- Key People:
- John Henry Comstock
- Related Topics:
- caterpillar
- moth
- butterfly
- skipper
- Pterophoroidea
Within the limitations imposed by their almost entirely plant-eating habits, the lepidopterans show a great diversity of size, structure, and other characteristics. Furthermore, some members of the order have retained many primitive features, while others have evolved very advanced and specialized ones. This broad range, from primitive to advanced and from generalized to specialized, is evident not only in the adults but also in the larvae and pupae and to some degree in the eggs.
The adult
Nearly all external surfaces of the adult are covered with scales, which may be broad and flat or long and hairlike. Each of these is the outgrowth of a single epidermal cell. Similar scales occur in a few groups of other insects but never to the same extent.
Like most insects, adult lepidopterans have three distinct body segments, or tagmata—the head, the thorax, and the abdomen—each with special functions. The head bears the main sensory organs and those of feeding and ingestion. The thorax is chiefly concerned with locomotion. The abdomen contains the main organs of digestion, excretion, and reproduction and bears the external accessory reproductive structures.
Head
The head is relatively small and round or elliptical. With regard to its evolutionary development, it is derived from the first six primitive body segments (somites, or metameres), but these have become so coalesced that none of the primitive segmentation is evident. The antennae are prominent and multisegmented, with many microscopic receptors (sensilla) for detecting odors. In most moths the antennae are slender and tapering, although in some the surface area of the antennae is greatly enlarged by many side branches that make them comblike or featherlike. In skippers and butterflies the terminal part is expanded into a clublike shape, on which most of the sensilla are concentrated. In many families there is also a cluster of sensory bristles (the chaetosema) on each side of the head near the eye. On either side of the head is a large compound eye, sometimes consisting of thousands of units (ommatidia). Most moths have, in addition to the compound eyes, a pair of very small simple eyes (ocelli), which have limited light-sensing ability but do not form an image.
The compound eyes are very efficient for distinguishing motion but not for resolving clear images of distant objects. Diurnal species may be able to distinguish flower shapes, and courting individuals have been shown to respond to specific wing patterns. Color perception, especially in the blue-violet end of the spectrum, is acute in some species. Moths are generally able to see by ultraviolet light, to which they are often strongly attracted at night.
The sense of smell is acute in some groups, especially in males with large antennal surfaces. The sense of taste, especially for sugars, is also acute. In moths and butterflies, taste receptors are located chiefly on the palpi and on the feet (tarsi).
Nearly all caterpillars have chewing mouthparts, but these are retained among adults of only the most primitive moths—the mandibulate moths of family Micropterigidae. Caterpillar mouthparts basically consist of an anterior flap (labrum), a pair of chewing jaws (mandibles), a pair of complex first maxillae, and a pair of similar second maxillae joined together behind the mouth to form a structure called the labium. Each of the first and second maxillae bears a jointed sensory appendage, or palpus. All these structures function together for chewing and manipulating solid foods. In the vast majority of adults the mandibles are either vestigial and nonfunctional or entirely absent. Parts of the first maxillae, however, are elongated to form the two halves of a tubular proboscis (haustellum) through which liquids may be sucked. The segmented palpi of the first and second maxillae are present and function as sensory organs. Not all adults have all these parts fully formed and functional. In numerous families the proboscis has become considerably reduced and even vestigial, resulting in adults that cannot feed. In advanced moths and in skippers and butterflies, the maxillary palpi are vestigial or lost, so that only the labial palpi remain functional.
Mouthpart reduction in adults appears to have occurred independently in a number of evolutionary lines. The result in these species has been to limit feeding to the larval stage and thus enable the adult stage to be dedicated only to reproduction and dispersal.
Thorax
The thorax consists of three segments, the prothorax, mesothorax, and metathorax, each derived from a primitive segment. The prothorax bears the first pair of legs and a pair of respiratory openings (spiracles). The much larger mesothorax bears the second pair of legs, a second pair of spiracles, and the pair of forewings. The metathorax bears the third pair of legs and the pair of hind wings. In many moths the metathorax bears a pair of complex auditory organs (tympana). In some species these organs serve as receptors of the high-frequency echolocation signals emitted by hunting bats, allowing the moths to initiate escape maneuvers. In other species the auditory organs are receptors of mate location calls. Sound signals are produced in some species by timbal organs and in others by a mechanical clicking of the wing base.
The wings begin development in the maturing larva as invaginations of the epidermis. As the pupa is formed, these fold out (evaginate) to lie externally and become large and flat. Within them, branching tubes (tracheae) carry an air supply and also mark the pathways along which will develop the tubular “veins” that support the fully formed wings. When the adult emerges from the pupa, the wings expand to full size. On the wing the scales lie in overlapping rows, like shingles. The usual scale is a flattened, rigid, air-filled sac attached by a peglike base. It is usually ribbed longitudinally and toothed terminally. Many males have special glandular scent scales (androconia) scattered or concentrated in patches (brands or stigmata) on the wings, sometimes forming expansible, hairlike tufts. These have a scent-distributing function that is essential in specialized courtship.
The scales and hairs of the adults, especially on the wings, are responsible for colors and patterns. There are many different pigments of several chemically diverse types such as melanins, uric acid derivatives, and flavones. In addition, the microstructure of hairs and scales refracts light in various ways so as to produce colors as well as metallic, pearly, iridescent, and white effects. Very fine parallel ridges in scales may produce an iridescence by acting as diffraction gratings. More common are the effects of very thin superimposed layers (laminae) in the walls of the scales, a color phenomenon comparable to that produced by a thin film of oil on water. Many of the most striking effects are caused by combinations of pigmental and structural colors.
The forewings and hind wings on each side are coupled together in various ways. In primitive moths a fingerlike lobe on the forewing overlaps the base of the hind wing. In most moths a strong bristle or cluster of bristles (frenulum) near the base of the hind wing engages a catch (retinaculum) on the forewing. In some moths and in the skippers and butterflies, the frenulum mechanism has been lost, and the wings are coordinated by the friction of the overlapping areas. In the most primitive moths the forewings and hind wings are similar in size, shape, and veinage. In most moths and in the skippers and butterflies, the hind wings have become shorter and more rounded, with reduced veining except in the posterior wing section. The anterior, or leading (costal), edge of the forewing is thickened, with stronger veins, while the outer and posterior (anal) wing section margins are thinner and weaker. This accords with the function of the wings as airfoils having a stiff leading edge and a flexible trailing edge.
The wings are characteristically modified in various families. In many moths of the superfamily Tineoidea, both wings have become extremely narrow, with much-reduced venation. However, along the margins there are long, dense fringes of hairs that maintain the functional wing area. In the plume moths (family Pterophoridae) the wings are deeply cleft into two or three narrow plumes, and in the many-plumed moths (family Alucitidae) each wing is cleft into six plumes. In a number of moth families the females are wingless, although the males are fully winged (bagworm moths [family Psychidae], some tussock moths [family Lymantriidae]). In the aquatic snout moth (Acentropus) some females are wingless, while some females and males are winged. There are eight main wing veins, each with a characteristic pattern. These are usually designated according to the modified Comstock-Needham system. The names of the veins (with their symbols in parentheses) and the usual number of branches of each (subscript designations) are as follows, in progression from the costal margin of the wing to the anal margin:
- Costa (C), along the anterior margin, no branches; Subcosta (Sc), usually with no branches, rarely with two (Sc1, Sc2);
- Radius (R), typically with five branches (R1–R5); Media (M), with three branches (M1–M3; four in some other insects);
- Cubitus (Cu), primitively with three branches (Cu1a, Cu1b, Cu2);
- Anal veins, First (1A), Second (2A), and Third (3A), all unbranched.
Abdomen and genitalia
The abdomen has 10 segments, although the posterior ones are indistinct. Each of the first eight segments bears a pair of spiracles. The first or second segments bear paired auditory organs in the snout moths and measuring worm moths. Segmental appendages are absent except for vestiges that may form parts of the genitalia. Various segments may bear special structures that produce and disperse pheromones. The genitalia of both sexes are often complex and bear characteristic spines, teeth, setae, and scale tufts. These structures are important in complex courtships and matings, preventing hybridization between males and females of different species.
In males a ringlike structure is the base of attachment for a number of dorsal structures and a pair of lateral clasping organs (valvae). In copulation a median tubular organ (the aedeagus) is extended through an eversible sheath (vesica) to inseminate the female. These structures are derived evolutionarily from parts of segments 8 and 10 and from vestiges of abdominal appendages.
The female genitalia show a number of different types of organization of the internal genital ducts and openings. These are considered so fundamental that the lepidopterans can be classified into suborders largely on the basis of these traits and their correlation with characteristics of the mouthparts, wings, and early developmental stages.
The internal reproductive systems of both sexes contain the organs typical of most insects. The testes of the male are paired in primitive lepidopterans but fused into a single organ in advanced forms. In both cases the sperm ducts are paired. As in other insects, the sperm pass from the testes down these paired ducts (vasa deferentia) for storage in sacs called seminal vesicles. Accessory glands, providing fluids that lengthen the life of the sperm, open into the vasa deferentia.
The female reproductive system consists of paired ovaries, paired accessory glands that provide the yolks and shells of the eggs, and a system of receptacles and ducts for receiving, conducting, and storing sperm. The individual oviducts join to form a common oviduct that leads to the vagina. In copulation the male deposits a sperm capsule (spermatophore) in a receptacle of the female. The spermatophore releases the sperm, which swim into the oviduct and thence to the seminal receptacle, where they are stored until egg laying. This may occur hours, days, or months after mating.
The egg
The egg is enclosed in a protective layer (the chorion), through which a system of tiny canals (micropyle) permits the entrance of sperm. In some groups the micropyle is at the side, whereas in others it is on the surface away from the substrate. The egg passes along the individual oviduct and through the common oviduct to the vagina. Here, just before it is laid, it comes into contact with a droplet of seminal fluid that has been stored in the female, and fertilization takes place. Most eggs are more or less spherical, but those of a few families are flat or long and tapered at the ends. Their surface may be strongly sculpted with pits, sharp projections, or raised ridges.
The larva, or caterpillar
Compared with the highly specialized adult, the larva is simple and primitive. Many of the primitive characteristics retained in the larva are important in the classification of the suborders, superfamilies, and families. The traits also aid in making speculations on relationships among these groups (taxa).
The head bears a pair of very short antennae and on each side a cluster of minute simple eyes (stemmata). A short liplike labrum is in front of the mouth. Behind the labrum are paired jaws (mandibles) that are short, broad, and powerful to allow consumption of large amounts of plant material. Next is a pair of small first maxillae, each with a segmented palp. Then, more or less connected with the maxillae, is the labium-hypopharynx, a complex structure with a pair of labial palps between which is located a tubular spinneret through which silk is extruded. Each of the three thoracic segments bears a pair of short segmented legs. The remaining 10 segments constitute the abdomen. Abdominal segments 3 through 6 and segment 10 bear a pair of fleshy appendages called prolegs, which may be homologous with the primitive segmental appendages. Each proleg has one or two curved rows of minute hooklets and an eversible soft end, the planta. The crochets on the prolegs allow the larva to hold onto surfaces. Body fluids forced into the proleg cause it to expand, extending the hooklets. After the proleg has been placed on the substrate, the fluids are retracted into the body and the elasticity of the cuticle causes the tiny hooks to retract, thus gripping the substrate. Prothorax and abdominal segments 1 to 8 have a pair of spiracles on them.
The larval epidermis bears on the head and each body segment a number of bristles known as primary setae. The position and number of setae are constant in each species and so are important in classification. Often there are many secondary setae, giving the caterpillar a hairy appearance. Larvae that live and feed as borers, burrowers, or miners are mostly plain. Those that live and feed in the open often show a great diversity of shape, color, pattern, and ornamentation. Many have wartlike projections (verrucae) that may bear tufts of setae or spiny projections, or there may be prominent hornlike or spiny processes.
In some families the number of prolegs has been modified. In the measuring worms the prolegs of segments three, four, and five are missing, and in some owlet moths one or two pairs have been lost. In the puss moth caterpillars the last pair has evolved into a pair of long eversible whiplashes (stemmatopoda). The larvae of some leaf miners (family Gracillariidae) have lost some or all of the prolegs. Mandibulate moth larvae have eight pairs of abdominal legs, which are structured more like true thoracic legs than prolegs.
Internally, the larva is relatively simple, with the very large digestive tract being the most prominent organ. The paired silk glands are often very large, extending far back into the abdomen. The ovaries and testes, which begin to develop during embryonic life, continue to develop in the larva, as do the wings. Many special glands secrete repellent or toxic substances, which may circulate in the blood or be extruded from special openings as a means of defense.
Larval vision can detect little more than differences between light and darkness. Taste is acute, with highly developed sensory receptors in the antennae and palpi. Food discrimination is keen, and many larvae will starve rather than eat abnormal food plants. The sense of touch functions via setae widely distributed over the outer surface. Some of these appear to react to sound waves of low pitch, well within the limits of human hearing.
The pupa, or chrysalis
Lepidopteran pupae show the same sort of evolutionary gradation from primitive to advanced as do larvae and adults. In the primitive mandibulate moths and sparkling archaic sun moths (family Eriocraniidae), the pupa has free and movable appendages and functional mandibles. In some less-primitive groups the pupa retains the ability to move some appendages. In the higher moths and the butterflies, all appendages are tightly fastened to the body wall. Called obtect pupae, these are immobile and able to wriggle only one or two abdominal segments. In a few groups the pupa has special stridulating rasps for sound production. Nearly all of the external structures of the adult can be seen on the pupa. The wings are prominent, folded down flat along the ventral surface, with the proboscis halves, the legs, and the antennae between them. At the posterior end is a spiny pad or spike, the cremaster, which in many groups attaches the pupa to silk fibers spun by the larva.
Except for a very slight respiratory exchange and a little water loss, the pupa is physiologically self-sufficient. Within it most of the cells and tissues of the larva undergo considerable histolysis (breakdown) as the adult structures are built up from the existing rudiments. Some structures begin developing as far back as the first larval stages.
Growth, molting, and metamorphosis
As in other insects, growth and its structural changes are controlled by an interacting set of hormones. These are chiefly secreted by the corpora allata and other parts of the brain and by paired prothoracic glands. The prothoracic gland hormone is necessary for larval molting (ecdysis), metamorphosis to the pupa, and formation of adult characteristics. On the other hand, a hormone secreted by the corpora allata inhibits metamorphosis until late larval development. A hormone secreted by cells in the pupal brain stimulates the prothoracic glands and thereby brings about differentiation of the adult and the end of the obligatory resting stage (diapause) of the pupa.
Evolution and paleontology
The Lepidoptera belong to an important group of insect orders called the panorpoid complex. This ancestral stem began in the Permian Period (298.9 million to 251.9 million years ago) and split into a number of branches, from which evolved the modern scorpionflies (order Mecoptera); dobsonflies, alderflies, and lacewings (see neuropteran); true flies (order Diptera); caddisflies (order Trichoptera); and the lepidopterans. The nearest living relatives of the Lepidoptera are the caddisflies, and in fact the very primitive mandibulate moths (family Micropterigidae) have been grouped with the caddisflies by some systematists. As the Lepidoptera developed into a distinctive group, certain major trends began to be emphasized. The adult mandibles disappeared and were replaced by the proboscis formed from the galeae of the maxillae, which broadened the food base exploitable by the species to include not only sap and other plant juices but nectar and fruit juices as the flowering plants evolved. Various groups, however, independently lost the proboscis and concentrated on the larval stage as the sole source of nutrition. The characteristic vein pattern evolved, with the long, veinless discal cell in each wing, as well as an emphasis on the strength of the forewing and a de-emphasis of the hind wing, forming a particular pattern of aerodynamic efficiency. Quite a few groups of small moths, settling into restricted ecological niches where strong flight is not a necessity, evolved a great deal of wing reduction.
Concurrent with changes in the adults, the larvae were evolving a multiplicity of different ways of feeding on the evolving seed plants. As both larval and adult differentiations accumulated, it became more and more necessary that the pupal stage be a passive, resting phase to allow time for the metamorphosis of larva into adult.
Very few fossil Lepidoptera are known. The earliest, dating from the beginning of the Paleogene Period, 66 million years ago, are small moths related to the superfamilies Eriocranioidea and Tineoidea. However, given their characteristics, the order must have existed long before then. The closely related caddisflies are known from the Jurassic Period (201.3 million to 145 million years ago). The best-known fossil butterflies are relatively recent, found in the shales of the Florissant Formation, a deposit laid down in Colorado at the Eocene-Oligocene boundary some 33.9 million years ago. Some of those are undoubtedly brush-footed butterflies (family Nymphalidae), which are very similar to modern genera.
Classification
Distinguishing taxonomic features
The chief characteristics used in the ordinal, subordinal, and family-level classification of the Lepidoptera are: the mouthparts, ranging from mandibulate to haustellate (with a proboscis) or degenerate; the venation and shape of the wings, homoneurous (the venation of the forewings and hind wings alike) or heteroneurous (forewings and hind wings different), aculeate (more or less covered with specialized bristles called microsetae) or nonaculeate, and type of coupling (jugum or frenulum); the anatomy of the female reproductive system; the exarate or obtect condition of the pupa; and the larval structure and pattern of the primary setae.
The venation of the wings is perhaps the most important single criterion for establishing both differences and relationships in the classification. However, venation patterns must be considered in terms of the evolution of these patterns from primitive to advanced conditions within individual phyletic lines. The most primitive groups tend to have the maximum number of veins and branches in each wing. More advanced groups lack some veins or branches in the forewing and have the hind-wing venation considerably reduced.
The structures of the genitalia are extremely important in classification, often serving better than any other characteristics to distinguish the species. The various parts have been homologized with each other, thus enabling comparative studies of families, genera, and species.
The female genitalia exhibit a number of different patterns of the internal ducts and the openings. These vary from a condition in which there are no special genital openings, with insemination and egg laying (oviposition) taking place through a single aperture shared with the excretory system, to one in which there are two specialized openings, one for insemination and one for oviposition, both of which are distinct from the anus.
Annotated classification
The classification presented follows Niels P. Kristensen (1999). Classification systems vary in the way they categorize taxonomic groups. Traditional taxonomic schemes gather superfamilies and families into suborders; more recent systematic schemes have abandoned suborders in favour of less rigidly organized clades. In the scheme followed below, neither suborders nor clades are indicated; instead, only superfamilies and families are shown. Of the 46 lepidopteran superfamilies, 24 are presented below, representing all the commonly encountered families of moths, butterflies, and skippers. Superfamilies and families not presented below include all moths that are rarely seen.
- Order Lepidoptera
- Insects with complete metamorphosis; the wings covered with flat scales; larval mouthparts mandibulate; adult mouthparts mandibulate to haustellate, sometimes vestigial; wings with venation ranging from primitive (complex) to considerably reduced, especially in the hind wings; pupae exarate to obtect; almost all phytophagous (herbivorous).
- Superfamily Noctuoidea
- Almost 52,000 species in 8 families; adults with a pair of complex tympanic organs on metathorax.
- Family Noctuidae (owlet moths)
- More than 35,000 species worldwide; abundant everywhere with a great diversity of size, color, and habit; sizes range from a wingspan of 10 mm (0.4 inches; Hypenodes) to 275 mm (Thysania); many species brightly colored, but the majority are plain and cryptic; nearly every type of plant material is exploited as food: foliage, flowers, fruits, stems, rootstocks, and woodland litter; a few genera (e.g., Cosmia) cannibalistic; family includes the armyworms and cutworms, among the most injurious of all moth larvae to human interests.
- Family Arctiidae (tiger moths)
- Approximately 10,000 species worldwide; adults usually brightly colored, especially in tropics; aposematic coloration, with protective and toxic secretions; timbal organ for making very high-pitched sounds located on each side of the metathorax; larvae often with much secondary hair; several other families, chiefly tropical, are closely related and also have timbal organs.
- Family Lymantriidae (tussock moths)
- More than 2,500 species worldwide, but mainly in Old World tropics; adult females heavy-bodied, sometimes wingless; many larvae with prominent tussocks and pencils of hair, poison spines, and protrusions that emit repellent odors (osmeteria); some—e.g., the gypsy moth (Lymantria dispar) and the nun moth (L. monacha)—are very destructive to forests; closely related Thaumetopoeidae of the Old World are best known for the gregarious habits of the larvae (processionary caterpillars), which move in orderly columns.
- Family Notodontidae (prominent moths)
- More than 2,800 species worldwide, except New Zealand and Pacific islands; adult coloration mostly dull, cryptic; larvae very diverse with cryptic, disruptive, or aposematic coloration; some protected by toxic secretions; posterior prolegs often modified or aborted; chiefly foliage eaters.
- Superfamily Pyraloidea
- Approximately 17,800 species in 2 families worldwide; most with a pair of tympanal organs on the first abdominal segment; adults usually slender-bodied with long legs; many with narrow forewings and broad, often folded, hind wings.
- Family Pyralidae (pyralid, or snout, moths)
- Approximately 6,130 species of small moths, mostly plain, often abundant, with many important pest species; differentiated from other families by wing venation; small subfamily Nymphulinae has aquatic larvae with tracheal gills for living in still or running fresh water; larvae of subfamily Pyralinae are mostly scavengers, as are those of the Galleriinae, many of which live in bee or wasp nests; larvae of the large subfamily Phycitinae have very diverse habits, including predation on scale insects.
- Family Crambidae (webworms)
- Approximately 11,600 species worldwide; small, often abundant moths, many larvae producing silk webbing in feeding sites; subfamily Crambinae contains almost 1,900 species, larvae feeding mainly on roots, grasses, or mosses on the ground or boring into stems of grasses, sedges, or rushes; subfamily Pyraustinae contains more than 7,400 species, feeding mainly on stems and fruits of various plants; many Pyraustinae species are considered pests, but some have been used in management of aquatic weeds.
- Superfamily Geometroidea
- Almost 22,000 species; adults with abdominal tympana; some authorities classify each of the 3 major families as a separate superfamily.
- Family Geometridae (measuring worm, or inchworm, moths)
- Approximately 21,000 species, abundant worldwide; adults and larvae commonly very cryptic, resembling bark, dead leaves, and twigs; larvae (“inchworms”) long, slender, lacking most posterior abdominal prolegs, crawling with characteristic looping gait; pupation usually in the soil, with no cocoon; many species destructive to foliage—e.g., cankerworms (Alsophila and Paleacrita) and the winter moth (Operophtera brumata).
- Family Uraniidae (swallowtail moths)
- Approximately 700 chiefly tropical species; some adults are large, brilliantly iridescent diurnal moths; the Asian Epicopeia (family Epicopeiidae) mimic swallowtail butterflies.
- Superfamily Drepanoidea
- Approximately 700 species worldwide in 2 families.
- Family Drepanidae (hooktip moths)
- Approximately 650 species worldwide, chiefly Indo-Australian; many of the adults have the forewing apexes strongly hooked; larvae usually lack last pair of prolegs; subfamilies Thyatirinae and Epibleminae sometimes classified as families.
- Family Epicopeiidae (epicopeiid moths)
- 25 species in Arctic and tropical Asia; colorful day-flying moths that often mimic butterflies and other colorful moths such as the Arctiidae; larvae feed on foliage of woody plants.
- Superfamily Gelechioidea
- More than 16,000 species worldwide; adults mostly larger and broader winged than Tineoidea; larvae seldom leaf miners; pupae relatively immobile.
- Family Gelechiidae (twirler moths)
- More than 4,500 species of small to minute moths, worldwide in distribution; larvae diverse, eating leaves, stems, fruit, and tubers, including a few gall makers; serious economic pests include the pink bollworm of cotton and the Angoumois grain moth (Sitotroga cerealella).
- Family Cosmopterigidae (cosmopterigid moths)
- More than 1,600 species of small moths, worldwide in distribution; many adults are very narrow-winged with bright, often metallic markings; in addition to leaf miners, rollers, and tiers, larvae include stem, fruit, and seed borers as well as scavengers.
- Family Coleophoridae (casebearer moths)
- Approximately 1,400 species, mainly Holarctic in distribution; small, very narrow-winged moths; larvae mostly mine leaves or feed on seeds; many larvae construct portable cases with distinctive shapes; some are pests of fruit trees.
- Family Oecophoridae (oecophorid moths)
- More than 3,100 small species worldwide; adults tend to be flat-bodied and somewhat broader-winged than related groups; related families: Elachistidae, Xylorictidae.
- Superfamily Papilionoidea (butterflies)
- 14,000 species, all families worldwide; adults with clubbed but not hooked antennae; flight slower than moths and not darting; larvae lacking “neck”; general evolutionary trend has been from a primitive condition of fully developed forelegs in both sexes (Pieridae, Papilionidae) to one in which they are greatly reduced and useless for walking (Nymphalidae); intermediate conditions occur in Lycaenidae; pupae (“chrysalides”) often brightly colored and irregularly shaped, not enclosed in cocoons.
- Family Lycaenidae (blues, coppers, hairstreaks, and metalmarks)
- 6,000 small and diverse species; many iridescent blue, green, or metallic orange; some adults have thin tails on the trailing margin of the hind wing; larvae somewhat sluglike, hairy, many secreting honeydew and having mutualistic relationships with ants; some prey on the ant brood or on other insects; many pupae have stridulatory structures on the abdomen.
- Family Nymphalidae (brush-footed butterflies)
- Approximately 6,000 species, often split into several families that here are considered subfamilies; Nymphalinae is the main subfamily, with many familiar species such as the fritillaries, admirals, checkerspots, and anglewings; many tropical species brilliantly iridescent; Satyrinae contains the familiar wood nymphs, meadow browns, and heaths, usually with eyespots on the wings; larvae distinctively pointed at the rear; spin crude cocoons; the Libytheinae (snout butterflies) are so named because of their long protruding palps; the very large Brassolinae and iridescent Morphinae are Neotropical, as are the highly distasteful, aposematic Heliconiinae and Ithomiinae that, with the worldwide Danainae, are models in many mimicry complexes; most of the pantropical Acraeinae are also highly protected and aposematic models; some nymphalids, such as the monarch butterfly, are migratory.
- Family Pieridae (white, orange-tip, and sulfur butterflies)
- Approximately 1,000 small to medium-size species; no native species are found in New Zealand; mostly white, yellow, or orange, often with dark tips on wings; pupae usually with a frontal horn on head; many species noted for mass migrations.
- Family Papilionidae (swallowtails and parnassians)
- 600 medium to large species, often brightly colored; many have the “swallowtails” on the hind wings; some females of the Indo-Australian birdwings (Troides) are the largest butterflies; larvae often curiously patterned, with eversible, protective scent organs (osmeteria); adults of many are highly distasteful and much mimicked; parnassians sometimes placed in a separate family, Parnassiidae.
- Superfamily Tortricoidea
- 6,100 species in 1 family; adults with fairly broad, short-fringed wings that seldom span more than 2.5 cm (1 inch); most have cryptic coloration and patterns; larvae mostly leaf folders and rollers, but many bore in fruits, seeds, and soft stems.
- Family Tortricidae (leaf roller moths)
- Approximately 6,100 species worldwide; family large and diverse; most larvae are stem borers or leaf rollers or feed in leaf litter; larvae of the green leaf roller of Europe (Tortrix viridana) defoliate oak forests; the spruce budworm (Choristoneura fumiferana) is the worst forest pest of North America.
- Superfamily Tineoidea
- More than 4,000 species worldwide; a large group of families of mostly small moths of diverse habits; all have some primitive venation features and life cycles; wings narrow to very narrow.
- Family Tineidae (clothes moths and other tineid moths)
- Approximately 3,000 species worldwide; small narrow-winged moths with rough, hairy heads; larvae often casemakers, feeding on debris and fungi; clothes moths (Tineola, Tinea, Trichophaga) often serious household pests; related family: Acrolophidae (burrowing sod webworms).
- Superfamily Gracillarioidea
- Approximately 2,300 species worldwide; small moths; larvae are mainly leaf miners or stem borers
- Families Gracillariidae and Douglasiidae
- Approximately 2,000 species worldwide whose larvae have degenerative legs and mandibles; adults with narrow, long-fringed wings often with metallic markings; larvae mostly leaf miners or stem borers, sometimes greatly flattened.
- Superfamily Hesperioidea
- 3,500 species worldwide in 1 family; similar to true butterflies, distinguished from moths by diurnal habits, clubbed antennae, a functional proboscis, and lack of ocelli; adults are fast-flying, with short, usually pointed forewings, broad heads, and antennae usually hooked beyond the club; larvae usually have a pronounced necklike collar.
- Family Hesperiidae (skippers)
- 3,500 species worldwide; adults range from small and drab to large and iridescent, especially in the tropics; larvae mostly live concealed in individual leaf nests or in webs among grasses, forming flimsy cocoons.
- Superfamily Bombycoidea
- Approximately 3,400 species; adults large to very large; male antennae comblike in form.
- Family Bombycidae (silkworm moths)
- 350 species worldwide except Europe; most common in Asian and New World tropics; includes the domesticated silkworm (Bombyx mori); related family: Eupterotidae.
- Family Saturniidae (giant silkworm moths)
- 1,480 large species worldwide; larvae very large and fleshy, often with brightly colored knobs and spines; most species spin firm cocoons of brown, green, or silvery silk; adults often with bright colors and striking wing shapes and patterns; males with broadly branched antennae, wingspan up to 275 mm (10.8 inches) in hercules moth (Coscinoscera hercules) of Australia and New Guinea, though every continent has some well-known species—e.g., the North American cecropia (Hyalophora cecropia), Io (Automeris io), polyphemus (Antheraea polyphemus), and luna (Actias luna); European giant peacock or emperor (Saturnia pavonia); and Indo-Australian atlas moth (Attacus atlas). Subfamily Citheroniinae (regal moths) are medium to very large New World species of heavy-bodied moths, often brightly colored; larvae often with long spines—e.g., the North American Citheronia (hickory and pine horned devils); pupation in the ground, with no cocoon; related families: Brahmaeidae (African and Indo-Australian), Mimallonidae (New World).
- Family Sphingidae (hawk, or sphinx, moths)
- 1,200 medium-size to large species worldwide; adults powerful fliers with long, narrow forewings, thick antennae, and generally strong proboscises; feed by hovering before flowers and drawing nectar through the extended proboscis; some are diurnal, most are active at twilight (crepuscular); larvae large, fleshy, and mostly smooth; most have a single long posterior horn; typically rest with head and thorax reared, fancifully like the Sphinx at Giza, Egypt, in profile.
- Superfamily Lasiocampoidea
- Approximately 1,600 species worldwide.
- Family Lasiocampidae (tent caterpillar and lappet moths)
- 1,500 species worldwide; larvae usually hairy and brightly colored, some living gregariously in silk nests; adults medium-size, stout-bodied, short-winged, and exceptionally hairy.
- Superfamily Zygaenoidea
- Approximately 2,500 species worldwide; wings broad to very broad with short fringes; some primitive adult, pupal, and larval features; pupae quite mobile.
- Family Limacodidae, or Eucleidae (slug caterpillar moths)
- More than 1,000 species worldwide; larvae broad and flat, with reduced prolegs; move glidingly with head hidden beneath prothorax; many with toxic, irritant setae; adults with heavy hairy bodies and vestigial proboscises.
- Family Megalopygidae (flannel moths)
- 240 species in Central and South America; larvae similar to those of Limacodidae, but with normal prolegs and traces of additional ones; setae very toxic and nettling.
- Family Zygaenidae (burnet and forester moths)
- More than 1,000 species, mainly in subtropical and tropical Asia and Palearctic regions; adults usually diurnal flower visitors, with bright colors and strong proboscises; strongly protected by high concentrations of hydrogen cyanide in blood; larvae are leaf skeletonizers; related families: Aididae and Chalcosiidae (Old World tropics); Pyromorphidae and Dalceridae (New World).
- Family Epipyropidae (parasitic moths)
- 40 chiefly Asian species; larvae live as external parasites on plant hoppers; related family: Cyclotornidae (Australian; larvae live similarly when young, then move to ants’ nests).
- Superfamily Yponomeutoidea
- More than 1,500 species worldwide; a limited and not very distinctive superfamily; larvae possess distinctive primary setation.
- Family Yponomeutidae (ermine moths)
- Almost 600 species worldwide; adults brightly colored, especially in the tropics; pupae of some make lacework cocoons; related families: Plutellidae, Glyphipterygidae, Heliodinidae, Scythrididae.
- Superfamily Sesioidea
- Approximately 1,200 species worldwide; most sesioid moths are diurnal with many aposematic adults.
- Family Sesiidae (clearwing, or wasp, moths)
- More than 1,000 species worldwide; adults diurnal flower visitors; often brightly colored with yellow, orange, or scarlet, the wings usually mostly transparent, often very striking mimics of wasps; larvae often are stem, twig, and root borers, often injurious to fruit trees.
- Family Castniidae (castniid moths)
- Approximately 130 species in Central and South America; medium-size to large diurnal species of the New World and Indo-Australian tropics; adults powerful fliers, heavy-bodied and broad-winged; clubbed antennae, bright colors; often mimic other butterflies and diurnal moths; larvae are often stem borers.
- Superfamily Cossoidea
- Approximately 700 species described; adults range from small to large, usually robust moths; males often with bipectinate antennae; larvae mainly stem or wood borers.
- Family Cossidae (carpenterworm and goat moths)
- Almost 700 species described worldwide; medium-size to large moths; adults are the heaviest adult lepidopterans, with wingspans up to 25 cm (10 inches); larvae are wood borers, penetrating even solid heartwoods; may require at least 2 years to mature; some very destructive, including the coffee borer (Zeuzera coffeae) and leopard moth (Z. pyrina).
- Superfamily Pterophoroidea
- Almost 1,000 species in 1 family.
- Family Pterophoridae (plume moths)
- Almost 1,000 mainly tropical species; adults with very long, slender legs and bodies, the wings usually deeply cleft into plumes; larvae spin webs on and eat the leaves of various plants or bore into seeds, seedpods, roots, or stems.
- Superfamily Alucitoidea
- Almost 150 species worldwide; this superfamily and the related Pterophoroidea are the only families with deeply lobed wings.
- Family Alucitidae (many-plumed moths)
- 130 species worldwide; each wing is very deeply cleft into 6 or more narrow plumelike divisions.
- Superfamily Nepticuloidea
- Approximately 900 species worldwide; females with one genital opening and a soft ovipositor.
- Family Nepticulidae (midget moths)
- 800 species worldwide of very small to minute moths; antennae with broad “eyecaps” at the base; larvae mostly leaf and bark miners, a few gall makers.
- Family Opostegidae
- Approximately 100 worldwide species of small moths with narrow long-fringed wings; larvae leaf, stem, or bark miners.
- Superfamily Tischerioidea
- Approximately 80 species in a single family.
- Family Tischeriidae (trumpet leaf miner moths)
- Approximately 80 species predominantly in North America; not found in Australia or the rest of Oceania.
- Superfamily Incurvarioidea
- More than 500 species; all females with an extensible, piercing ovipositor for inserting eggs into plant tissue.
- Family Incurvariidae (fairy, or leafcutter, moths)
- Approximately 100 species worldwide; many are small brilliantly colored diurnal flower visitors; male antennae often several times as long as forewings; mutualistic relationships of the yucca moths (Prodoxinae) with their food plants are notable as an example of coevolution; family sometimes split into families Incurvariidae, Adelidae, and Prodoxidae; related family: Heliozelidae (shield bearers).
- Superfamily Hepialoidea
- 520 species; females with two genital openings; adult mouthparts reduced and nonfunctional, antennae very short.
- Family Hepialidae (swifts, or ghost moths)
- Almost 500 species found worldwide but chiefly in Australia and New Zealand; medium-size to very large moths, some brilliantly colored; larvae mostly bore in turf or wood; related families, less-known: Prototheoridae (Africa and Australia), Palaeosetidae (Australia), Anomosetidae (Australia).
- Superfamily Micropterigoidea
- The most primitive lepidopterans; females with no special genital opening; larvae, pupae, and adults with mandibulate mouthparts.
- Family Micropterigidae (mandibulate moths)
- 120 small species, a few found in the Northern Hemisphere, more in Australia and New Zealand; adults eat pollen; larvae eat mosses and liverworts.
- Superfamily Eriocranioidea
- 24 species in 1 family; females with one genital opening, a long cloaca, and a piercing ovipositor; adults with a short proboscis; pupae with functional mandibles.
- Family Eriocraniidae (sparkling archaic sun moths)
- 24 species with a Holarctic distribution; often brilliantly colored; adults feed on nectar; related families: Mnesarchiidae (New Zealand), Mesopseustidae (India and Taiwan).
