color influenced by background color.

distribution

Palearctic region with the exception of extreme north and south; introduced into North America and Australia.

habitat

Almost any type of open space, including weedy areas, gardens, roadsides, cities, and suburbs.

behavior

Female is active during the day, with slow, lumbering, erratic flight; male flies in a straight line. Larva is a slow crawler and sits on the upper surfaces of leaves of food plants in daylight, camouflaged by coloration; when disturbed it regurgitates a poisonous fluid.

feeding ecology and diet

Larva feeds on plants containing mustard oil (among them, cabbage, radish, broccoli, mustard, and other Brassicaceae species). Adult feeds on the nectar of several flowers, including mustards, dandelion, red clover, asters, and mints.

reproductive biology

Males patrol for females. Females lay single eggs on the under-sides of the host plant. Larval stage takes 17 days; pupa over-winters, and adults emerge in summer. Adults breed all year in subtropical areas; there are three to five generations in temperate regions.

conservation status

Not threatened.

significance to humans

Considered a pest; larvae feed on vegetable gardens and crops.

Yucca moth

Tegeticula yuccasella

family

Prodoxidae

taxonomy

Pronuba yuccasella Riley, 1872, central Missouri.

other common names

English: Tegeticula moth.

physical characteristics

Adults small and white; wingspan is 0.07–0.11 in (1.8–2.75 mm). Forewings are white, wide, and blunt; hind wings are brownish-gray. Females are larger than males, with a swordlike ovipositor and enlarged maxillary palps. Larvae are pink and more than 0.4 in (10 mm) long. They form a spherical or elongate cocoon, about 0.24–0.32 in (6–8 mm) long.

distribution

Southern Canada south through Mexico.

habitat

Associated with yucca plants growing in sand dunes, grasslands, pine forests, and glades.

behavior

Adults hide inside yucca flowers during the daytime and fly at night. Between dusk and midnight females gather pollen from flowers using their maxillary palps; they form balls of sticky pollen and push them into the receptive tips of yucca pistils (stigmata). Larval stages grow inside developing yucca fruits.

feeding ecology and diet

Larvae feed on developing yucca seeds, consuming a small percentage of the hundreds of seeds within capsules.

reproductive biology

Mating occurs inside yucca flowers; the gravid female visits flowers and lays an egg in one of the six locules (cavities within the ovary). The larva remains inside the little feeding cavity of fused seeds within the capsule until the first autumn rains and then drops to the ground, burrows into soil, and overwinters in a silk-and-sand cocoon. Pupation occurs after the onset of spring rains, and the adult emerges in late May or in June. Adults stay close to their home yucca clusters and remain active for less than a week.

conservation status

Not threatened.

significance to humans

The association between yucca and moth constitutes a classic example of "mutualism" and "co-evolution" and provides a unique opportunity for studying these phenomena. The yucca plant cannot produce seeds without the moth, and without the plant the moths would die off in one generation.

Bagworm

Thyridopteryx ephemeraeformis

family

Psychidae

taxonomy

Sphinx ephemeraeformis Haworth, 1803, Great Britain (apparently in error).

other common names

English: Evergreen bagworm moth.

physical characteristics

Adult males have short (0.5 in, or 12.5 mm; wingspan of 1.12 in, or 24 mm), clear wings; hairy black bodies; and feathery antennae. Females resemble maggots, with no functional eyes, legs, wings, mouthparts, or antennae. Larvae live inside a bag made of silk and bits of needles, bark, or twigs and are up to 1.5–2.5 in (40–65 mm) long.

distribution

Eastern United States.

habitat

A wide range of broadleaf and evergreen trees and shrubs (128 species) serve as hosts, including arborvitae and other ornamental conifers, sycamore, and willow.

behavior

The female remains inside the bag her entire life; males are nimble fliers and in the fall fly around infested trees in search of a mate. During feeding, caterpillars emerge from the top of the bag and hang on to the host plant with their legs and sometimes with a silken thread; the bottom of the bag remains open to allow fecal material to pass out. Young larvae disperse, walking or using wind currents. During molts and pupation, caterpillars seal the bags.

feeding ecology and diet

Larvae feed on conifers, maple, oak, dogwood, and willow, consuming one branch at a time and leaving only the middle rib of leaves.

reproductive biology

Univoltine, they overwinter as eggs inside the mother's bag. Eggs hatch in spring. Larvae crawl out in search of food and construct a bag, where they molt four or more times before pupation. Adults emerge in the fall. Males are attracted to the female's bags by pheromones and mate. Female lay 500–1,600 eggs within bag, after which they drop to the ground and die.

conservation status

Not threatened.

significance to humans

Pest; defoliates evergreen ornamentals.

No common name

Parargyractis confusalis

family

Pyralidae

taxonomy

Cataclysta confusalis Walker, 1865, North America.

other common names

None known.

physical characteristics

Adult is small, with a forewing span of 0.20–0.44 in (5–11 mm). Light tan with well-defined darker areas of brown. Hind wing broadly triangular with a marginal row of small, eyelike spots. Larva (0.60 in, or 15 mm) is greenish with brown head capsule, thoracic shields, and filamentous clusters of gills along the side. Pupa (0.32 in, or 8 mm) is brown in feltlike cocoon. All immature stages found on rocks in creeks and streams.

distribution

Northern California east to Montana, Idaho, and Nevada and north to southern British Columbia.

habitat

Near and in creeks and streams.

behavior

Adults often found at rest on bridge supports crossing streams or on bushes, trees, and other vegetation bordering streams.

feeding ecology and diet

Larvae feed on algae and diatoms on substrate (e.g., rock crevices) in streams.

reproductive biology

Female swims to the bottom of riffle and lays a few to several hundred eggs. Larvae live under protected silken areas on alga-covered substrate; pupation occurs in these waterproof silken linings. Adults emerge and swim to the surface.

conservation status

Not threatened.

significance to humans

None known.

Indian mealmoth

Plodia interpunctella

family

Pyralidae

taxonomy

Tinea interpunctella Hübner, 1813, Europe.

other common names

French: Pyrale indienne de la Farine, teigne des fruits secs; German: Kupferrote, Dörrobstmotte; Spanish: Polilla de la fruta seca; Portuguese: Traça dos cereais.

physical characteristics

Adults are small, with a wing length of 0.20–0.34 in (5.0–8.5 mm) and a wingspan of 0.62–0.79 in (16–20 mm). Forewing has reddish-brown apical half and ocher basal half; hind wing is dull translucent white. Full-grown larvae are 0.36–0.60 in (9–15 mm), with yellow-brown head capsule and prothoracic shield; remainder of the body is dull white to pinkish.

distribution

Cosmopolitan.

habitat

Often seen in pantry areas of homes, in supermarkets, and in feed stores, but adults also occur outdoors and are attracted to light at night. Larvae feed on stored foods.

behavior

Adults fly at night. Hatched larvae tunnel into food often associated with frass and silk webbing.

feeding ecology and diet

Larvae are significant nuisance pests of dried stored foods; characteristic webbing is associated with larval feeding. Adults often are seen flying in these areas.

reproductive biology

Females can lay eggs three to four days after emerging as adults. Larval development can take 13–288 days, depending on temperature and humidity. Generations overlap in home and warehouse situations.

conservation status

Not threatened.

significance to humans

A minor to significant pantry pest of stored dried food materials. Infestations can be persistent, requiring cleanliness and exclusion practices such as hermetically sealed containers for food.

Atlas moth

Attacus atlas

family

Saturniidae

taxonomy

Phalaena atlas Linnaeus, 1758, "Citro Asiae, Americae (fixed as Bogor, Djawa, Indonesia)."

other common names

English: Giant comet moth, giant silkmoth; French: Papillon géant; Italian: Farfalla cobra; German: Tropischer Spinner.

physical characteristics

Wings are reddish-brown with a triangular transparent spot (membrane without scales). The wingspan is 8 in (20 cm), and the tips of the forewings are curved. Stout, hairy body. Males have feathery antennae; females are larger than males. Large, spherical, reddish eggs. Larvae are bluish-green with shades of pink; they have bumps and are covered with a fine white powder. Pupate in cocoons made of broken strands of silk.

distribution

Tropics of Asia, including India and Southeast Asia.

habitat

Lowland to upper mountain forests.

behavior

Active at night, irregular flight, attracted to light. Females position themselves to enable maximum dispersal of their sexual attractants by wind. Males up to three miles downwind can detect these scents. Larvae wander, looking for food.

feeding ecology and diet

Larvae are polyphagous: feed on a wide range of trees (e.g., Jamaican cherry tree, soursop, cinnamon, rambutan, guava, and citrus). Adults have atrophied mouthparts and do not eat.

reproductive biology

After mating, the female lays small groups of up to several hundred eggs on the undersides of leaves; adults die a few hours later. Eggs require eight to 14 days to hatch, depending on temperature. Pupal stage lasts about four weeks.

conservation status

Not threatened.

significance to humans

Larva are cultivated commercially around the world by collectors. In Taiwan cocoons are used as pocket purses and in northern India to make Fagara silk.

Death's head hawk moth

Acherontia atropos

family

Sphingidae

taxonomy

Sphinx atropos Linnaeus, 1758, Europe.

other common names

English: Bee robber; German: Totenkopfschwärmer; French: Sphinx têtede-mort, fluturele cap-mort; Spanish: Mariposa de la muerte; Estonian: Tontsuru; Swedish: Dödskallefjäril;

physical characteristics

Adult has skull-like pattern on thorax. Large and heavily built, with 4.4–4.8 in (11–12 cm) wingspan. Dark forewings and yellow hind wings with black submarginal lines. Proboscis is short, stout, and hairy. Abdomen has yellow riblike markings. Larva is 4.8–5.2 in (12–13 cm) and colored yellow, green, or brown with a large posterior horn. Pupa grows to 3–3.2 in (7.57ndash;8 cm) long and is mahogany brown and glossy.

distribution

Afrotropical, extending north to Mediterranean; migrant in central and northern Europe.

habitat

Prefers dry and sunny locations; frequents open scrubs with solanaceous (nightshade) plants and cultivated areas where potato is grown.

behavior

Larvae are inactive, moving only to find a fresh leaf; when disturbed, they click their mandibles and may even bite. Adults are active from dusk to midnight; during the day they rest on tree trunks, walls, or leaves on the ground. Attracted to light and occasionally to blossoms. Frequent beehives, where they rob honey. In defense, they mimic honeybees' cutaneous fatty acids and raise their wings, run, and hop around. When disturbed, they emit loud, shrill squeaks, forcing air out the proboscis, followed sometimes by secretion of moldy smell from glandular hairs in the abdomen.

feeding ecology and diet

Larvae prefer solanaceaous plants, especially potato. Short proboscis prevents adult from taking nectar from deep-set flowers; instead, they imbibe honey, juice from rotten fruits, and sap from trees.

reproductive biology

Breed year-round in Africa; adults migrate to Europe from May to September but do not overwinter there. Eggs are laid singly underneath old leaves of host plant. Pupation takes place in a very fragile cocoon 6–16 in (15–40 cm) deep in the soil in a smooth-sided cavity.

conservation status

Not as common as it used to be, owing to use of insecticides.

significance to humans

In Greek mythology, Atropos is the eldest of the three Fates, who severs the thread of life. The moth is considered a sinister creature because of its skull design and the loud sound emitted when it is disturbed. It once was thought to be a harbinger of war, pestilence, and death, and it has entered modern mythology as an emblem of perverted evil in the book and film Silence of the Lambs, in which the trademark of the serial murderer is a pupa placed in the mouth of his female victims, whom he later skins.

Webbing clothes moth

Tineola bisselliella

family

Tineidae

taxonomy

Tinea bisselliella Hummel, 1823, most likely Europe.

other common names

English: Common clothes moth; French: Mite des vêtements; German: Kleidermotte; Spanish: Polilla de la ropa; Dutch: Kleermot; Finnish: Vaatekoi; Norwegian: Klädesmal.

physical characteristics

Adults grow to 0.16–0.28 in (4–7 mm) in length, with a wingspan of 0.6 in (1.5 cm). Golden brown with a tuft of reddish-yellow hairs on the head. Narrow wings fringed with long hairs are reddish-yellow with no markings. Larvae are yellowish with dark brown heads and grow to 0.4 in (1 cm) long; they live in a tube of silk, excreta, and other debris found in the area where they feed.

distribution

Originally from the Old World; now cosmopolitan with the exception of the tropics.

habitat

Can inhabit any storage area. Breeds at 50–91.4°F (10–33°C). Its optimum relative humidity is 70%.

behavior

Adults tend to fly in darkened areas. Males are the fliers, whereas females generally walk or run. Soon after hatching, larvae begin constructing a silken tube. This case acts as shelter during the day, offering the larva good camouflage, from which it emerges at night to feed.

feeding ecology and diet

Caterpillar commonly feeds in dark, protected areas on woolens, furs, feathers, animal bristle brushes, dead insects, dried animal carcasses, pollen, and other dried plant and animal products. Adult mouthparts are atrophied, and they do not feed.

reproductive biology

The female lays about 50 eggs in darkened areas, such as natural fibers and cloth; afterward it dies. Larvae pass through five instars, although under adverse conditions there may be as many as 40. Pupation takes place within the silken case. Total life span varies from five to nine months; it may exceed two years if the larva goes into a dormant period. In heated buildings females can mate and lay eggs at any time during the year.

conservation status

Not threatened.

significance to humans

This household and industrial pest of natural fibers can also infest dried vegetable material.

Resources

Books

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.

Periodicals

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

Organizations

Association for Tropical Lepidoptera. P. O. Box 141210, Gainesville, FL 32614-1210 United States. Phone: (352) 392-5894. Fax: (352) 373-3249. E-mail: [email protected] Web site: <http://www.troplep.org>.

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 [email protected].

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

Natalia von Ellenrieder, PhD

Rosser W. Garrison, PhD