Lepidoptera

Lepidoptera

Overview
Lepidoptera is a large order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 of insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s that includes moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...

s and butterflies
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...

 (called lepidopterans). It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...

, skipper butterflies
Skipper (butterfly)
A skipper or skipper butterfly is a butterfly of the family Hesperiidae. They are named after their quick, darting flight habits. There are more than 3500 recognized species of skippers and they occur worldwide, but with the greatest diversity in the Neotropical regions of Central and South...

, and moth-butterflies
Hedylidae
Hedylidae, the "American moth-butterflies", is a family of insects in the lepidopteran order, representing the superfamily Hedyloidea. They are an extant sister group of the butterfly superfamilies Papilionoidea and Hesperioidea...

. The term was coined by Linnaeus in 1735 and is derived from Ancient Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

  (scale) and (wing). Comprising an estimated 174,250 species, in 126 families
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 and 46 superfamilies
Taxonomic rank
In biological classification, rank is the level in a taxonomic hierarchy. Examples of taxonomic ranks are species, genus, family, and class. Each rank subsumes under it a number of less general categories...

, the Lepidoptera show many variations of the basic body structure that have evolved to gain advantages in lifestyle and distribution.
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Encyclopedia
Lepidoptera is a large order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 of insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s that includes moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...

s and butterflies
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...

 (called lepidopterans). It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...

, skipper butterflies
Skipper (butterfly)
A skipper or skipper butterfly is a butterfly of the family Hesperiidae. They are named after their quick, darting flight habits. There are more than 3500 recognized species of skippers and they occur worldwide, but with the greatest diversity in the Neotropical regions of Central and South...

, and moth-butterflies
Hedylidae
Hedylidae, the "American moth-butterflies", is a family of insects in the lepidopteran order, representing the superfamily Hedyloidea. They are an extant sister group of the butterfly superfamilies Papilionoidea and Hesperioidea...

. The term was coined by Linnaeus in 1735 and is derived from Ancient Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

  (scale) and (wing). Comprising an estimated 174,250 species, in 126 families
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 and 46 superfamilies
Taxonomic rank
In biological classification, rank is the level in a taxonomic hierarchy. Examples of taxonomic ranks are species, genus, family, and class. Each rank subsumes under it a number of less general categories...

, the Lepidoptera show many variations of the basic body structure that have evolved to gain advantages in lifestyle and distribution. Recent estimates suggest that the order may have more species than earlier thought, and is among the four most speciose orders, along with the Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

, Diptera
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...

, and the Coleoptera
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...

.

Lepidopteran species are characterized by more than 3 derived features, some of the most apparent being the scales covering their bodies and wings, and a proboscis
Proboscis
A proboscis is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In simpler terms, a proboscis is the straw-like mouth found in several varieties of species.-Etymology:...

. The scales are modified, flattened "hairs", and give butterflies and moths their extraordinary variety of colors and patterns. Almost all species have some form of membranous wings, except for a few that have reduced wings or are wingless. Like most other insects, butterflies and moths are holometabolous, meaning they undergo complete metamorphosis
Holometabolism
Holometabolism, also called complete metamorphism, is a term applied to insect groups to describe the specific kind of insect development which includes four life stages - as an embryo or egg, a larva, a pupa and an imago or adult. Holometabolism is a monophyletic trait that all insects in the...

. Mating and the laying of eggs are carried out by adults, normally near or on host plants for the larvae. The larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

e are commonly called caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

s, and are completely different from their adult moth or butterfly form, having a cylindrical body with a well-developed head, mandible mouth parts, and from 0 to 11 (usually 8) pairs of proleg
Proleg
A Proleg is the small fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the order Lepidoptera, though they can also be found on other larval insects such as sawflies and a few types of flies....

s. As they grow, these larvae will change in appearance, going through a series of stages called instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...

s. Once fully matured, the larva develops into a pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

, referred to as a chrysalis in the case of butterflies. A few butterflies and many moth species spin a silk case or cocoon prior to pupating, while others do not, instead going underground.

The Lepidoptera have, over millions of years, evolved a wide range of wing patterns and coloration ranging from drab moths akin to the related order Trichoptera
Trichoptera
The caddisflies are an order, Trichoptera, of insects with approximately 12,000 described species. Also called sedge-flies or rail-flies, they are small moth-like insects having two pairs of hairy membranous wings...

, to the brightly colored and complex-patterned butterflies. Accordingly, this is the most recognized and popular of insect orders with many people involved in the observation, study, collection, rearing of and commerce in these insects. A person who collects or studies this order is referred to as a lepidopterist
Lepidopterist
A lepidopterist is a person who specialises in the study of Lepidoptera, members of an order encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...

.

Butterflies and moths play an important role in the natural ecosystem as pollinator
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...

s and as food in the food chain; conversely, their larva are considered very problematic to vegetation in agriculture, as their main source of food is often live plant matter. In many species, the female may produce anywhere from 200 to 600 eggs, while in others the number may go as high as 30,000 eggs in one day. The caterpillars hatching from these eggs can cause damage to large quantities of crops. Many moth and butterfly species are of economic interest by virtue of their role as pollinators, the silk
Silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...

 they produce or as pest species.

Etymology


The word Lepidoptera comes from the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 word for "scaly wing", from the Ancient Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

  () meaning scale and () meaning wing. Sometimes the term Rhopalocera is used to group the species that are butterflies
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...

, derived from the Ancient Greek () and () meaning club and horn respectively; coming from the shape of the antenna
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....

e of butterflies.

The origins of the common names "butterfly"and "moth" are varied and often obscure. The English
English language
English is a West Germanic language that arose in the Anglo-Saxon kingdoms of England and spread into what was to become south-east Scotland under the influence of the Anglian medieval kingdom of Northumbria...

 word butterfly is from Old English , with many variations in spelling. Other than that, the origin is unknown, although it could be derived from the pale yellow color of many species' wings suggesting the color of butter or margarine. The species of Heterocera are commonly called moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...

s. The origins of the English word moth are more clear, deriving from the Old English " (cf. Northumbrian dialect ) from Common Germanic (compare Old Norse
Old Norse
Old Norse is a North Germanic language that was spoken by inhabitants of Scandinavia and inhabitants of their overseas settlements during the Viking Age, until about 1300....

 , Dutch
Dutch language
Dutch is a West Germanic language and the native language of the majority of the population of the Netherlands, Belgium, and Suriname, the three member states of the Dutch Language Union. Most speakers live in the European Union, where it is a first language for about 23 million and a second...

  and German
German language
German is a West Germanic language, related to and classified alongside English and Dutch. With an estimated 90 – 98 million native speakers, German is one of the world's major languages and is the most widely-spoken first language in the European Union....

  all meaning "moth"). Perhaps its origins are related to Old English meaning "maggot
Maggot
In everyday speech the word maggot means the larva of a fly ; it is applied in particular to the larvae of Brachyceran flies, such as houseflies, cheese flies, and blowflies, rather than larvae of the Nematocera, such as mosquitoes and Crane flies...

" or from the root of "midge
Midge
A midge is a very small, two-winged flying insect. "Midge" may also refer to:-Real:* Midge Costanza , American politician* Mildred Gillars , aka "Midge", American broadcaster of Nazi propaganda during World War II...

", which until the 16th century was used mostly to indicate the larva, usually in reference to devouring clothes.

The etymological origins of the word caterpillar, the larval form of butterflies and moths, are from the early 16th century, from Middle English
Middle English
Middle English is the stage in the history of the English language during the High and Late Middle Ages, or roughly during the four centuries between the late 11th and the late 15th century....

 catirpel, catirpeller, probably an alteration of Old North French
Old French
Old French was the Romance dialect continuum spoken in territories that span roughly the northern half of modern France and parts of modern Belgium and Switzerland from the 9th century to the 14th century...

 catepelose: cate, cat (from Latin cattus) + pelose, hairy (from Latin pilōsus).

Distribution and diversity


Lepidoptera are among the most successful groups of insects. They are found on all continents, except Antarctica. Lepidoptera inhabit all terrestrial habitats ranging from desert to rainforest, from lowland grasslands to montane plateaus but almost always associated with higher plants, especially angiosperms (flowering plant
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...

s). Among the most northern dwelling species of butterflies and moths is the Arctic Apollo (Parnassius arcticus
Parnassius arcticus
Siberian Apollo Parnassius arcticus is a high altitude butterfly which is found in NE. Yakutia. It is a member of the Snow Apollo genus Parnassius of the Swallowtail family....

), which is found in the Arctic Circle in northeastern Yakutia, at an altitude of 1500 meters above sea level. In the Himalayas
Himalayas
The Himalaya Range or Himalaya Mountains Sanskrit: Devanagari: हिमालय, literally "abode of snow"), usually called the Himalayas or Himalaya for short, is a mountain range in Asia, separating the Indian subcontinent from the Tibetan Plateau...

, various Apollo species such as Parnassius epaphus
Parnassius epaphus
Common Red Apollo Parnassius epaphus is a high altitude butterfly which is found in India. It is a member of the Snow Apollo genus Parnassius of the Swallowtail family...

, besides others, have been recorded to occur up to an altitude of 6,000 meters above sea level.

Some lepidopteran species exhibit symbiotic
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

, phoretic
Commensalism
In ecology, commensalism is a class of relationship between two organisms where one organism benefits but the other is neutral...

, or parasitic
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...

 life-styles inhabiting the bodies of organisms rather than the environment. Coprophagous
Coprophagia
Coprophagia or coprophagy is the consumption of feces, from the Greek κόπρος copros and φαγεῖν phagein . Many animal species practice coprophagia as a matter of course; other species do not normally consume feces but may do so under unusual conditions...

 pyralid
Pyralidae
The Pyralidae or snout moths are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many classifications, the grass moths are included in the Pyralidae as a subfamily, making the combined group one of the largest families in the Lepidoptera...

 moth species, called as sloth moth
Sloth moth
Sloth moth is a generic term used to refer to coprophagous moths which have evolved to exclusively inhabit the fur of sloths and to use sloth dung as a substrate for the early stages of reproduction....

s, such as Bradipodicola hahneli
Bradipodicola hahneli
Bradypodicola hahneli is a sloth moth in the Pyralidae family that lives exclusively in the fur of the Pale-throated Three-toed Sloth , a three-toed sloth found in South America. It is the only species of the Bradypodicola genus.The three-toed sloth’s fur forms a micro-ecozone inhabited by green...

and Cryptoses choloepi
Cryptoses choloepi
Cryptoses choloepi is a sloth moth in the snout moth family that lives exclusively in the fur of sloths, mammals found in South and Central America....

, are unusual in that they are exclusively found inhabiting the fur of the sloth
Sloth
Sloths are the six species of medium-sized mammals belonging to the families Megalonychidae and Bradypodidae , part of the order Pilosa and therefore related to armadillos and anteaters, which sport a similar set of specialized claws.They are arboreal residents of the jungles of Central and South...

s, mammals found in Central
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...

 and South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

.
Two species of Tinea
Tinea (moth)
Tinea is a genus of the fungus moth family, Tineidae. Therein, it belongs to the subfamily Tineinae. As evident by its name, it is the type genus of its subfamily and family...

moths have been recorded as feeding on horny tissue and have been bred from the horns of cattle. The larva of Zenodochium
Zenodochium
Zenodochium is a genus of moth in the family Blastobasidae....

 coccivorella
is an internal parasite of the coccid
Scale insect
The scale insects are small insects of the order Hemiptera, generally classified as the superfamily Coccoidea. There are about 8,000 species of scale insects.-Ecology:...

 Kermes
Kermes (insect)
Kermes is a genus of scale insects in the order Hemiptera. They feed on the sap of evergreen oaks; the females produce a red dye, also called "kermes", that is the source of natural crimson...

species. Many species have been recorded as breeding in natural materials or refuse such as owl pellets, bat caves, honey-combs or diseased fruit.

Of the approximately 174,250 lepidopteran species described until 2007, it is estimated that butterflies and skippers comprise approximately 17,950 with moths making up the rest. The vast majority of Lepidoptera are to be found in the tropics but substantial diversity exists on most continents. North America has over 700 species of butterflies and over 11,000 species of moths, while there are about 400 species of butterflies and 20,000 species of moths reported from Australia. The diversity of Lepidoptera in each faunal region has been estimated by John Heppner in 1991 based partly on actual counts from the literature, partly on the card indexes in the Natural History Museum
Natural History Museum
The Natural History Museum is one of three large museums on Exhibition Road, South Kensington, London, England . Its main frontage is on Cromwell Road...

 (London) and the National Museum of Natural History
National Museum of Natural History
The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. Admission is free and the museum is open 364 days a year....

 (Washington), and partly on estimates:
Diversity of Lepidoptera in each faunal region
Palearctic Nearctic Neotropic Afrotropic Indo-Australian
(comprising Indomalayan and Australian regions)
Estimated number of species 22,465 11,532 44,791 20,491 47,286

External morphology



Lepidoptera are morphologically distinguished from other orders principally by the presence of scale
Scale (Lepidoptera)
The presence of scales on the wings of Lepidoptera, comprising moths and butterflies, characterises this order of insects. The name is derived from Ancient Greek λεπίδος and πτερόν . The wings of Lepidoptera are minutely scaled, which feature gives the name to this order...

s on the external parts of the body and appendages, especially the wing
Wing
A wing is an appendage with a surface that produces lift for flight or propulsion through the atmosphere, or through another gaseous or liquid fluid...

s. Butterflies and moths vary in size from microlepidoptera
Microlepidoptera
Microlepidoptera is an artificial grouping of moth families, commonly known as the 'smaller moths' . These generally have a wingspan of under 20 mm, and are thus harder to identify by external phenotypic markings than macrolepidoptera...

 only a few millimeters long, to conspicuous animals with a wingspan of many inches, such as the Monarch butterfly and Atlas moth.
Lepidopterans undergo a four-stage life cycle
Biological life cycle
A life cycle is a period involving all different generations of a species succeeding each other through means of reproduction, whether through asexual reproduction or sexual reproduction...

: egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...

; larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

 or caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

; pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

 or chrysalis; and imago (plural: imagines)
Imago
In biology, the imago is the last stage of development of an insect, after the last ecdysis of an incomplete metamorphosis, or after emergence from the pupa where the metamorphosis is complete...

 / adult and show many variations of the basic body structure, which have evolved to gain advantages in lifestyle and distribution.

Head



The head is where many sensing organs and the mouth parts are found. Like the adult, the larva also have a toughened, or sclerotized
Sclerite
A sclerite is a hardened body part. The term is used in various branches of biology for various structures including hardened portions of sponges, but it is most commonly used for the hardened portions of arthropod exoskeletons....

 head capsule. Here, there are two compound eyes, and chaetosema, raised spots or clusters of sensory bristles unique to Lepidoptera, even though many taxa have lost one or both of these spots. The antennae have a wide variation in form among species and even between different sexes. The antennae of butterflies are usually filiform and shaped like clubs, those of the skippers are hooked while those of moths have flagellar segments variously enlarged or branched. Some moths have antennae that are enlarged, or tapered and hooked at the ends.

The maxillary galeae are modified and form an elongated proboscis
Proboscis
A proboscis is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In simpler terms, a proboscis is the straw-like mouth found in several varieties of species.-Etymology:...

. The proboscis consists of one to five segments, usually kept coiled up under the head by small muscles when it is not being used to suck up nectar from flowers or other liquids. Some basal
Basal (phylogenetics)
In phylogenetics, a basal clade is the earliest clade to branch in a larger clade; it appears at the base of a cladogram.A basal group forms an outgroup to the rest of the clade, such as in the following example:...

 moths still have mandibles, or separate moving jaws, like their ancestors and these form the family Micropterigidae
Micropterigidae
Micropterigoidea is the superfamily of "mandibulate archaic moths", all placed in the single family Micropterigidae, containing currently about 20 living genera. They are considered the most primitive extant lineage of Lepidoptera ....

.

The larvae, called caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

s, have a toughened head capsule. Caterpillars lack the proboscis and have separate chewing mouthparts
Insect mouthparts
Insects exhibit a range of mouthparts, adapted to particular modes of feeding. The earliest insects had chewing mouthparts...

. These mouthparts, called mandibles, are used to chew up the plant matter that the larvae eat. The lower jaw, or labium, is weak but may carry a spinneret, an organ used to create silk. The head is made of large lateral lobes, each having an ellipse of up to six simple eyes.

Thorax


The thorax is made of three fused segments, the prothorax
Prothorax
The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites are the pronotum , the prosternum , and the propleuron on each side. The prothorax never bears wings in extant insects, though some fossil groups possessed...

, mesothorax
Mesothorax
The mesothorax is the middle of the three segments in the thorax of an insect, and bears the second pair of legs. Its principal sclerites are the mesonotum , the mesosternum , and the mesopleuron on each side...

, and metathorax
Metathorax
The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites are the metanotum , the metasternum , and the metapleuron on each side...

, each with a pair of legs. The first segment contains the first pair of legs. In some males of the butterfly family Nymphalidae
Nymphalidae
The Nymphalidae is a family of about 5,000 species of butterflies which are distributed throughout most of the world. These are usually medium sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called...

, the fore-legs are greatly reduced and are not used for walking or perching. The three pairs of legs are covered with scales. Lepidoptera also have olfactory organs on their feet, which aid the butterfly in "tasting" or "smelling" out its food. In the larval form there are 3 pairs of true legs, with 0–11 pairs of abdominal legs (usually 8) and hooklets, called apical crochets.

The two pairs of wings are found on the middle and third segment, or mesothorax
Mesothorax
The mesothorax is the middle of the three segments in the thorax of an insect, and bears the second pair of legs. Its principal sclerites are the mesonotum , the mesosternum , and the mesopleuron on each side...

 and metathorax
Metathorax
The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites are the metanotum , the metasternum , and the metapleuron on each side...

 respectively. In the more recent genera, the wings of the second segment are much more pronounced, although some more primitive forms have similarly sized wings of both segments. The wings are covered in scales arranged like shingles, which form an extraordinary variety of colors and patterns. The mesothorax is designed to have more powerful muscles to propel the moth or butterfly through the air, with the wing of this segment (forewing) having a stronger vein structure. The largest superfamily, Noctuidae
Noctuidae
The Noctuidae or owlet moths are a family of robustly-built moths that includes more than 35,000 known species out of possibly 100,000 total, in more than 4,200 genera. They constitute the largest family in the Lepidoptera....

, have the wings modified to act as Tympanal or hearing organ
Tympanal organ
A tympanal organ is a hearing organ in insects, consisting of a membrane stretched across a frame backed by an air sac and associated sensory neurons...

s

The caterpillar has an elongated soft body that may have hair-like or other projections, 3 pairs of true legs, with 0–11 pairs of abdominal legs (usually 8) and hooklets, called apical crochets. The thorax will usually have a pair of legs on each segment. The thorax is also lined with many spiracles on both the mesothorax and metathorax, except for a few aquatic species, who instead have a form of gill
Gill
A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist...

.

Abdomen



The abdomen, which is less sclerotized than the thorax, consists of 10 segments with membranes in between allowing for articulated movement. The sternum, on the first segment, is small in some families and is completely absent in others. The last 2 or 3 segments form the external parts of the species' sex organs. The genitalia
Lepidoptera genitalia
The study of the genitalia of Lepidoptera is important for Lepidoptera taxonomy in addition to development, anatomy and natural history. The genitalia are complex and provide the basis for species discrimination in most families and also in family identification. The genitalia are attached onto the...

 of Lepidoptera are highly varied and are often the only means of differentiating between species. Male genitals include a valva, which is usually large, as it is used to grasp the female during mating. Female genitalia include three distinct sections.

In the females of basal moths, there is only one sex organ, which is used for copulation and as a ovipositor
Ovipositor
The ovipositor is an organ used by some animals for oviposition, i.e., the laying of eggs. It consists of a maximum of three pairs of appendages formed to transmit the egg, to prepare a place for it, and to place it properly...

, or egg laying organ. 98% of moth species have a separate organ for mating, and an external duct that carries the sperm from the male.

The abdomen of the caterpillar has 4 pairs of prolegs, normally located on the third to sixth segments of the abdomen, and a separate pair of prolegs by the anus, which have a pair of tiny hooks called crotchets. These aid in gripping and walking, especially in species that lack many prolegs (e. g. larvae of Geometridae). In some basal moths, these prolegs may be on every segment of the body, while prolegs may be lost completely in other groups, which are more adapted to boring and living in sand (e. g., Prodoxidae
Prodoxidae
Prodoxidae is a family of primitive monotrysian Lepidoptera. Some of these small-to-medium sized moths are day flying, like Lampronia capitella, known to European gardeners as the "Currant Shoot Borer". Others occur in Africa and Asia.Tetragma gei feeds on Mountain Avens Geum triflorum in USA....

 and Nepticulidae
Nepticulidae
Nepticulidae is a family of very small moths with a worldwide distribution. They are characterised by eyecaps over the eyes . These pigmy moths or midget moths, as they are commonly known, include the smallest of all living moths, with a wingspan that can be as little as 3 mm...

 respectively).

Scales


The wings, head parts of thorax and abdomen of Lepidoptera are covered with minute scales, from which feature the order 'Lepidoptera' derives its names, the word "lepteron" in Ancient Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

 meaning 'scale'. Most scales are lamellar, or blade-like and attached with a pedicel, while other forms may be hair-like or specialized as secondary sexual characteristics. The lumen or surface of the lamella, has a complex structure. It gives color either due to the pigmentary colors contained within or due to its three-dimensional structure
Photonic crystal
Photonic crystals are periodic optical nanostructures that are designed to affect the motion of photons in a similar way that periodicity of a semiconductor crystal affects the motion of electrons...

. Scales provide a number of functions, which include insulation, thermoregulation, aiding gliding flight, among others, the most important of which is the large diversity of vivid or indistinct patterns they provide, which help the organism protect itself by camouflage
Camouflage
Camouflage is a method of concealment that allows an otherwise visible animal, military vehicle, or other object to remain unnoticed, by blending with its environment. Examples include a leopard's spotted coat, the battledress of a modern soldier and a leaf-mimic butterfly...

, mimicry, and to seek mates.

Internal morphology


In the reproductive system
Reproductive system
The reproductive system or genital system is a system of organs within an organism which work together for the purpose of reproduction. Many non-living substances such as fluids, hormones, and pheromones are also important accessories to the reproductive system. Unlike most organ systems, the sexes...

 of butterflies and moths, the male genitalia
Sex organ
A sex organ, or primary sexual characteristic, as narrowly defined, is any of the anatomical parts of the body which are involved in sexual reproduction and constitute the reproductive system in a complex organism; flowers are the reproductive organs of flowering plants, cones are the reproductive...

 are complex and unclear. In females there are three types of genitalia based on the relating taxa: monotrysian, exoporian, and ditrysian. In the monotrysian type there is an opening on the fused segments of the sterna 9 and 10, which act as insemination and oviposition. In the exoporian type (in Hepaloidae and Mnesarchaeoidea) there are two separate places for insemination and oviposition, both occurring on the same sterna as the monotrysian type, i.e. 9 and 10. The ditrysian groups have an internal duct that carry sperm, with separate openings for copulation and egg-laying In most species the genitalia are flanked by two soft lobes, although they may be specialized and sclerotized in some species for ovipositing in area such as crevices and inside plant tissue. Hormones and the glands that produce them run the development of butterflies and moths as they go through their life cycle, called the endocrine system
Endocrine system
In physiology, the endocrine system is a system of glands, each of which secretes a type of hormone directly into the bloodstream to regulate the body. The endocrine system is in contrast to the exocrine system, which secretes its chemicals using ducts. It derives from the Greek words "endo"...

. The first insect hormone PTTH
Prothoracicotropic hormone
Prothoracicotropic hormone was the first insect hormone that was discovered. It was originally described simply as "brain hormone" by early workers such as Stefan Kopeć and Vincent Wigglesworth , who realized that ligation of the head of immature insects could prevent molting or pupation of the...

 (Prothoracicotropic hormone) operates the species life cycle and diapause (see the relates section). This hormone is produced by corpora allata
Corpus allatum
The corpus allatum , in insect physiology, is an endocrine gland which generates juvenile hormone; as such, it plays a crucial role in metamorphosis. Surgical removal of the corpora allata can cause an immature larva to pupate at its next molt, resulting in a miniature adult...

 and corpora cardiaca, where it is also stored. Some glands are specialized to perform certain task such as producing silk or producing saliva in the palpi. While the corpora cardiaca produce PTTH, the corpora allata also produces jeuvanile hormones, and the prothorocic glands produce moulting hormones.

In the digestive system, the anterior region of the foregut has been modified to form a pharyngeal sucking pump as they need it for the food they eat, which are for the most part liquids. An esophagus
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...

 follows and leads to the posterior of the pharynx and in some species forms a form of crop. The midgut is short and straight, with the hindgut being longer and coiled. Ancestors of lepidopteran species, stemming from Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

, had midgut ceca, although this is lost in current butterflies and moths. Instead, all the digestive enzymes other than initial digestion, are immobilized at the surface of the midgut cells. In larvae, long-necked and stalked goblet cell
Goblet cell
Goblet cells are glandular simple columnar epithelial cells whose sole function is to secrete mucin, which dissolves in water to form mucus. They use both apocrine and merocrine methods for secretion....

s are found in the anterior and posterior midgut regions, respectively. In insects, the goblet cells excrete positive potassium
Potassium
Potassium is the chemical element with the symbol K and atomic number 19. Elemental potassium is a soft silvery-white alkali metal that oxidizes rapidly in air and is very reactive with water, generating sufficient heat to ignite the hydrogen emitted in the reaction.Potassium and sodium are...

 ions, which are absorbed from leaves ingested by the larvae. Most butterflies and moths display the usual digestive cycle, however species that have a different diet require adaptations to meet these new demands.
In the circulatory system
Circulatory system
The circulatory system is an organ system that passes nutrients , gases, hormones, blood cells, etc...

, hemolymph
Hemolymph
Hemolymph, or haemolymph, is a fluid in the circulatory system of some arthropods and is analogous to the fluids and cells making up both blood and interstitial fluid in vertebrates such as birds and mammals...

, or insect blood, is used to circulate heat in a form of thermoregulation
Thermoregulation
Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different...

, where muscles contraction produces heat, which is transferred to the rest of the body when conditions are unfavorable. In lepidopteran species, hemolymph is circulated through the veins in the wings by some form of pulsating organ, either by the heart or by the intake of air into the trachea
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....

. Air is taken in through spiracles along the sides of the abdomen and thorax supplying the trachea with oxygen as it goes through the lepidopteran's respiratory system
Respiratory system
The respiratory system is the anatomical system of an organism that introduces respiratory gases to the interior and performs gas exchange. In humans and other mammals, the anatomical features of the respiratory system include airways, lungs, and the respiratory muscles...

. There are three different tracheae supplying oxygen diffusing oxygen throughout the species body: The dorsal, ventral, and visceral. The dorsal tracheae supply oxygen to the dorsal musculature and vessels, while the ventral tracheae supply the ventral musculature and nerve cord, and the visceral tracheae supply the guts, fat bodies, and gonads.

Polymorphism


Polymorphism is the appearance of forms or "morphs", which differ in color and number of attributes within a single species. In Lepidoptera, polymorphism can be seen not only between individuals in a population, but also between the sexes as sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...

, between geographically separated populations in geographical polymorphism and also between generations flying at different seasons of the year (seasonal polymorphism or polyphenism
Polyphenism
A polyphenic trait is a trait for which multiple, discrete phenotypes can arise from a single genotype as a result of differing environmental conditions.-Definition:A polyphenism is a biological mechanism that causes a trait to be polyphenic...

). In some species, the polymorphism is limited to one sex, typically the female. This often includes the phenomenon of mimicry when mimetic morphs fly alongside non-mimetic morphs in a population of a particular species. Polymorphism occurs both at specific level with heritable variation in the overall morphological design of individuals as well as in certain specific morphological or physiological traits within a species.

Environmental polymorphism, in which trait
Trait (biology)
A trait is a distinct variant of a phenotypic character of an organism that may be inherited, environmentally determined or be a combination of the two...

s are not inherited, is often termed as polyphenism
Polyphenism
A polyphenic trait is a trait for which multiple, discrete phenotypes can arise from a single genotype as a result of differing environmental conditions.-Definition:A polyphenism is a biological mechanism that causes a trait to be polyphenic...

. Polyphenism in Lepidoptera is commonly seen in the form of seasonal morphs especially in the butterfly families of Nymphalidae
Nymphalidae
The Nymphalidae is a family of about 5,000 species of butterflies which are distributed throughout most of the world. These are usually medium sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called...

 and Pieridae
Pieridae
The Pieridae are a large family of butterflies with about 76 genera containing approximately 1,100 species, mostly from tropical Africa and Asia. Most pierid butterflies are white, yellow or orange in coloration, often with black spots...

. An Old World pierid butterfly, the Common Grass Yellow (Eurema hecabe
Eurema hecabe
The Large Grass Yellow or Common Grass Yellow is a small pierid butterfly species found in Asia or Africa. They are found flying close to the ground and are found in open grass and scrub habitats...

) has a darker summer adult morph, triggered by a long day exceeding 13 hours in duration, while the shorter diurnal period of 12 hours or less induces a paler morph in the post-monsoon period. Polyphenism also occurs in caterpillars, an example being the Peppered Moth
Peppered moth
The peppered moth is a temperate species of night-flying moth. Peppered moth evolution is often used by educators as an example of natural selection.- Distribution :...

, Biston betularia.

Geographical polymorphism is where geographical isolation causes a divergence of a species into different morphs. A good example is the Indian White Admiral Limenitis procris
Limenitis procris
The Commander , sometimes included in the genus Limenitis), is a medium-sized, strikingly coloured brush-footed butterfly found in Asia. It is notable for the mode of concealment employed by its caterpillar and the cryptic camouflage of its pupa.-Description:The Commander has a wingspan of about 6...

, which has five forms, each geographically separated from the other by large mountain ranges. An even more dramatic showcase of geographical polymorphism is the Apollo butterfly
Apollo (butterfly)
The Apollo or Mountain Apollo , is a butterfly of the Papilionidae family.-Distribution and status:It is found on mountains in Europe usually above up to , preferring flowery meadows and mountain pastures...

 (Parnassius apollo). Due to the Apollos living in small local populations, having no contact with each other, but because of the strong stenotopic species and weak migration ability interbreeding between populations of one species practically does not occur; they form over 600 different morphs, with the size of spots on the wings of which varies greatly.



Sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...

 is the occurrence of differences between males and females in a species. In Lepidoptera, sexual dimorphism is widespread and almost completely determined by genetic determination. Sexual dimorphism is present in all families of the Papilionoidoea and more prominent in the Lycaenidae
Lycaenidae
The Lycaenidae are the second-largest family of butterflies, with about 6000 species worldwide, whose members are also called gossamer-winged butterflies...

, Pieridae
Pieridae
The Pieridae are a large family of butterflies with about 76 genera containing approximately 1,100 species, mostly from tropical Africa and Asia. Most pierid butterflies are white, yellow or orange in coloration, often with black spots...

 and certain taxa of the Nymphalidae
Nymphalidae
The Nymphalidae is a family of about 5,000 species of butterflies which are distributed throughout most of the world. These are usually medium sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called...

. Apart from color variation, which may differ from slight to completely different color-pattern combinations, secondary sexual characteristics may also be present. Different genotypes maintained by natural selection may also be expressed at the same time. Polymorphic and/or mimetic females occur in the case of some taxa in the Papilionidae primarily to obtain a level of protection not available to the male of their species. The most distinct case of sexual dimorphism is that of adult females of many Psychidae species who have only vestigial wings, legs, and mouthparts as compared to the adult males who are strong fliers with well-developed wings and feathery antennae.

Reproduction and development


Species of Lepidoptera undergo holometabolism
Holometabolism
Holometabolism, also called complete metamorphism, is a term applied to insect groups to describe the specific kind of insect development which includes four life stages - as an embryo or egg, a larva, a pupa and an imago or adult. Holometabolism is a monophyletic trait that all insects in the...

 or "complete metamorphosis". Their life cycle normally consists of an egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...

, larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

, pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

, and an imago
Imago
In biology, the imago is the last stage of development of an insect, after the last ecdysis of an incomplete metamorphosis, or after emergence from the pupa where the metamorphosis is complete...

 or adult. The larvae are commonly called caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

s, and the pupae of moths that are encapsulated in silk are called cocoons while the uncovered pupae of butterflies are called chrysalides.

Mating


Males usually get a head start, and start eclosion or emergence, earlier than females and peak in numbers before females. Both of the sexes are sexually mature by the time of eclosion. Butterflies and moths normally don't associate with each other, except for migrating species, staying relatively asocial. Mating begins with an adult (female or male) attracting a mate, normally using visual stimuli, especially in diurnal species like most butterflies
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...

. However, the females of most nocturnal species, including almost all moth species, use pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...

s to attract males, sometimes from long distances. Some species engage in a form of acoustic courtship, or attract mates using sound or vibration such as the polka-dot wasp moth, Syntomeida epilais
Syntomeida epilais
The Polka-Dot Wasp Moth is a species of moth thought to be native to the Caribbean. The species is also called the Oleander Moth after the Oleander plant, from which its young feed. Like most wasp moths, these moths are day fliers....

.

Adaptations include undergoing one seasonal generation, two or even more, called voltinism
Voltinism
Voltinism is a term used in biology to indicate the number of broods or generations of an organism in a year. The term is particularly in use in sericulture, where silkworm varieties vary in their voltinism....

 (Univoltism, bivoltism and multivism respectively). Most lepidoptera in temperate climates are univoltine, while in tropical climates most have two seasonal broods. Some others may take advantage of any opportunity they can get, and mate continuously throughout the year. These seasonal adaptations are controlled by hormones, and these delays in reproduction are called diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

. Many lepidopteran species, after mating and laying their eggs, die shortly afterwards, having only lived for a few days after eclosion. Others may still be active for several weeks and then overwinter and become sexually active again when the weather becomes more favorable, or diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

. The sperm of the male that mated most recently with the female is most likely to have fertilized the eggs but the sperm from a prior mating may still prevail.

Life cycle



Eggs


Lepidoptera usually reproduce sexually and are oviparous
Oviparity
Oviparous animals are animals that lay eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, reptiles, all birds, the monotremes, and most insects, some molluscs and arachnids....

 (egg-laying), though some species exhibit live birth in a process called ovoviviparity
Ovoviviparity
Ovoviviparity, ovovivipary, or ovivipary, is a mode of reproduction in animals in which embryos develop inside eggs that are retained within the mother's body until they are ready to hatch...

. There are a variety of differences in egg-laying and the number of eggs laid. Some species simply drop their eggs in flight (these species normally have polyphagous larvae, meaning they eat a variety of plants e. g., hepialid
Hepialidae
The Hepialidae is a family of insects in the lepidopteran order. Moths of this family are often referred to as swift moths or ghost moths.-Taxonomy and systematics:...

s and some nymphalids
Nymphalidae
The Nymphalidae is a family of about 5,000 species of butterflies which are distributed throughout most of the world. These are usually medium sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called...

) while most Lepidoptera will lay their eggs near or on the host plant that the larvae feed on. The number of eggs laid may vary from only a few to several thousand. The females of both butterflies and moths select the host plant instinctively primarily by chemical cues.

The egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...

 is covered by a hard-ridged protective outer layer of shell, called the chorion. It is lined with a thin coating of wax, which prevents the egg from drying out before the larva has had time to fully develop. Each egg contains a number of micropyles, or tiny funnel-shaped openings at one end, the purpose of which is to allow sperm to enter and fertilize the egg. Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate.

The egg stage lasts a few weeks in most butterflies but eggs laid close to winter, especially in temperate regions, go through a diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

, and hatching may be delayed until spring. Other butterflies may lay their eggs in the spring and have them hatch in the summer. These butterflies are usually northern species (e. g. Nymphalis antiopa
Nymphalis antiopa
Nymphalis antiopa, known as the Mourning Cloak in North America and the Camberwell Beauty in Britain, is a large butterfly native to Eurasia and North America. See also Anglewing butterflies. The immature form of this species is sometimes known as the spiny elm caterpillar. Other older names for...

).

Larvae


The larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

e or caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

s are the first stage in the life cycle after hatching. Caterpillars, are "characteristic polypod larvae with cylindrical bodies, short thoracic legs and abdominal prolegs (pseudopods)". They have a toughened (sclerotised
Sclerite
A sclerite is a hardened body part. The term is used in various branches of biology for various structures including hardened portions of sponges, but it is most commonly used for the hardened portions of arthropod exoskeletons....

) head capsule, mandibles (mouthparts) for chewing, and a soft tubular, segmented body, that may have hair-like or other projections, three pairs of true legs, and additional proleg
Proleg
A Proleg is the small fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the order Lepidoptera, though they can also be found on other larval insects such as sawflies and a few types of flies....

s (up to five pairs). The body consists of thirteen segments, of which three are thoracic and ten are abdominal. Most larvae are herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

s, but a few are carnivore
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

s (some eat ants or other caterpillars) and detritivore
Detritivore
Detritivores, also known as detritophages or detritus feeders or detritus eaters or saprophages, are heterotrophs that obtain nutrients by consuming detritus . By doing so, they contribute to decomposition and the nutrient cycles...

s.

Different herbivore species have adapted to feed on every part of the plant and are normally considered pests to their host plant; some species have been found to lay their eggs on the fruit and other species lay their eggs on clothing or fur (e. g., Tineola bisselliella, the common clothes moth). Some species are carnivorous and others are even parasitic. Some lycaenid species such as Maculinea rebeli
Maculinea rebeli
Phengaris rebeli , common name Mountain Alcon Blue, is a species of butterfly in the Lycaenidae family. It is found in Andorra, Austria, Belgium, Bulgaria, Croatia, Czech Republic, France, Germany, Hungary, Italy, Liechtenstein, Poland, Russia, Slovakia, Slovenia, Spain, and Switzerland. Its...

are social parasites of Myrmica
Myrmica
Myrmica is a genus of ants within the subfamily Myrmicinae. It is widespread throughout the temperate regions of the Holarctic and high mountains in Southeast Asia. The genus consists of around 200 known species, and additional subspecies, although this figure is likely only to rise as soon as...

ants nests. A species of Geometridae from Hawaii
Hawaii
Hawaii is the newest of the 50 U.S. states , and is the only U.S. state made up entirely of islands. It is the northernmost island group in Polynesia, occupying most of an archipelago in the central Pacific Ocean, southwest of the continental United States, southeast of Japan, and northeast of...

 has carnivorous larvae that catch and eat flies. Some pyralid caterpillars are aquatic.

The larvae develop rapidly with several generations in a year; however, some species may take up to 3 years to develop and exceptional examples like Gynaephora groenlandica
Gynaephora groenlandica
Gynaephora groenlandica is a Lymantriid moth found in the Arctic circle found in Greenland and Canada. It is best known for its very slow rate of development. It was once estimated that it had a 14 year life cycle from egg to adult moth, a unique life-cycle among the Lepidoptera with the ability...

take as long as seven years. The larval stage is where the feeding and growing stages occur, and the larvae periodically undergo hormone-induced ecdysis
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...

, developing further with each instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...

, until they undergo the final larval-pupal molt. Lepidoptera pupa, known as Pupa#Chrysalis, have functional mandibles and with appendages fused or glued to the body in most species, while the pupal mandibles are not functional in others.

The larvae of both butterflies and moths exhibit mimicry to deter potential predators. Some caterpillars have the ability to inflate parts of their head to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called osmeteria
Osmeterium
The osmeterium is a fleshy organ found in the prothoracic segment of larvae of Swallowtail butterflies including Birdwings. This organ emits smelly compounds believed to be pheromones. Normally hidden, this forked structure can be everted when the caterpillar is threatened, and used to emit a...

, which are averted to produce smelly chemicals. These are used in defense. Host plants often have toxic substances in them and caterpillars are able to sequester these substances and retain them into the adult stage. This helps making them unpalatable to birds and other predators. Such unpalatability is advertised using bright red, orange, black or white warning colors. The toxic chemicals in plants are often evolved specifically to prevent them from being eaten by insects. Insects in turn develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants.

Wing development


Any form of wings are externally visible on the larva, however when larvae are dissected, developing wings can be seen as disks, which can be found on the second and third thoracic segments, in place of the spiracles that are apparent on abdominal segments. Wing disks develop in association with a trachea that runs along the base of the wing, and are surrounded by a thin peripodial membrane, which is linked to the outer epidermis of the larva by a tiny duct. Wing disks are very small until the last larval instar, when they increase dramatically in size, are invaded by branching tracheae
Invertebrate trachea
The invertebrate trachea refers to the open respiratory system composed of spiracles, tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues....

 from the wing base that precede the formation of the wing veins, and begin to develop patterns associated with several landmarks of the wing.

Near pupation, the wings are forced outside the epidermis under pressure from the hemolymph
Hemolymph
Hemolymph, or haemolymph, is a fluid in the circulatory system of some arthropods and is analogous to the fluids and cells making up both blood and interstitial fluid in vertebrates such as birds and mammals...

, and although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval cuticle they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled without damage to the wings.

Pupa


After about 5 to 7 instars, or molts, certain hormones, like prothoracicotropic hormone
Prothoracicotropic hormone
Prothoracicotropic hormone was the first insect hormone that was discovered. It was originally described simply as "brain hormone" by early workers such as Stefan Kopeć and Vincent Wigglesworth , who realized that ligation of the head of immature insects could prevent molting or pupation of the...

, stimulate the production of ecdysone
Ecdysone
Ecdysone is a steroidal prohormone of the major insect molting hormone 20-hydroxyecdysone, which is secreted from the prothoracic glands. Insect molting hormones are generally called ecdysteroids. Ecdysteroids act as moulting hormones of arthropods but also occur in other related phyla where they...

, which initiates insect molting. Then, the larva puparium, a sclerotized or hardened cuticle of the last larval instar, develops into the pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

. Depending on the species, the pupa may be covered in silk and attached to many different types of debris or may not be covered at all. The pupa stays attached to the leaf by silk spun by the caterpillar before it spins the silk for the full pupa. Features of the imago are externally recognizable in the pupa. All the appendages that are found on the adult head and thorax are found cased inside the cuticle (antenna
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....

e, mouthpart
Insect mouthparts
Insects exhibit a range of mouthparts, adapted to particular modes of feeding. The earliest insects had chewing mouthparts...

s, etc.), with the wings wrapped around, adjacent to the antennae.

While encased, some of the lower segments are not fused, and are able to move using small muscles found in between the membrane. Moving may help the pupa, for example, escape the sun, which would otherwise kill it. The pupa of the Mexican jumping bean
Mexican jumping bean
A Mexican jumping bean is an occurrence native to Mexico, where it is known as Frijoles saltarines . Physically, Mexican jumping beans resemble small beans, tan to brown in colour. They are a seed pod through which the larva of a small moth has chewed. The bean "jumps" because when it gets in a...

 moth (Cydia deshaisiana
Cydia deshaisiana
Cydia deshaisiana or jumping bean moth is a moth from Mexico that is most widely known as its larva, where it inhabits the carpels of seeds from several species of the shrub genus Sebastiania . These seeds are commonly known as Mexican jumping beans.The moth lays the egg on the young pod...

) does this. The larvae cut a trapdoor in the bean (species of Sebastiania
Sebastiania
Sebastiania is a plant genus in the Euphorbiaceae family, comprising about 100 species. These are shrubs found in tropical and warm areas.Species include:*Sebastiania alpina*Sebastiania crenulata*Sebastiania fasciculata...

) and use the bean as a shelter. When there is a sudden rise in temperature, the pupa inside twitches and jerks, pulling on the threads inside. Wiggling may also help to deter parasitoid
Parasitoid
A parasitoid is an organism that spends a significant portion of its life history attached to or within a single host organism in a relationship that is in essence parasitic; unlike a true parasite, however, it ultimately sterilises or kills, and sometimes consumes, the host...

 wasps from laying eggs on the pupa. Other species of moth are able to make clicks to deter predators.

The length of time before the pupa ecloses (emerges) varies greatly. The monarch butterfly may stay in its chrysalis for two weeks, while other species may need to stay for more than 10 months in diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

. The adult will emerge from the pupa either by using abdominal hooks or from projections located on the head. The mandibles found in the most primitive moth families are used to escape from their cocoon (e. g., Micropterigoidea).

Adult


Most lepidopteran species do not live long after eclosion, only needing a few days to find a mate and then lay their eggs. Others may remain active for from one to several weeks or go through diapause, overwintering as monarch butterflies do, or waiting out environmental stress. Some adult species of Microlepidoptera
Microlepidoptera
Microlepidoptera is an artificial grouping of moth families, commonly known as the 'smaller moths' . These generally have a wingspan of under 20 mm, and are thus harder to identify by external phenotypic markings than macrolepidoptera...

 go through a stage where there is no reproductive-related activity lasting through summer and winter, followed by mating and oviposition
Oviposition
Oviposition is the process of laying eggs by oviparous animals.Some arthropods, for example, lay their eggs with an organ called the ovipositor.Fish , amphibians, reptiles, birds and monetremata also lay eggs....

, or egg laying, in the early spring.

While most butterflies and moths are terrestrial
Terrestrial animal
Terrestrial animals are animals that live predominantly or entirely on land , as compared with aquatic animals, which live predominantly or entirely in the water , or amphibians, which rely on a combination of aquatic and terrestrial habitats...

, many species of Pyralidae
Pyralidae
The Pyralidae or snout moths are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many classifications, the grass moths are included in the Pyralidae as a subfamily, making the combined group one of the largest families in the Lepidoptera...

 are truly aquatic
Aquatic animal
An aquatic animal is an animal, either vertebrate or invertebrate, which lives in water for most or all of its life. It may breathe air or extract its oxygen from that dissolved in water through specialised organs called gills, or directly through its skin. Natural environments and the animals that...

 with all stages except the adult occurring in water. Many species from other families such as Arctiidae
Arctiidae
Arctiidae is a large and diverse family of moths with around 11,000 species found all over the world, including 6,000 neotropical species. This family includes the groups commonly known as tiger moths , which usually have bright colours, footmen , lichen moths and wasp moths...

, Nepticulidae
Nepticulidae
Nepticulidae is a family of very small moths with a worldwide distribution. They are characterised by eyecaps over the eyes . These pigmy moths or midget moths, as they are commonly known, include the smallest of all living moths, with a wingspan that can be as little as 3 mm...

, Cosmopterygidae, Tortricidae
Tortricidae
Tortricidae is a family of moths, commonly known as tortrix moths, in the order Lepidoptera. Tortricidae is a large family with over 9,400 species described, and is the sole member of the superfamily Tortricoidea. Many of these are economically important pests. Olethreutidae is a junior synonym...

, Olethreutidae, Noctuidae
Noctuidae
The Noctuidae or owlet moths are a family of robustly-built moths that includes more than 35,000 known species out of possibly 100,000 total, in more than 4,200 genera. They constitute the largest family in the Lepidoptera....

, Cossidae
Cossidae
Cossidae, the cossid millers or carpenter millers, make up a family of mostly large miller moths. Ths family contains over 110 genera with almost 700 known species, and many more species await description...

 and Sphingidae
Sphingidae
Sphingidae is a family of moths , commonly known as hawk moths, sphinx moths and hornworms, that includes about 1,200 species . It is best represented in the tropics but there are species in every region . They are moderate to large in size and are distinguished among moths for their rapid,...

 are aquatic or semi-aquatic.

Flight


Flight is an important aspect of the lives of butterflies and moths and is used for evading predators, searching for food and finding mates in a timely manner as lepidopteran species do not live long after eclosion. It is the main form of locomotion in most species. In lepidoptera, the forewings and hindwings are mechanically coupled and flap in synchrony. Flight is anteromotoric, or being driven primarily by action of the forewings. Although it has been reported that lepidopteran species can still fly when their hindwings are cut off, it reduces their linear flight and turning capabilities.

Lepidopteran species have to be warm, about 77 to 79 °F (25 to 26.1 C) in order to fly. They depend on their body temperature being sufficiently high and since they can't regulate it themselves, this is dependent on their environment. Butterflies living in cooler climates may use their wings to warm their bodies. They will bask in the sun, spreading out their wings so that they get maximum exposure to the sunlight. In hotter climates butterflies can easily overheat, so they are usually active only during the cooler parts of the day, early morning, late afternoon or early evening. During the heat of the day they rest in the shade. Some larger thick-bodied moths (e. g. Sphingidae
Sphingidae
Sphingidae is a family of moths , commonly known as hawk moths, sphinx moths and hornworms, that includes about 1,200 species . It is best represented in the tropics but there are species in every region . They are moderate to large in size and are distinguished among moths for their rapid,...

) can generate their own heat to a limited degree by vibrating their wings. The heat generated by the flight muscles warms the thorax while the temperature of the abdomen is unimportant for flight. To avoid overheating some moths rely on hairy scales, internal air sacs, and other structures to separate the thorax and abdomen and keep the abdomen cooler.

Some species of butterfly can reach fast speeds, such as the Southern Dart
Ocybadistes walkeri
Ocybadistes walkeri, commonly known as the Greenish Grass-dart, Green Grass-dart, Southern Dart or Yellow-banded Dart, is a type of butterfly known as a Skipper found in eastern and southern Australia, with one subspecies found in the Northern Territory.The larvae feed on Dianella, Brachypodium...

, which can go as fast as 48.4  km/h. Sphingids
Sphingidae
Sphingidae is a family of moths , commonly known as hawk moths, sphinx moths and hornworms, that includes about 1,200 species . It is best represented in the tropics but there are species in every region . They are moderate to large in size and are distinguished among moths for their rapid,...

 are some of the fastest flying insects, some are capable of flying at over 50  km/h (30 miles per hour), having a wingspan of 35–150 mm. In some species, there is sometimes a gliding component to their flight. Flight occurs either as hovering, or as forward or backward motion. In butterfly and in moth species, like hawk moth
Sphingidae
Sphingidae is a family of moths , commonly known as hawk moths, sphinx moths and hornworms, that includes about 1,200 species . It is best represented in the tropics but there are species in every region . They are moderate to large in size and are distinguished among moths for their rapid,...

s, hovering is important in that they need to hover over flowers when feeding on the nectar.

Navigation


Navigation
Navigation
Navigation is the process of monitoring and controlling the movement of a craft or vehicle from one place to another. It is also the term of art used for the specialized knowledge used by navigators to perform navigation tasks...

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

Many studies have also shown that moths navigate. One study showed that many moths may use the Earth's magnetic field
Earth's magnetic field
Earth's magnetic field is the magnetic field that extends from the Earth's inner core to where it meets the solar wind, a stream of energetic particles emanating from the Sun...

 to navigate, as a study of the moth Heart and Dart
Heart and Dart
The Heart and Dart is a moth of the family Noctuidae. A familiar moth to many, it is considered one of the most common of the European region....

 suggests. Another study, this time of the migratory behavior of the Silver Y
Silver Y
The Silver Y is a migratory moth of the family Noctuidae which is named for the silvery Y-shaped mark on each of its forewings.-Description:...

, showed that even at high altitudes the species can correct its course with changing winds, and prefers flying with favourable winds, suggesting a great sense of direction. Aphrissa statira
Aphrissa statira
The Statira Sulphur, Aphrissa statira, is a species of lepidoptera in the family Pieridae.-Description:Aphrissa statira have a wingspan of about to ....

in Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

 loses its navigational capacity when exposed to a magnetic field, suggesting it uses the Earth’s magnetic field.

Moths exhibit a tendency to circle artificial lights repeatedly. This suggests that they use a technique of celestial navigation
Celestial navigation
Celestial navigation, also known as astronavigation, is a position fixing technique that has evolved over several thousand years to help sailors cross oceans without having to rely on estimated calculations, or dead reckoning, to know their position...

 called transverse orientation
Transverse orientation
Transverse orientation, keeping a fixed angle on a distant source of light for orientation, is a proprioceptive response displayed by many moths.....

. By maintaining a constant angular relationship to a bright celestial light, such as the Moon
Moon
The Moon is Earth's only known natural satellite,There are a number of near-Earth asteroids including 3753 Cruithne that are co-orbital with Earth: their orbits bring them close to Earth for periods of time but then alter in the long term . These are quasi-satellites and not true moons. For more...

, they can fly in a straight line. Celestial objects are so far away, that even after traveling great distances, the change in angle between the moth and the light source is negligible; further, the moon will always be in the upper part of the visual field or on the horizon
Horizon
The horizon is the apparent line that separates earth from sky, the line that divides all visible directions into two categories: those that intersect the Earth's surface, and those that do not. At many locations, the true horizon is obscured by trees, buildings, mountains, etc., and the resulting...

. When a moth encounters a much closer artificial light and uses it for navigation, the angle changes noticeably after only a short distance, in addition to being often below the horizon. The moth instinctively attempts to correct by turning toward the light, causing airborne moths to come plummeting downwards, and—at close range—which results in a spiral flight path that gets closer and closer to the light source. Other explanations have been suggested, such as the idea that moths may be impaired with a visual distortion called a Mach band by Henry Hsiao in 1972. He stated that they fly towards the darkest part of the sky in pursuit of safety and are thus inclined to circle ambient objects in the Mach band region.

Migration



Lepidopteran migration is usually season
Season
A season is a division of the year, marked by changes in weather, ecology, and hours of daylight.Seasons result from the yearly revolution of the Earth around the Sun and the tilt of the Earth's axis relative to the plane of revolution...

al, the insects moving to escape dry seasons or other disadvantageous conditions. Most lepidopterans that migrate are butterflies, the distance travelled varying from short to very long journeys. Some butterflies that migrate include the Mourning Cloak
Nymphalis antiopa
Nymphalis antiopa, known as the Mourning Cloak in North America and the Camberwell Beauty in Britain, is a large butterfly native to Eurasia and North America. See also Anglewing butterflies. The immature form of this species is sometimes known as the spiny elm caterpillar. Other older names for...

, Painted Lady
Painted Lady
The Cynthia group of colourful butterflies, commonly called painted ladies, comprises a subgenus of the genus Vanessa in the Family Nymphalidae...

, American Lady
American Painted Lady
The American Painted Lady or American Lady is a butterfly found throughout North America.Vanessa virginiensis lives in flowery habitats, usually in mountains. The larvae feed on various Asteraceae, especially the cudweeds of genus Gnaphalium...

, Red Admiral, and the Common Buckeye
Junonia coenia
The Buckeye is a butterfly in the family Nymphalidae. It is found in southern Manitoba, Ontario, Quebec, and Nova Scotia and all parts of the United States except the northwest, and is especially common in the south, the California coast, and throughout Central America and Colombia...

. Particularly famous migrations are those of the Monarch butterfly from Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...

 to northern United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

 and southern Canada
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...

, a distance of about 4000 kilometre. Other well known migratory species include the Painted Lady
Vanessa cardui
Vanessa cardui is a well-known colourful butterfly, known as the Painted Lady, or in North America as the Cosmopolitan. This butterfly has a strange pattern of flying in a sort of screw shape.-Distribution:...

 and several of the danaine butterflies. Spectacular and large scale migrations associated with the Monsoons are seen in peninsular India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

. Migrations have been studied in more recent times using wing tags and also using stable hydrogen isotopes.

Moths also undertake migrations, an example being the uraniids
Uraniidae
The Uraniidae are a family of moths containing four subfamilies, ninety genera, and roughly seven-hundred species. The family is distributed throughout the tropics of Americas, Africa and Indo-Australia. Some of the tropical species are known for their bright, butterfly-like colors and are called...

. Urania fulgens
Urania fulgens
Urania fulgens is a day-flying moth of the Uraniidae family. It is found from Veracruz, Mexico, through Central America to northwestern South America . It is highly migratory and has been recorded as a vagrant to Texas, USA.It is sometimes confused with the similar U...

undergoes population explosions and massive migrations that may be not surpassed by any other insect in the Neotropic
Neotropic
In biogeography, the Neotropic or Neotropical zone is one of the eight terrestrial ecozones. This ecozone includes South and Central America, the Mexican lowlands, the Caribbean islands, and southern Florida, because these regions share a large number of plant and animal groups.It is sometimes used...

s. In Costa Rica
Costa Rica
Costa Rica , officially the Republic of Costa Rica is a multilingual, multiethnic and multicultural country in Central America, bordered by Nicaragua to the north, Panama to the southeast, the Pacific Ocean to the west and the Caribbean Sea to the east....

 and Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

, the first population movements may begin in July and early August and, depending on the year, may be very massive, continuing unabated for as long as five months.

Communication


Pheromones are commonly involved in mating rituals among species, especially moths, but they are also an important aspect of other forms of communication. Usually the pheromones are produced by either the male or the female and detected by members of the opposite sex with their antenna
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....

e. In many species, a gland between the eighth and ninth segment under the abdomen in the female produces the pheromones. Communication can also occur through stridulation, or producing sounds by rubbing various parts of the body together.

Moths are known to engage in acoustic forms of communication; most often species engage use it in a form of acoustic courtship, attracting mates using sound or vibration. Like most other insects, moths pick up these sounds using tympanic membranes in the abdomen. An example is that of the polka-dot wasp moth
Syntomeida epilais
The Polka-Dot Wasp Moth is a species of moth thought to be native to the Caribbean. The species is also called the Oleander Moth after the Oleander plant, from which its young feed. Like most wasp moths, these moths are day fliers....

 (Syntomeida epilais), which produce sounds with a frequency above that normally detectable by humans (~20kHZ). These sounds also function as tactile communication, or communication through touch, as they stridulate, or vibrate a substrate like leaves and stems.

Most moths lack bright colors as many species use coloration as camouflage
Camouflage
Camouflage is a method of concealment that allows an otherwise visible animal, military vehicle, or other object to remain unnoticed, by blending with its environment. Examples include a leopard's spotted coat, the battledress of a modern soldier and a leaf-mimic butterfly...

 but butterflies engage in visual communication. Female cabbage butterflies, for example, use ultraviolet light to communicate, with scales colored in this range on the dorsal wing surface. When they fly, each down stroke of the wing creates a brief flash of ultraviolet light that the males apparently recognize as the flight signature of a potential mate. These flashes from the wings may attract several males who engage in aerial courtship displays.

Diapause


One of the most important adaptations is diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

, or delay in development in response to regularly and recurring periods of adverse environmental conditions (winter, dry season, etc.). Diapause normally occurs in eggs, or as a reproductive delay in adults. Butterflies like the monarch may undergo diapause during winter, where they undergo a form of hibernation, lying dormant on trees for protection after their large scale migration. Seasonal adaptations such as voltism, where they may reproduce one or more times annually are due to diapause. This response to environmental stress is controlled by hormones and is necessary to survival during unfavorable times, especially in northern areas and high mountains where winter is regular and harsh. For example, in the Mediterranean
Mediterranean Sea
The Mediterranean Sea is a sea connected to the Atlantic Ocean surrounded by the Mediterranean region and almost completely enclosed by land: on the north by Anatolia and Europe, on the south by North Africa, and on the east by the Levant...

 area, larvae feed during the spring when the vegetation flourishes, then undergo diapause in the summer during drought, and hibernate in the winter.c

Ecology


Moths and Butterflies are important in the natural ecosystem. They are integral participants in the food chain, having co-evolved with flowering plants and predators, lepidopteran species have formed a network of trophic
Trophic level
The trophic level of an organism is the position it occupies in a food chain. The word trophic derives from the Greek τροφή referring to food or feeding. A food chain represents a succession of organisms that eat another organism and are, in turn, eaten themselves. The number of steps an organism...

 relationships between autotroph
Autotroph
An autotroph, or producer, is an organism that produces complex organic compounds from simple inorganic molecules using energy from light or inorganic chemical reactions . They are the producers in a food chain, such as plants on land or algae in water...

s and heterotroph
Heterotroph
A heterotroph is an organism that cannot fix carbon and uses organic carbon for growth. This contrasts with autotrophs, such as plants and algae, which can use energy from sunlight or inorganic compounds to produce organic compounds such as carbohydrates, fats, and proteins from inorganic carbon...

s, which are included in the stages of Lepidoptera larvae, pupae and adults. Larvae and pupae are links in the diet of birds and parasitic entomophagous
Entomophagy
Entomophagy is the consumption of insects as food. Insects are eaten by many animals, but the term is generally used to refer to human consumption of insects; animals that eat insects are known as insectivores...

 insects. The adults are included in food webs in a much broader range of consumers (including birds, small mammals, reptiles, etc.).

Defense and predation



Lepidopteran species are soft bodied, fragile and almost defenseless while the immature stages move slowly or are immobile, hence all stages are exposed to predation
Predation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...

. Adult butterflies and moths are predated upon by bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s, lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s, amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...

s, dragonflies
Dragonfly
A dragonfly is a winged insect belonging to the order Odonata, the suborder Epiprocta or, in the strict sense, the infraorder Anisoptera . It is characterized by large multifaceted eyes, two pairs of strong transparent wings, and an elongated body...

 and spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...

s, besides others. Caterpillars and pupa fall prey, not only to birds but invertebrate predators, small mammals, as well as fungi and bacteria. Parasitoid
Parasitoid
A parasitoid is an organism that spends a significant portion of its life history attached to or within a single host organism in a relationship that is in essence parasitic; unlike a true parasite, however, it ultimately sterilises or kills, and sometimes consumes, the host...

 and parasitic wasps and flies
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...

 may lay eggs in the caterpillar, which eventually kill it as they hatch inside its body and eat its tissues. Insect-eating birds are probably the worst predators. Lepidoptera, especially the immature stages, are an ecologically important food to many insectivorous birds, such as the Great Tit
Great Tit
The Great Tit is a passerine bird in the tit family Paridae. It is a widespread and common species throughout Europe, the Middle East, Central and Northern Asia, and parts of North Africa in any sort of woodland. It is generally resident, and most Great Tits do not migrate except in extremely...

 in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

.

An "evolutionary arms race
Evolutionary arms race
In evolutionary biology, an evolutionary arms race is an evolutionary struggle between competing sets of co-evolving genes that develop adaptations and counter-adaptations against each other, resembling an arms race, which are also examples of positive feedback...

" can be seen between predator and prey species. Lepidoptera have developed a number of strategies for defense and protection including evolution of morphological characters and changes in ecological life-style and in behavior. These include aposematism
Aposematism
Aposematism , perhaps most commonly known in the context of warning colouration, describes a family of antipredator adaptations where a warning signal is associated with the unprofitability of a prey item to potential predators...

, mimicry, camouflage
Camouflage
Camouflage is a method of concealment that allows an otherwise visible animal, military vehicle, or other object to remain unnoticed, by blending with its environment. Examples include a leopard's spotted coat, the battledress of a modern soldier and a leaf-mimic butterfly...

, development of threat patterns and displays and so on. Only a few birds, such as the nightjar
Nightjar
Nightjars are medium-sized nocturnal or crepuscular birds with long wings, short legs and very short bills. They are sometimes referred to as goatsuckers from the mistaken belief that they suck milk from goats . Some New World species are named as nighthawks...

s, hunt nocturnal Lepidoptera and their main enemy are bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...

s. Again, an "evolutionary race" exists, which has led to numerous evolutionary adaptations of moths to escape from their main predators, such as the ability to hear ultrasonic sounds, or even to emit sounds in some cases. Lepidoptera eggs are also predated upon. Some caterpillars, such as the zebra swallowtail butterfly
Zebra Swallowtail Butterfly
The Zebra Swallowtail is a swallowtail butterfly native to the eastern United States and southeast Canada. Its distinctive wing shape and long tails make it easy to identify, and its black and white-striped pattern is reminiscent of a zebra...

 larvae, are cannibalistic and may eat other larvae of the same species. Lepidopteran species rely on a variety of strategies.

Some species of lepidoptera are poisonous to predators, such as the Monarch butterfly in the Americas, Atrophaneura
Atrophaneura
Atrophaneura, commonly referred to as the Red-bodied Swallowtails is a genus of butterflies in the Swallowtail family that are generally found in Asia.-Species:Listed alphabetically within groups:The latreillei species-group:...

 species (roses, windmills etc.) in Asia, as well as Papilio antimachus
Papilio antimachus
Papilio antimachus is a butterfly in the family Papilionidae. With a wingspan between , it is the largest butterfly in Africa and among the largest butterflies in the world. Papilio antimachus live in the tropical rainforests of West and Central Africa...

 and the birdwing
Birdwing
Birdwings are papilionid butterflies native to the Indian Subcontinent, mainland and archipelagic Southeast Asia and Australasia, and are usually regarded as belonging to three genera: Ornithoptera, Trogonoptera and Troides. Some authorities include additional genera...

s, the largest butterflies in Africa and Asia respectively. They obtain their toxicity by sequestering the chemicals from the plants they eat into their own tissues. Some Lepidoptera manufacture their own toxins. Predators that eat poisonous butterflies and moths may become sick and vomit violently, learning not to eat those types of species. A predator who has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future, thus saving many other species as well. Toxic butterflies and larvae tend to develop bright colors, striking patterns as an indicator to predators about their toxicity. This phenomenon is known as aposematism
Aposematism
Aposematism , perhaps most commonly known in the context of warning colouration, describes a family of antipredator adaptations where a warning signal is associated with the unprofitability of a prey item to potential predators...

. Other caterpillars emit bad smells to ward off predators. Some caterpillars, especially members of Papilionidae, contain an osmeterium
Osmeterium
The osmeterium is a fleshy organ found in the prothoracic segment of larvae of Swallowtail butterflies including Birdwings. This organ emits smelly compounds believed to be pheromones. Normally hidden, this forked structure can be everted when the caterpillar is threatened, and used to emit a...

, a Y-shaped protrusible gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...

 found in the prothoracic
Prothorax
The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites are the pronotum , the prosternum , and the propleuron on each side. The prothorax never bears wings in extant insects, though some fossil groups possessed...

 segment of the larvae. When threatened, the caterpillar emits unpleasant smells from the organ to ward off the predators.

Camouflage
Camouflage
Camouflage is a method of concealment that allows an otherwise visible animal, military vehicle, or other object to remain unnoticed, by blending with its environment. Examples include a leopard's spotted coat, the battledress of a modern soldier and a leaf-mimic butterfly...

 is also important defense strategies, which involves the use of coloration or shape to blend into the surrounding environment. Some lepidopteran species blend with its surroundings, making them difficult to be spotted by predators. Caterpillars can be shades of green that matches its host plant. Others look like inedible objects, such as twigs or leaves. The larvae of some species, such as the Common Mormon (Papilio polytes
Papilio polytes
The Common Mormon is a common species of swallowtail butterfly widely distributed across Asia. This butterfly is known for the mimicry displayed by the numerous forms of its females which mimic inedible Red-bodied Swallowtails, such as the Common Rose and the Crimson Rose.- Names :The common name...

) and the Western Tiger Swallowtail
Western Tiger Swallowtail
The Western Tiger Swallowtail, Papilio rutulus, is a common swallowtail butterfly of western North America, frequently seen in urban parks and gardens as well as in rural woodlands and riparian areas...

 look like bird droppings. For example, adult Sesiidae
Sesiidae
The Sesiidae or clearwing moths are family of the Lepidoptera in which the wings partially have hardly any of the normal lepidopteran scales, leaving them transparent. The bodies are generally striped with yellow, red or white, sometimes very brightly, and they have simple antennae...

 species (also known as clearwing moths) have a general appearance that is sufficiently similar to a wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...

 or hornet
Hornet
Hornets are the largest eusocial wasps; some species can reach up to in length. The true hornets make up the genus Vespa and are distinguished from other vespines by the width of the vertex , which is proportionally larger in Vespa and by the anteriorly rounded gasters .- Life cycle :In...

 to make it likely that the moths gain a reduction in predation by Batesian mimicry. Eyespot
Eyespot (mimicry)
An eyespot is an eye-like marking. They are found on butterflies, reptiles, birds and fish. In members of the Felidae family , the white circular markings on the backs of the ears are termed ocelli, and they are functionally similar to eyespots in other animals.Eyespots may be a form of...

s are a type of automimicry used by some butterflies and moths. In butterflies, the spots are composed of concentric rings of scales of different colors. The proposed role of the eyespots is to deflect attention to predators. Their resemblance to eyes provokes the predator's instinct to attack these wing patterns.

Batesian
Batesian mimicry
Batesian mimicry is a form of mimicry typified by a situation where a harmless species has evolved to imitate the warning signals of a harmful species directed at a common predator...

 and Müllerian
Müllerian mimicry
Müllerian mimicry is a natural phenomenon when two or more harmful species, that may or may not be closely related and share one or more common predators, have come to mimic each other's warning signals...

 mimicry complexes are commonly found in Lepidoptera. Genetic polymorphism and natural selection give rise to otherwise edible species (the mimic) gaining a survival advantage by resembling inedible species (the model). Such a mimicry complex is referred to as Batesian and is most commonly known by the mimicry by the limenitidine
Limenitidinae
Limenitidinae is a subfamily of butterflies that includes the admirals and relatives. The common names of many species and genera reference military ranks or – namely the Adoliadini – titles of nobility , in reference to these butterflies' large size, bold pattern and dashing flight...

 Viceroy butterfly
Viceroy butterfly
The Viceroy Butterfly is a North American butterfly with a range from the Northwest Territories along the eastern edges of the Cascade Range and Sierra Nevada mountains, southwards into central Mexico....

 of the inedible danaine Monarch. Later research has discovered that the Viceroy is, in fact more toxic than the Monarch and this resemblance should be considered as a case of Müllerian mimicry. In Müllerian mimicry, inedible species, usually within a taxonomic order, find it advantageous to resemble each other so as to reduce the sampling rate by predators who need to learn about the insects' inedibility. Taxa from the toxic genus Heliconius
Heliconius
Heliconius comprise a colorful and widespread brush-footed butterfly genus distributed throughout the tropical and subtropical regions of the New World. These butterflies utilize Passion flower plants as their larval food source and rely on bright wing color patterns to signal their distastefulness...

form one of the most well known Müllerian complexes. The adults of the various species now resemble each other so well that the species cannot be distinguished without close morphological observation and, in some cases, dissection or genetic analysis.

There is evidence moths are able to hear the range emitted by bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...

s, which in effect causes flying moths to make evasive maneuvers because bats are a main predator of moths. Ultrasonic frequencies trigger a reflex action in the noctuid moth that cause it to drop a few inches in its flight to evade attack. Tiger moth
Arctiidae
Arctiidae is a large and diverse family of moths with around 11,000 species found all over the world, including 6,000 neotropical species. This family includes the groups commonly known as tiger moths , which usually have bright colours, footmen , lichen moths and wasp moths...

s in a defense emit clicks within the same range of the bats, which interfere with the bats, and foil their attempts to echolocate it.

Pollination



Most species of Lepidoptera engage in some form of entomophily
Entomophily
Entomophily is a form of pollination whereby pollen is distributed by insects. Several insect are reported to be responsible for the pollination of many plant species, particularly bees, Lepidoptera , wasps, flies, ants and beetles. Some plant species co-evolved with a particular pollinator, such...

 (more specifically psychophily and phalaenophily for butterflies and moths respectively), or the pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...

 of flowers. Most adult butterflies and moths feed on the nectar inside flowers, using their proboscis to reach the nectar hidden at the base of the petals. In the process, the adult brushes against the flower's stamen
Stamen
The stamen is the pollen producing reproductive organ of a flower...

, on which the flower's reproductive pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

 is made and stored. The pollen is transferred on appendages on the adult, who flies to the next flower to feed and unwittingly deposits the pollen on the stigma
Stigma (botany)
The stigma is the receptive tip of a carpel, or of several fused carpels, in the gynoecium of a flower. The stigma receives pollen at pollination and it is on the stigma that the pollen grain germinates. The stigma is adapted to catch and trap pollen with various hairs, flaps, or sculpturings...

 of the next flower, where the pollen germinates
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

 and fertilizes the seeds.

Flowers pollinated by butterflies tend to be large and flamboyant, being pink or lavender in color, frequently having a landing area, and are usually scented, as butterflies are typically day-flying. Since butterflies do not digest
Digestion
Digestion is the mechanical and chemical breakdown of food into smaller components that are more easily absorbed into a blood stream, for instance. Digestion is a form of catabolism: a breakdown of large food molecules to smaller ones....

 pollen (except for Heliconid species
Heliconius
Heliconius comprise a colorful and widespread brush-footed butterfly genus distributed throughout the tropical and subtropical regions of the New World. These butterflies utilize Passion flower plants as their larval food source and rely on bright wing color patterns to signal their distastefulness...

), more nectar is offered than pollen. The flowers have simple nectar guides with the nectaries usually hidden in narrow tubes or spurs, reached by the long tongue of the butterflies. Butterflies like the Thymelicus flavus
Thymelicus
Thymelicus is a genus in the skipper butterfly family, Hesperiidae. It is presently the only member of the tribe Thymelini, but many skipper butterflies are yet to be assigned to tribes, so this might change eventually.-Species:...

have been observed to engage in flower constancy
Flower constancy
Flower constancy or pollinator constancy is defined as the tendency of individual pollinators to exclusively visit certain flower species or morphs within a species, bypassing other available flower species that could potentially be more rewarding...

, which means that they are more likely to transfer pollen to other conspecific plants. This can be beneficial for the plants being pollinated, as flower constancy prevents the loss of pollen during different flights and the pollinators from clogging stigmas with pollen of other flower species.

Among the more important moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...

 pollinators are the hawk moths (Sphingidae
Sphingidae
Sphingidae is a family of moths , commonly known as hawk moths, sphinx moths and hornworms, that includes about 1,200 species . It is best represented in the tropics but there are species in every region . They are moderate to large in size and are distinguished among moths for their rapid,...

). Their behavior is similar to hummingbird
Hummingbird
Hummingbirds are birds that comprise the family Trochilidae. They are among the smallest of birds, most species measuring in the 7.5–13 cm range. Indeed, the smallest extant bird species is a hummingbird, the 5-cm Bee Hummingbird. They can hover in mid-air by rapidly flapping their wings...

s: Using rapid wing beats to keep hovered in front of flowers. Most being nocturnal or crepuscular
Crepuscular
Crepuscular animals are those that are active primarily during twilight, that is during dawn and dusk. The word is derived from the Latin word crepusculum, meaning "twilight." Crepuscular is, thus, in contrast with diurnal and nocturnal behavior. Crepuscular animals may also be active on a bright...

, so moth-pollinated flowers (e.g., Silene latifolia ) tend to be white, night-opening, large and showy with tubular corollas and a strong, sweet scent produced in the evening, night or early morning. A lot of nectar is produced to fuel the high metabolic rates needed to power their flight. Other moths (e.g., noctuids
Noctuidae
The Noctuidae or owlet moths are a family of robustly-built moths that includes more than 35,000 known species out of possibly 100,000 total, in more than 4,200 genera. They constitute the largest family in the Lepidoptera....

, geometrids
Geometer moth
The geometer moths or Geometridae are a family of the order Lepidoptera...

, pyralids
Pyraloidea
The Pyraloidea are a moth superfamily containing about 16,000 described species worldwide , and probably at least as many more remain to be described. They are generally fairly small moths....

) fly slowly and settle on the flower. They do not require as much nectar as the fast-flying hawk moths, and the flowers tend to be small (though they may be aggregated in heads).

Mutualism


Mutualism is a form of biological interaction
Biological interaction
Biological interactions are the effects organisms in a community have on one another. In the natural world no organism exists in absolute isolation, and thus every organism must interact with the environment and other organisms...

 where each individual involves benefits in some shape or form. An example of a mutualistic relationship would be the relationship shared by yucca moths
Tegeticula
Tegeticula is a genus of moths of the Prodoxidae family.-Species:* Tegeticula altiplanella* Tegeticula baccatella* Tegeticula california* Tegeticula cassandra* Tegeticula corruptrix* Tegeticula elatella...

 (Tegeculidae) and their host, yucca flowers
Yucca
Yucca is a genus of perennial shrubs and trees in the family Asparagaceae, subfamily Agavoideae. Its 40-50 species are notable for their rosettes of evergreen, tough, sword-shaped leaves and large terminal panicles of white or whitish flowers. They are native to the hot and dry parts of North...

 (Liliaceae). Female yucca moths enter the host flowers, collect the pollen into a ball using specialized maxillary palps, then move to the apex of the pistil where pollen is deposited on the stigma, and lay eggs into the base of the pistil where seeds will develop. The larvae develop in the fruit pod and feed on a portion of the seeds. Thus, both insect and plant benefit, forming a highly mutualistic relationship. Another form of mutualism occurs between some larvae of butterflies and certain species of ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s (e. g. Lycaenidae
Lycaenidae
The Lycaenidae are the second-largest family of butterflies, with about 6000 species worldwide, whose members are also called gossamer-winged butterflies...

). The larvae communicate with the ants using vibrations that are transmitted through a substrate, such as the wood of a tree or stems, as well as using chemical signals. The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions
Honeydew (secretion)
Honeydew is a sugar-rich sticky liquid, secreted by aphids and some scale insects as they feed on plant sap. When their mouthpart penetrates the phloem, the sugary, high-pressure liquid is forced out of the gut's terminal opening. Honeydew is particularly common as a secretion in the Hemipteran...

.

Parasitism


There are only 41 known species of parasitoid
Parasitoid
A parasitoid is an organism that spends a significant portion of its life history attached to or within a single host organism in a relationship that is in essence parasitic; unlike a true parasite, however, it ultimately sterilises or kills, and sometimes consumes, the host...

 lepidoptera (1-Pyralidae
Pyralidae
The Pyralidae or snout moths are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many classifications, the grass moths are included in the Pyralidae as a subfamily, making the combined group one of the largest families in the Lepidoptera...

; 40-Epipyropidae
Epipyropidae
Epipyropidae is a small family of moths. This family and the closely related Cyclotornidae are unique among the Lepidoptera in that the larvae are ectoparasites, the hosts typically being fulgoroid planthoppers, thus the common name Planthopper Parasite Moths.-References:*...

). The larvae of the Greater
Galleria mellonella
The Greater Wax Moth or Honeycomb Moth is a moth of the family Pyralidae. It is the only member of the genus Galleria. It is found in most of the world, including Europe and adjacent Eurasia , and as an introduced species e.g...

 and Lesser wax moth
Lesser wax moth
Achroia grisella, the Lesser Wax Moth, is a small moth of the snout moth family , wherein it belongs to the subfamily Galleriinae. It is the only unequivocally recognized species of genus Achroia...

s feed on the honeycomb
Honeycomb
A honeycomb is a mass of hexagonal waxcells built by honey bees in their nests to contain their larvae and stores of honey and pollen.Beekeepers may remove the entire honeycomb to harvest honey...

 inside bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...

 nests and may become pests; they are also found in bumblebee
Bumblebee
A bumble bee is any member of the bee genus Bombus, in the family Apidae. There are over 250 known species, existing primarily in the Northern Hemisphere although they are common in New Zealand and in the Australian state of Tasmania.Bumble bees are social insects that are characterised by black...

 and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...

 nests, albeit to a lesser extent. In northern Europe the wax moth is regarded as the most serious parasitoid of the bumblebee, and is found only in bumblebee nests. In some areas in southern England as many as eighty percent of nests can be destroyed. Other parasitic larvae are known to prey upon cicada
Cicada
A cicada is an insect of the order Hemiptera, suborder Auchenorrhyncha , in the superfamily Cicadoidea, with large eyes wide apart on the head and usually transparent, well-veined wings. There are about 2,500 species of cicada around the world, and many of them remain unclassified...

s and leaf hoppers.


File: Parasitism_in_Gypsy_moths.png|thumb|center|500px| The different parasitoids affecting the Gypsy Moth (Lymantaria dispar). What stage they affect and eventually kill and its duration are denoted by arrows.
rect 11 156 222 176 Brachymeria intermedia
rect 11 222 179 182 Coccygomimus instigator
rect 3 194 664 669 Compsilura concinnata
rect 3 686 188 701 Parasetigena silvestris
Parasetigena
Parasetigena is a genus of flies in the family Tachinidae.-Species:*P. amurensis *P. bicolor *P. silvestris *P. takaoi...


rect 231 756 401 774 Blepharipa pratensis
Blepharipa
Blepharipa is a genus of flies in the family Tachinidae.-Species:*B. albocinta *B. carbonata *B. chaetoparafacialis Chao, 1982*B. fimbriata *B. fusiformis...


rect 568 777 833 794 Aphantorhaphopsis samerensis
rect 574 797 779 816 Glyptapanteles liparidis
Glyptapanteles
Glyptapanteles is a genus of endoparasitoid wasp found in Central and North America. The larvae of the members of Glyptapanteles sp. are distinguished by their ability to manipulate their hosts into serving as bodyguards.-Reproduction:...


rect 573 821 766 837 Meteorus pulchricornis
rect 779 541 963 564 Cotesia melanoscelus
rect 783 569 998 592 Glyptapanteles porthetriae
Glyptapanteles
Glyptapanteles is a genus of endoparasitoid wasp found in Central and North America. The larvae of the members of Glyptapanteles sp. are distinguished by their ability to manipulate their hosts into serving as bodyguards.-Reproduction:...


rect 783 596 968 614 Hyposoter tricoloripes
rect 783 619 961 637 Phobocampe disparis
rect 783 131 934 157 Anastatus disparis
desc bottom-left


In reverse, moths and butterflies may be subject to parasitic wasp
Parasitic wasp
The term parasitoid wasp refers to a large evolutionary grade of hymenopteran superfamilies, mainly in the Apocrita. They are primarily parasitoids of other animals, mostly other arthropods...

s and flies
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...

, which may lay eggs on the caterpillars, which hatch and feed inside its body resulting in death. Although, in a form of parasitism called idiobiont, the adult paralyzes the host, so as not to kill it but for it to live as long as possible, in order for the parasitic larvae to benefit the most. Another form of parasitism, is koinobiont, where the species live off their host while inside or endoparasitic. These parasites live inside the host caterpillar throughout its life cycle, or may affect it later on as an adult. In other orders, koinobionts include flies, a majority of coleopteran
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...

, and many hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

n parasitoids. Some species may be subject to a variety of parasites, such as the Gypsy moth
Gypsy moth
The gypsy moth, Lymantria dispar, is a moth in the family Lymantriidae of Eurasian origin. Originally ranging from Europe to Asia, it was introduced to North America in the late 1860s and has been expanding its range ever since...

 (Lymantaria dispar), which is attacked by a series of 13 species, in 6 different taxa throughout its life cycle.

In response to a parsitoid egg or larvae in the caterpillar's body, the plasmatocyte
Hemocyte
A hemocyte is a cell that plays a role in the immune system of invertebrates. It is found within the hemolymph.Hemocytes are phagocytes of invertebrates....

s, or simply the host's cells can form a multilayered capsule that eventually cause the endoparasite to asphyxiate, and die. The process is called encapsulation, and is one of the caterpillar's only means of defense against parasitoids.

Other biological interactions


A few species of Lepidoptera are secondary consumers, or predator
Predation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...

s. These species typically prey upon the eggs of other insects, aphids, scale insects, or ant larvae. Some caterpillars are cannibals, and others prey on caterpillars of other species (e. g. Hawaiian Eupithecia
Eupithecia
Eupithecia is a large genus of moths of the family Geometridae. There are hundreds of described species, found in all parts of the world , and new species are discovered on a regular basis....

). Those of the 15 species in Eupithecia
Eupithecia
Eupithecia is a large genus of moths of the family Geometridae. There are hundreds of described species, found in all parts of the world , and new species are discovered on a regular basis....

 that mirror inchworms, are the only known species of butterflies and moths that are ambush predators. There are 4 known species that eat snails. For example, the Hawai'ian caterpillar, (H. molluscivora
Hyposmocoma molluscivora
Hyposmocoma molluscivora is a Hawaiian moth whose larvae are predators, capturing snails in their silk, much like a hunting spider's web, and then crawling inside the snail's shell to eat it alive...

), uses silk traps, in a manner similar to that of spiders to capture certain species of snails (typically Tornatellides
Tornatellides
Tornatellides is a genus of minute, air-breathing land snails, terrestrial gastropod mollusks, or micromolluscs in the family Achatinellidae.-Predators:These very small snails are preyed upon by caterpillars of the moth Hyposmocoma molluscivora....

).

Larvae of some species of moths of Tineidae
Tineidae
Tineidae is a family of moths in the order Lepidoptera. Collectively, they are known as fungus moths or tineid moths. The family contains considerably more than 3,000 species in over 300 genera. Most of the tineid moths are small or medium-sized, with wings held roofwise over the body at rest...

, Gelechioidae and Noctuidae
Noctuidae
The Noctuidae or owlet moths are a family of robustly-built moths that includes more than 35,000 known species out of possibly 100,000 total, in more than 4,200 genera. They constitute the largest family in the Lepidoptera....

, besides others, feed on detritus
Detritus
Detritus is a biological term used to describe dead or waste organic material.Detritus may also refer to:* Detritus , a geological term used to describe the particles of rock produced by weathering...

, or organic material that is not living, such as fallen leaves and fruit, fungi, and animal products and turn it into humus. Well known species include the cloth moth
Tineidae
Tineidae is a family of moths in the order Lepidoptera. Collectively, they are known as fungus moths or tineid moths. The family contains considerably more than 3,000 species in over 300 genera. Most of the tineid moths are small or medium-sized, with wings held roofwise over the body at rest...

s (Tineola bisselliella, T. pellionella
Tinea pellionella
The Case-bearing Clothes Moth is a species of tineoid moth. It belongs to the fungus moth family , and therein to the nominate subfamily Tineinae...

, and T. tapetzella
Trichophaga tapetzella
The Tapestry Moth or Carpet Moth is a moth of the Tineidae family. It is found worldwide.The wingspan is 14–18 mm...

), which feed on detritus containing keratin
Keratin
Keratin refers to a family of fibrous structural proteins. Keratin is the key of structural material making up the outer layer of human skin. It is also the key structural component of hair and nails...

, including hair
Hair
Hair is a filamentous biomaterial, that grows from follicles found in the dermis. Found exclusively in mammals, hair is one of the defining characteristics of the mammalian class....

, feather
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...

s, cobweb
Spider web
A spider web, spiderweb, spider's web or cobweb is a device built by a spider out of proteinaceous spider silk extruded from its spinnerets....

s, bird nest
Bird nest
A bird nest is the spot in which a bird lays and incubates its eggs and raises its young. Although the term popularly refers to a specific structure made by the bird itself—such as the grassy cup nest of the American Robin or Eurasian Blackbird, or the elaborately woven hanging nest of the...

s (particularly of Domestic Pigeon
Domestic Pigeon
The Domestic Pigeon was derived from the Rock Pigeon. The Rock Pigeon is the world's oldest domesticated bird. Mesopotamian cuneiform tablets mention the domestication of pigeons more than 5,000 years ago, as do Egyptian hieroglyphics.Research suggests that domestication of pigeons was as early as...

s, Columba livia domestica) and fruits or vegetables. These species are important to ecosystems as they remove substances that would otherwise take a long time to decompose.

History of study


Linnaeus in Systema Naturae
Systema Naturae
The book was one of the major works of the Swedish botanist, zoologist and physician Carolus Linnaeus. The first edition was published in 1735...

(1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx and Phalaena
Phalaena
Phalaena is an obsolete genus Lepidoptera used by Carl Linnaeus to house most moths.Phalaena was one of three genera used by Linnaeus to cover all Lepidoptera. Papilio included all butterflies at that time, Sphinx included all hawk moths, and Phalaena included all the remaining moths...

, with seven subgroups in Phalaena. These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Michael Denis
Michael Denis
Johann Nepomuk Cosmas Michael Denis, also: SinedSined is an anagram of Denis. the Bard, was an Austrian poet, bibliographer, and lepidopterist....

 & Ignaz Schiffermüller
Ignaz Schiffermüller
Ignaz Schiffermüller was an Austrian naturalist mainly interested in Lepidoptera....

 (1775), Johan Christian Fabricius
Johan Christian Fabricius
Johan Christian Fabricius was a Danish zoologist, specialising in "Insecta", which at that time included all arthropods: insects, arachnids, crustaceans and others...

 (1775) and Pierre André Latreille
Pierre André Latreille
Pierre André Latreille was a French zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained his freedom after recognising a rare species he found in the prison, Necrobia ruficollis...

 (1796). Jacob Hübner
Jacob Hübner
Jacob Hübner was a German entomologist. He was the author of Sammlung Europäischer Schmetterlinge , a founding work of entomology.-Scientific career:...

 described many genera, and the Lepidopteran genera were catalogued by Ferdinand Ochsenheimer
Ferdinand Ochsenheimer
Ferdinand Ochsenheimer was a German actor and entomologist .-Life:Ochsenheimer was born and brought up in in Mainz, Germany, and began to show an interest in butterflies and moths in his early youth...

 and Georg Friedrich Treitschke
Georg Friedrich Treitschke
Georg Friedrich Treitschke was a German librettist, translator and lepidopterist....

 in a series of volumes on the Lepidopteran fauna of Europe published between 1807 and 1835. Gottlieb August Wilhelm Herrich-Schäffer
Gottlieb August Wilhelm Herrich-Schäffer
Dr Gottlieb August Wilhelm Herrich-Schäffer was a German entomologist and physician. He was born, and died, in Regensburg. Herrich-Schäffer studied and collected particularly butterflies and moths...

 (several volumes, 1843–1856), and Edward Meyrick
Edward Meyrick
Edward Meyrick FRS was an English schoolmaster and amateur entomologist. He was an expert on microlepidoptera and some consider him one of the founders of modern Microlepidoptera systematics....

 (1895) based their classifications primarily on wing venation. Sir George Francis Hampson
George Francis Hampson
Sir George Francis Hampson, 10th Baronet was a British entomologist.Hampson studied at Charterhouse School and Exeter College, Oxford. He travelled to India to become a tea-planter in the Nilgiri Hills of the Madras presidency , where he became interested in moths and butterflies...

 worked on the 'Microlepidoptera' during this period and Philipp Christoph Zeller
Philipp Christoph Zeller
Philipp Christoph Zeller was a German entomologist.Zeller was born at Steinheim Württemberg, two miles from Marbach, the birthplace of Schiller. The family moved to Frankfurt where Philip went to the gymnasium where natural history was not taught. Instead, helped by Alois Metzner, he taught...

 published The Natural History of the Tineinae also on Microlepidoptera (1855).

Among the first entomologists to study fossil insects and their evolution was Samuel Hubbard Scudder
Samuel Hubbard Scudder
Samuel Hubbard Scudder was an American entomologist and palaeontologist.Born in Boston, Massachusetts, Scudder may be most widely known for his essay on the importance of first-hand, careful observation in the natural sciences...

 (1837–1911), who worked on butterflies. He published a study of the Florissant deposits of Colorado, including the exceptionally preserved Prodryas persephone
Prodryas
Prodryas persephone is an extinct butterfly, known from a single specimen from Eocene rocks. It was the first fossil butterfly to be found in North America, and is unusually well preserved...

. Andreas V. Martynov (1879–1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.

Major contributions in the 20th century included the creation of the monotrysia and ditrysia (based on female genital structure) by Borner in 1925 and 1939. Willi Hennig
Willi Hennig
Emil Hans Willi Hennig was a German biologist who is considered the founder of phylogenetic systematics, also known as cladistics. With his works on evolution and systematics he revolutionised the view of the natural order of beings...

 (1913–1976) developed the cladistic
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among monotrysia
Monotrysia
The Monotrysia is a group of insects in the Lepidopteran order which is not currently considered to be a natural group or clade. The group contains only moths and most of these are small and are relatively understudied in many regions of the world...

n families and Kristensen worked more generally on insect phylogeny and higher Lepidoptera too. While it is often found that DNA-based phylogenies differ from those based on morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

, this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.

Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not monophyletic
Monophyly
In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...

: Microlepidotera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.

Fossil record


The fossil record for Lepidoptera is lacking in comparison to other winged species, and tending not to be as common as some other insects in the habitats that are most conducive to fossilization, such as lakes and ponds, and their juvenile stage has only the head capsule as a hard part that might be preserved. The location and abundance of the most common moth species are indicative that mass migrations of moths occurred over the Palaeogene North Sea
North Sea
In the southwest, beyond the Straits of Dover, the North Sea becomes the English Channel connecting to the Atlantic Ocean. In the east, it connects to the Baltic Sea via the Skagerrak and Kattegat, narrow straits that separate Denmark from Norway and Sweden respectively...

, which is why there is a serious lack of moth fossils. Yet there are fossils, some preserved in amber and some in very fine sediments. Leaf mines
Leaf miner
Leaf miner is a term used to describe the larvae of many different species of insect which live in and eat the leaf tissue of plants. The vast majority of leaf-mining insects are moths , sawflies and flies , though some beetles and wasps also exhibit this behavior.Like Woodboring beetles, leaf...

 are also seen in fossil leaves, although the interpretation of them is tricky.

Putative fossil stem group representatives of Amphiesmenoptera (the clade comprising Trichoptera and Lepidoptera) are known from the Triassic
Triassic
The Triassic is a geologic period and system that extends from about 250 to 200 Mya . As the first period of the Mesozoic Era, the Triassic follows the Permian and is followed by the Jurassic. Both the start and end of the Triassic are marked by major extinction events...

. The earliest known fossil lepidopteran is Archaeolepis mane
Archaeolepis
Archaeolepis mane is the earliest known Lepidopteran fossil. It dates from the Lower Jurassic and according to Grimaldi & Engel a recent re-examination of the specimen has given additional support to its ordinal placement...

from the Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to  Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...

, about in Dorset
Dorset
Dorset , is a county in South West England on the English Channel coast. The county town is Dorchester which is situated in the south. The Hampshire towns of Bournemouth and Christchurch joined the county with the reorganisation of local government in 1974...

, UK. The fossil belongs to a small primitive moth-like species, and its wings are showing scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared with Trichoptera
Trichoptera
The caddisflies are an order, Trichoptera, of insects with approximately 12,000 described species. Also called sedge-flies or rail-flies, they are small moth-like insects having two pairs of hairy membranous wings...

 (Caddisflies). Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

, which all belong to primitive moth-like families. Many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber
Baltic amber
The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite, with about 80% of the world's known amber found there. It dates from 44 million years ago...

. The oldest genuine butterflies of the superfamily Papilionoidea have been found in the Paleocene
Paleocene
The Paleocene or Palaeocene, the "early recent", is a geologic epoch that lasted from about . It is the first epoch of the Palaeogene Period in the modern Cenozoic Era...

 MoClay
MoClay
The Moclay strata are formally designated as the Fur Formation.It is a diatomitic sediment of Lower Eocene Epoch, c. 56-54,5 Ma) age which crops out in the Limfjord region of Denmark from Silstrup via Mors and Fur to Ertebølle, and can be seen in many cliffs and quarries in the area...

 or Fur Formation
Fur Formation
Essentially the Fur Formation is defined as a clayey diatomite with a lage number of volcanic ash layers.The Fur Formation is a marine geological formation of Ypresian age which crops out in the Limfjord region of Denmark from Silstrup via Mors and Fur to Ertebølle, and can be seen in many cliffs...

 of Denmark. The best preserved fossil lepidopteran is the Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...

 Prodryas persephone
Prodryas
Prodryas persephone is an extinct butterfly, known from a single specimen from Eocene rocks. It was the first fossil butterfly to be found in North America, and is unusually well preserved...

from the Florissant Fossil Beds
Florissant Fossil Beds National Monument
Florissant Fossil Beds National Monument is a United States National Monument in Teller County, Colorado, that is noted for its fossils. It is located in a mountain valley just west of Pikes Peak and holds spectacular remnants of prehistoric life...

.

Phylogeny


Lepidoptera and Trichoptera
Trichoptera
The caddisflies are an order, Trichoptera, of insects with approximately 12,000 described species. Also called sedge-flies or rail-flies, they are small moth-like insects having two pairs of hairy membranous wings...

 (caddisflies) are more closely related than any other taxa, sharing many similarities that are lacking in other insect orders; for example the females of both orders are heterogametic
ZW sex-determination system
The ZW sex-determination system is a system that determines the sex of offspring in birds, some fish and crustaceans such as the giant river prawn, some insects , and some reptiles, including Komodo dragons...

, meaning they have two different sex chromosomes
Gamete
A gamete is a cell that fuses with another cell during fertilization in organisms that reproduce sexually...

, whereas in most species the males are heterogametic and the females have two identical sex chromosomes. The adults in both orders display a particular wing venation pattern on their forewings. The larvae of both orders have mouth structures and gland with which they make and manipulate silk
Silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...

. Willi Hennig
Willi Hennig
Emil Hans Willi Hennig was a German biologist who is considered the founder of phylogenetic systematics, also known as cladistics. With his works on evolution and systematics he revolutionised the view of the natural order of beings...

 grouped the two sister orders into the Amphiesmenoptera
Amphiesmenoptera
Amphiesmenoptera is an insect superorder, established by Willi Hennig in his revision of insect taxonomy for two sister orders: Lepidoptera and Trichoptera ....

 superorder. This group probably evolved in the Jurassic, having split from the now extinct order Necrotaulidae.

Micropterigidae
Micropterigidae
Micropterigoidea is the superfamily of "mandibulate archaic moths", all placed in the single family Micropterigidae, containing currently about 20 living genera. They are considered the most primitive extant lineage of Lepidoptera ....

, Agathiphagidae and Heterobathmiidae are the oldest and most basal
Basal (phylogenetics)
In phylogenetics, a basal clade is the earliest clade to branch in a larger clade; it appears at the base of a cladogram.A basal group forms an outgroup to the rest of the clade, such as in the following example:...

 lineages of Lepidoptera. The adults of these families do not have the curled tongue or proboscis
Proboscis
A proboscis is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In simpler terms, a proboscis is the straw-like mouth found in several varieties of species.-Etymology:...

, that are found in most members order, but instead have chewing mandibles adapted for a special diet. Micropterigidae larvae feed on leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

, fungi, or liverworts
Marchantiophyta
The Marchantiophyta are a division of bryophyte plants commonly referred to as hepatics or liverworts. Like other bryophytes, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of genetic information....

 (much like the Trichoptera
Trichoptera
The caddisflies are an order, Trichoptera, of insects with approximately 12,000 described species. Also called sedge-flies or rail-flies, they are small moth-like insects having two pairs of hairy membranous wings...

). Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live inside kauri pines and feed on seeds. In Heterobathmiidae the larvae feed on the leaves of Nothofagus
Nothofagus
Nothofagus, also known as the southern beeches, is a genus of 35 species of trees and shrubs native to the temperate oceanic to tropical Southern Hemisphere in southern South America and Australasia...

, the southern beech tree. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon after metamorphosis
Metamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...

.

The Eriocraniidae have a short coiled proboscis in the adult stage, and though they retain their pupal mandibles with which they escaped the cocoon, their mandibles are non-functional thereafter. Most of these non-ditrysian families, are primarily leaf miner
Leaf miner
Leaf miner is a term used to describe the larvae of many different species of insect which live in and eat the leaf tissue of plants. The vast majority of leaf-mining insects are moths , sawflies and flies , though some beetles and wasps also exhibit this behavior.Like Woodboring beetles, leaf...

s in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.

With the evolution of the Ditrysia
Ditrysia
The Ditrysia are a natural group or clade of insects in the Lepidopteran order containing both butterflies and moths. They are so named because the female has two distinct sexual openings: one for mating, and the other for laying eggs .About 98% of described species of Lepidoptera belong to Ditrysia...

 in the mid-Cretaceous, there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is one cloaca
Cloaca
In zoological anatomy, a cloaca is the posterior opening that serves as the only such opening for the intestinal, reproductive, and urinary tracts of certain animal species...

, or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, Gracillarioidea
Gracillarioidea
Gracillarioidea is a large superfamily containing four families of insects in the order Lepidoptera. These generally small moths are miners in plant tissue as caterpillars. There are about 113 described genera distributed worldwide, the most commonly encountered of which are leaf-miners in the...

 and Gelechioidea
Gelechioidea
| name = Curved-horn moths| image = Xylorycta assimilis.jpg| image_width = 240px| image_caption = Adult Xylorycta assimilis of the Xyloryctidae, photographed in Aranda Note prominent "horns" and long antennae| regnum = Animalia| phylum = Arthropoda...

 are mostly leaf miners, but more recent lineages feed externally. In the Tineoidea
Tineoidea
Tineoidea is the superfamily of moths that includes clothes moths, bagworms and relatives. There are six families usually included within it, Eriocottidae, Arrhenophanidae, Lypusidae, Acrolophidae, Tineidae and Psychidae, whose relationships are currently uncertain.The Lypusidae, for example, might...

, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.

The Yponomeutoidea
Yponomeutoidea
Yponomeutoidea is a superfamily of Ermine moths and relatives.-Familiae:*Acrolepiidae*Bedelliidae*Glyphipterigidae*Heliodinidae*Lyonetiidae*Plutellidae*Yponomeutidae*Ypsolophidae- Etymology :...

 is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants. They evolved about the time that flowering plants underwent an expansive adaptive radiation
Adaptive radiation
In evolutionary biology, adaptive radiation is the evolution of ecological and phenotypic diversity within a rapidly multiplying lineage. Starting with a recent single ancestor, this process results in the speciation and phenotypic adaptation of an array of species exhibiting different...

 in the mid-Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

, and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved coevolution or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.

In the so-called "Macrolepidoptera
Macrolepidoptera
Macrolepidoptera is a group within the insect order Lepidoptera. Traditionally used for the larger butterflies and moths as opposed to the "Microlepidoptera", this group is unnatural. However, it seems that by moving some taxa about, a monophyletic Macrolepidoptera can be easily achieved...

", which constitutes about 60% of lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant. Many also have tympanal organ
Tympanal organ
A tympanal organ is a hearing organ in insects, consisting of a membrane stretched across a frame backed by an air sac and associated sensory neurons...

s, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.
The main lineages in the Macrolepidoptera are the Noctuoidea
Noctuoidea
Noctuoidea is the superfamily of noctuid or "owlet" moths, and has the largest number of species described for any Lepidopteran superfamily. Its classification has not yet reached a satisfactory or stable state. The most recent classifications include only four families in the superfamily;...

, Bombycoidea
Bombycoidea
Bombycoidea is a superfamily of moths. It contains the silk moths, emperor moths, sphinx moths and relatives. The Lasiocampoidea are close relatives and sometimes merged in the present group. Their larvae exhibit horns.-Sources:...

, Lasiocampidae
Lasiocampidae
The Lasiocampidae family of moths are also known as eggars, snout moths or lappet moths. There are over 2000 species worldwide, and probably not all have been named or studied....

, Mimallonoidea
Mimallonoidea
Mimallonoidea is the superfamily of sack bearer moths, containing the single family Mimallonidae.-Genera:*Aceclostria*Adalgisa*Aleyda*Alheita*Bedosia*Biterolfa*Cicinnus*Druentica*Eadmuna*Euphaneta...

, Geometroidea
Geometroidea
Geometroidea is the superfamily of geometrid moths in the Lepidoptera. It includes the families Geometridae, Uraniidae, and Sematuridae...

 and Rhopalocera. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a monophyletic group. The Rhopalocera, comprising the Papilionoidea
Papilionoidea
The superfamily Papilionoidea contains all the butterflies except for the skippers, which are classified in superfamily Hesperioidea, and the moth-like Hedyloidea....

 (butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved. There is quite a good fossil record for this group, with the oldest skipper dating from .

Taxonomy


Taxonomy is the classification of species in selected taxa, the process of naming being called nomenclature
Biological classification
Biological classification, or scientific classification in biology, is a method to group and categorize organisms by biological type, such as genus or species. Biological classification is part of scientific taxonomy....

. There are over 120 families in lepidoptera, in 45 to 48 superfamilies. Lepidoptera have always been, historically, classified in five suborders, one of which is of primitive moths that never lost the morphological features of its ancestors. The rest of the moths and butterflies make up ninety-eight percent of the other taxa, making Ditrysia
Ditrysia
The Ditrysia are a natural group or clade of insects in the Lepidopteran order containing both butterflies and moths. They are so named because the female has two distinct sexual openings: one for mating, and the other for laying eggs .About 98% of described species of Lepidoptera belong to Ditrysia...

. More recently, new findings of new taxa and larvae and pupa have aided in detailing the relationships of primitive taxa, phylogenetic analysis showing the primitive lineages to be paraphyletic
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...

 compared to the rest of Lepidoptera lineages. Recently lepidopterists have abandoned clades like suborders, and those between orders and superfamilies.
  • Zeugloptera is a clade with Micropterigoidea being its only family. Species of Micropterigoidea are practically living fossils, being one of the most primitive lepidopteran species, still retaining mandible mouthparts, unlike other clades of butterflies and moths. About 120 species are known worldwide, with more than half the species in the genus Micropteryx in the Paleartic region. There are only 2 known in North America (Epimartyria
    Epimartyria
    Epimartyria is a genus of small primitive metallic moths in the insect order Lepidoptera within the family Micropterigidae.-Species:*Epimartyria auricrinella Walsingham, 1898*Epimartyria pardella...

    ), with many more being found Asia and the southwest Pacific, particularly New Zealand with about 50 species.

  • Glossata
    Glossata
    Glossata is the suborder of the insect order Lepidoptera that includes all the superfamilies of moths and butterflies that have a coilable proboscis. ....

     contains a majority of the species, with the most obvious difference is non-functioning mandibles, and elongated maxillary galeae or the proboscis. The basal clades still retaining some of the ancestral features of the wings such as similarly shaped fore- and hindwings with relatively complete venation. Glossata also contains the division Ditrysia
    Ditrysia
    The Ditrysia are a natural group or clade of insects in the Lepidopteran order containing both butterflies and moths. They are so named because the female has two distinct sexual openings: one for mating, and the other for laying eggs .About 98% of described species of Lepidoptera belong to Ditrysia...

    , which contains 98% of all described species in Lepidoptera.

  • Aglossata it is the second most primitive lineage
    Lineage (evolution)
    An evolutionary lineage is a sequence of species, that form a line of descent, each new species the direct result of speciation from an immediate ancestral species. Lineages are subsets of the evolutionary tree of life. Lineages are often determined by the techniques of molecular systematics.-...

     of lepidoptera; being first described in 1952 by Lionel Jack Dumbleton
    Lionel Jack Dumbleton
    Lionel Jack Dumbleton was a New Zealand entomologist. He was born in Hampden, New Zealand and was a founding member of the Entomological Society of New Zealand....

    . Agathiphagidae and Heterobathmiidae are the only families in Aglossata. Agathiphagidae only contains about 2 species in its genus Agathiphaga
    Agathiphaga
    Agathiphaga is a genus of moths in the family Agathiphagidae, known as kauri moths. This caddis fly-like lineage of primitive moths was first reported by Lionel Jack Dumbleton in 1952, as a new genus of Micropterigidae....

    .. Agathiphaga queenslandensis
    Agathiphaga queenslandensis
    Agathiphaga queenslandensis is a moth of the Agathiphagidae family. It is found along the north-eastern coast of Queensland, Australia.The wingspan is about 13 mm. Adults are night active. The forewings are unicolorous, without spots or pattern....

    and Agathiphaga vitiensis
    Agathiphaga vitiensis
    Agathiphaga vitiensis is a moth of the Agathiphagidae family. It is found from Fiji to Vanuatu and Solomon Islands.The length of the forewings is about 4 mm...

    , being found along the north-eastern coast of Queensland
    Queensland
    Queensland is a state of Australia, occupying the north-eastern section of the mainland continent. It is bordered by the Northern Territory, South Australia and New South Wales to the west, south-west and south respectively. To the east, Queensland is bordered by the Coral Sea and Pacific Ocean...

    , Australia
    Australia
    Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

    , and in Fiji
    Fiji
    Fiji , officially the Republic of Fiji , is an island nation in Melanesia in the South Pacific Ocean about northeast of New Zealand's North Island...

     to Vanuatu
    Vanuatu
    Vanuatu , officially the Republic of Vanuatu , is an island nation located in the South Pacific Ocean. The archipelago, which is of volcanic origin, is some east of northern Australia, northeast of New Caledonia, west of Fiji, and southeast of the Solomon Islands, near New Guinea.Vanuatu was...

     and the Solomon Islands
    Solomon Islands
    Solomon Islands is a sovereign state in Oceania, east of Papua New Guinea, consisting of nearly one thousand islands. It covers a land mass of . The capital, Honiara, is located on the island of Guadalcanal...

    , respectively.

  • Heterobathmiina was first described by Kristensen and Nielsen in 1979. There are about 10 species, which are day-flying, metallic moths, confined to southern South America, the adults eat the pollen of Nothofagus
    Nothofagus
    Nothofagus, also known as the southern beeches, is a genus of 35 species of trees and shrubs native to the temperate oceanic to tropical Southern Hemisphere in southern South America and Australasia...

    or Southern Beech and the larvae mine the leaves.

In culture


Artistic depictions of butterflies have been used in many cultures including as early as 3500 years ago, in Egyptian hieroglyphs. Today, butterflies are widely used in various objects of art and jewelry: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and in some mixed media artworks and furnishings. Butterflies have also inspired the "butterfly fairy
Fairy
A fairy is a type of mythical being or legendary creature, a form of spirit, often described as metaphysical, supernatural or preternatural.Fairies resemble various beings of other mythologies, though even folklore that uses the term...

" as an art and fictional character, including in the Barbie Mariposa
Barbie Mariposa
Barbie Mariposa is a 2008 direct to video computer animated Barbie film which was released on February 26, 2008. This film is a part of the Barbie "Fairytopia" series, but is not a canon sequel to the previous films...

film.

In many cultures the soul of a dead person is associated with the butterfly. As in Ancient Greece
Ancient Greece
Ancient Greece is a civilization belonging to a period of Greek history that lasted from the Archaic period of the 8th to 6th centuries BC to the end of antiquity. Immediately following this period was the beginning of the Early Middle Ages and the Byzantine era. Included in Ancient Greece is the...

, where the word for butterfly ψυχή (psyche) also means soul and breath. In Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

, as in Ancient Greece, the word for "butterfly" papillio was associated with the soul of the dead. The skull-like marking on the thorax of the Death's-head Hawkmoth
Death's-head Hawkmoth
The name Death's-head Hawkmoth refers to any one of the three species of moth in the genus Acherontia. The former species is primarily found in Europe, the latter two are Asian, and most uses of the common name refer to the European species...

 has helped these moths, particularly A. atropos, earn a negative reputation, such as associations with the supernatural and evil. The moth has been prominently featured in art and movies such as (by Buñuel
Luis Buñuel
Luis Buñuel Portolés was a Spanish-born filmmaker — later a naturalized citizen of Mexico — who worked in Spain, Mexico, France and the US..-Early years:...

 and Dalí
Salvador Dalí
Salvador Domènec Felip Jacint Dalí i Domènech, Marquis de Púbol , commonly known as Salvador Dalí , was a prominent Spanish Catalan surrealist painter born in Figueres,Spain....

) and The Silence of the Lambs, and in the artwork of the Japanese metal band Sigh
Sigh (band)
is a Japanese extreme metal band from Tokyo, formed in 1990. They are credited as being one of the first Japanese black metal bands, when the majority of black metal in early 1990s came from Scandinavia. They gradually shifted from a more traditional black/thrash metal sound, to a more...

's album Hail Horror Hail
Hail Horror Hail
Hail Horror Hail is an album by the band Sigh. It was released by Cacophonous Records originally in 1997.This is the first full-length album of Sigh's to explore their more well known avant-garde/experimental sound...

. According to Kwaidan: Stories and Studies of Strange Things
Kwaidan: Stories and Studies of Strange Things
, often shortened to Kwaidan, is a book by Lafcadio Hearn that features several Japanese ghost stories and a brief non-fiction study on insects...

, by Lafcadio Hearn
Lafcadio Hearn
Patrick Lafcadio Hearn , known also by the Japanese name , was an international writer, known best for his books about Japan, especially his collections of Japanese legends and ghost stories, such as Kwaidan: Stories and Studies of Strange Things...

, a butterfly was seen in Japan as the personification of a person's soul; whether they be living, dying, or already dead. One Japanese superstition says that if a butterfly enters your guestroom and perches behind the bamboo
Bamboo
Bamboo is a group of perennial evergreens in the true grass family Poaceae, subfamily Bambusoideae, tribe Bambuseae. Giant bamboos are the largest members of the grass family....

 screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad omen
Omen
An omen is a phenomenon that is believed to foretell the future, often signifying the advent of change...

s. When Taira no Masakado
Taira no Masakado
was a samurai in the Heian period of Japan, who led one of the largest insurgent forces in the period against the central government of Kyoto.-History:...

 was secretly preparing for his famous revolt, there appeared in Kyoto
Kyoto
is a city in the central part of the island of Honshū, Japan. It has a population close to 1.5 million. Formerly the imperial capital of Japan, it is now the capital of Kyoto Prefecture, as well as a major part of the Osaka-Kobe-Kyoto metropolitan area.-History:...

 so vast a swarm of butterflies that the people were frightened—thinking the apparition to be a portent of coming evil.

In the ancient Mesoamerica
Mesoamerica
Mesoamerica is a region and culture area in the Americas, extending approximately from central Mexico to Belize, Guatemala, El Salvador, Honduras, Nicaragua, and Costa Rica, within which a number of pre-Columbian societies flourished before the Spanish colonization of the Americas in the 15th and...

n city of Teotihuacan
Teotihuacan
Teotihuacan – also written Teotihuacán, with a Spanish orthographic accent on the last syllable – is an enormous archaeological site in the Basin of Mexico, just 30 miles northeast of Mexico City, containing some of the largest pyramidal structures built in the pre-Columbian Americas...

, the brilliantly colored image of the butterfly was carved into many temples, buildings, jewelry, and emblazoned on incense burners
Censer
Censers are any type of vessels made for burning incense. These vessels vary greatly in size, form, and material of construction. They may consist of simple earthenware bowls or fire pots to intricately carved silver or gold vessels, small table top objects a few centimetres tall to as many as...

 in particular. The butterfly was sometimes depicted with the maw of a jaguar
Jaguar
The jaguar is a big cat, a feline in the Panthera genus, and is the only Panthera species found in the Americas. The jaguar is the third-largest feline after the tiger and the lion, and the largest in the Western Hemisphere. The jaguar's present range extends from Southern United States and Mexico...

 and some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies to fire
Fire
Fire is the rapid oxidation of a material in the chemical process of combustion, releasing heat, light, and various reaction products. Slower oxidative processes like rusting or digestion are not included by this definition....

 and warfare persisted through to the Aztec civilization and evidence of similar jaguar-butterfly images has been found among the Zapotec
Zapotec civilization
The Zapotec civilization was an indigenous pre-Columbian civilization that flourished in the Valley of Oaxaca of southern Mesoamerica. Archaeological evidence shows their culture goes back at least 2500 years...

, and Maya civilization
Maya civilization
The Maya is a Mesoamerican civilization, noted for the only known fully developed written language of the pre-Columbian Americas, as well as for its art, architecture, and mathematical and astronomical systems. Initially established during the Pre-Classic period The Maya is a Mesoamerican...

s.

As pests


The larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

e of many Lepidopteran species are major pests in agriculture
Agriculture
Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...

. Some of the major pests include Tortricidae
Tortricidae
Tortricidae is a family of moths, commonly known as tortrix moths, in the order Lepidoptera. Tortricidae is a large family with over 9,400 species described, and is the sole member of the superfamily Tortricoidea. Many of these are economically important pests. Olethreutidae is a junior synonym...

, Noctuidae
Noctuidae
The Noctuidae or owlet moths are a family of robustly-built moths that includes more than 35,000 known species out of possibly 100,000 total, in more than 4,200 genera. They constitute the largest family in the Lepidoptera....

, and Pyralidae
Pyralidae
The Pyralidae or snout moths are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many classifications, the grass moths are included in the Pyralidae as a subfamily, making the combined group one of the largest families in the Lepidoptera...

. The larvae of the Noctuidae genus Spodoptera
Spodoptera
Spodoptera is a genus of moths of the Noctuidae family. Many are known as pest insects. The larvae are sometimes called armyworms.Species include:* Spodoptera abyssinia Guenée, 1852...

(armyworms) and Helicoverpa
Helicoverpa
Helicoverpa is a genus of moth in the Noctuidae family.-Extant species:* Helicoverpa armigera - Cotton Bollworm Hübner, 1805* Helicoverpa assulta * Helicoverpa atacamae Hardwick, 1965...

(corn earworm) can cause extensive damage to certain crops. Helicoverpa zea
Helicoverpa zea
The larva of the moth Helicoverpa zea is a major agricultural pest. It can feed on many different plants during the larval stage. Accordingly, the species has been given many different common names. When the larva consumes cotton, it is known as the cotton bollworm...

larvae (cotton bollworms or tomato fruitworms) are polyphagous, meaning they eat a variety of crops, including tomato
Tomato
The word "tomato" may refer to the plant or the edible, typically red, fruit which it bears. Originating in South America, the tomato was spread around the world following the Spanish colonization of the Americas, and its many varieties are now widely grown, often in greenhouses in cooler...

es and cotton
Cotton
Cotton is a soft, fluffy staple fiber that grows in a boll, or protective capsule, around the seeds of cotton plants of the genus Gossypium. The fiber is almost pure cellulose. The botanical purpose of cotton fiber is to aid in seed dispersal....

.

Butterflies and moths are one of the largest taxa to solely feed and be dependent on living plants, in terms of the number of species, and they are in many ecosystems make up the largest biomass to do so. In many species, the female may produce anywhere from 200 to 600 eggs, while in some others it may go as high as 30,000 eggs in one day. This creates many problems for agriculture, where many caterpillars can mow down acres of vegetation. Some reports estimate that there have been over 80,000 caterpillars of several different taxa feeding on a single oak tree. In some cases, phytophagous larvae can lead to the destruction of entire trees in relatively short periods of time.

Ecological ways of removing pest lepidoptera species are becoming more economically viable, as research has shown ways like introducing parasitic wasp and flies. For example Sarcophaga aldrichi
Sarcophaga aldrichi
The friendly fly or large flesh fly, Sarcophaga aldrichi, is a fly that is a parasitoid of the forest tent caterpillar. It strongly resembles the house fly but is in a different family, the Sarcophagidae, or flesh-flies.. It is a little larger than the house fly, and has the same three black...

, which the larvae feed upon the larvae of the Forest Tent Caterpillar Moth. Pesticides can affect other species other than the species they are targeted to eliminate, damaging the natural ecosystem. Another good biological pest control method is the use of pheromone trap
Pheromone trap
A pheromone trap is a type of insect trap that uses pheromones to lure insects. Sex pheromones and aggregating pheromones are the most common types used. A pheromone-impregnated lure is encased in a conventional trap such as a Delta trap, water-pan trap, or funnel trap.-Sensitivity:Pheromone traps...

s. A pheromone trap is a type of insect trap
Insect trap
Insect traps are used to monitor or directly reduce insect populations. They typically use food, visual lures, chemical attractants and pheromones as bait and are installed so that they do not injure other animals or humans or result in residues in foods or feeds. Visual lures use light, bright...

 that uses pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...

s to lure insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s. Sex pheromones and aggregating pheromones are the most common types used. A pheromone-impregnated lure is encased in a conventional trap such as a Delta trap, water-pan trap, or funnel trap.

Species of moths that are detrivores would naturally eat detritus
Detritus
Detritus is a biological term used to describe dead or waste organic material.Detritus may also refer to:* Detritus , a geological term used to describe the particles of rock produced by weathering...

 containing keratin
Keratin
Keratin refers to a family of fibrous structural proteins. Keratin is the key of structural material making up the outer layer of human skin. It is also the key structural component of hair and nails...

, such as hair
Hair
Hair is a filamentous biomaterial, that grows from follicles found in the dermis. Found exclusively in mammals, hair is one of the defining characteristics of the mammalian class....

s or feather
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...

s. Well known species are cloth moth
Tineidae
Tineidae is a family of moths in the order Lepidoptera. Collectively, they are known as fungus moths or tineid moths. The family contains considerably more than 3,000 species in over 300 genera. Most of the tineid moths are small or medium-sized, with wings held roofwise over the body at rest...

s (T. bisselliella, T. pellionella
Tinea pellionella
The Case-bearing Clothes Moth is a species of tineoid moth. It belongs to the fungus moth family , and therein to the nominate subfamily Tineinae...

, and T. tapetzella
Trichophaga tapetzella
The Tapestry Moth or Carpet Moth is a moth of the Tineidae family. It is found worldwide.The wingspan is 14–18 mm...

), feeding on foodstuffs that people find economically important, such cotton
Cotton
Cotton is a soft, fluffy staple fiber that grows in a boll, or protective capsule, around the seeds of cotton plants of the genus Gossypium. The fiber is almost pure cellulose. The botanical purpose of cotton fiber is to aid in seed dispersal....

, linen
Linen
Linen is a textile made from the fibers of the flax plant, Linum usitatissimum. Linen is labor-intensive to manufacture, but when it is made into garments, it is valued for its exceptional coolness and freshness in hot weather....

, silk
Silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...

 and wool fabric
Fabric
A fabric is a textile material, short for "textile fabric".Fabric may also refer to:*Fabric , the spatial and geometric configuration of elements within a rock*Fabric , a nightclub in London, England...

s as well as fur
Fur
Fur is a synonym for hair, used more in reference to non-human animals, usually mammals; particularly those with extensives body hair coverage. The term is sometimes used to refer to the body hair of an animal as a complete coat, also known as the "pelage". Fur is also used to refer to animal...

s; furthermore they have been found on shed feather
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...

s and hair
Hair
Hair is a filamentous biomaterial, that grows from follicles found in the dermis. Found exclusively in mammals, hair is one of the defining characteristics of the mammalian class....

, bran
Bran
Bran is the hard outer layer of grain and consists of combined aleurone and pericarp. Along with germ, it is an integral part of whole grains, and is often produced as a by-product of milling in the production of refined grains. When bran is removed from grains, the grains lose a portion of their...

, semolina
Semolina
Semolina is the coarse, purified wheat middlings of durum wheat used in making pasta, and also used for breakfast cereals and puddings. Semolina is also used to designate coarse middlings from other varieties of wheat, and from other grains such as rice and corn.-Name:The term semolina derives from...

 and flour
Flour
Flour is a powder which is made by grinding cereal grains, other seeds or roots . It is the main ingredient of bread, which is a staple food for many cultures, making the availability of adequate supplies of flour a major economic and political issue at various times throughout history...

 (possibly preferring wheat
Wheat
Wheat is a cereal grain, originally from the Levant region of the Near East, but now cultivated worldwide. In 2007 world production of wheat was 607 million tons, making it the third most-produced cereal after maize and rice...

 flour), biscuit
Biscuit
A biscuit is a baked, edible, and commonly flour-based product. The term is used to apply to two distinctly different products in North America and the Commonwealth Nations....

s, casein
Casein
Casein is the name for a family of related phosphoprotein proteins . These proteins are commonly found in mammalian milk, making up 80% of the proteins in cow milk and between 60% and 65% of the proteins in human milk....

, and insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

 specimens in museum
Museum
A museum is an institution that cares for a collection of artifacts and other objects of scientific, artistic, cultural, or historical importance and makes them available for public viewing through exhibits that may be permanent or temporary. Most large museums are located in major cities...

s.

As beneficial


Even though most butterflies and moths affect the economy negatively, some species are a valuable economic resource. The most prominent example is that of the Domesticated silkworm moth
Bombyx mori
The silkworm is the larva or caterpillar of the domesticated silkmoth, Bombyx mori . It is an economically important insect, being a primary producer of silk...

 (Bombyx mori), the larvae of which make their cocoons out of silk
Silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The best-known type of silk is obtained from the cocoons of the larvae of the mulberry silkworm Bombyx mori reared in captivity...

, which can be spun into cloth. Silk is and has been an important economic resource throughout history
History of silk
According to Chinese tradition, the history of silk begins in the 27th century BCE. Its use was confined to China until the Silk Road opened at some point during the latter half of the first millennium BCE. China maintained its virtual monopoly over silk for another thousand years...

. The species Bombyx mori has been domesticated to the point where it is completely dependent on mankind for survival. A number of wild moths such as Bombyx mandarina
Bombyx mandarina
Bombyx mandarina, the Wild Silkmoth, is an insect from the moth family Bombycidae. It is the closest relative of Bombyx mori the Domesticated Silkmoth or "silkworm" . Unlike the domesticated relative which is unable to fly or indeed persist outside human care, the Wild Silkmoth is a fairly ordinary...

, and Antheraea
Antheraea
Antheraea is a moth genus belonging to the family Saturniidae. Several species of this genus have caterpillars which produce wild silk of commercial importance...

species, besides others, provide commercially important silks.

The preference of the larvae of most Lepidopteran species to feed on a single species or limited range of plants is used as a mechanism for biological control of weeds in place of herbicides. The pyralid
Pyralidae
The Pyralidae or snout moths are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many classifications, the grass moths are included in the Pyralidae as a subfamily, making the combined group one of the largest families in the Lepidoptera...

 cactus moth was introduced from Argentina
Argentina
Argentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...

 to Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

, where it successfully suppressed millions of acres of Prickly pear cactus
Opuntia
Opuntia, also known as nopales or paddle cactus , is a genus in the cactus family, Cactaceae.Currently, only prickly pears are included in this genus of about 200 species distributed throughout most of the Americas. Chollas are now separated into the genus Cylindropuntia, which some still consider...

. Another species of the Pyralidae, called the alligator weed stem borer (Arcola malloi), was used to control the aquatic plant
Aquatic plant
Aquatic plants are plants that have adapted to living in aquatic environments. They are also referred to as hydrophytes or aquatic macrophytes. These plants require special adaptations for living submerged in water, or at the water's surface. Aquatic plants can only grow in water or in soil that is...

 known as alligator weed
Alligator weed
Alternanthera philoxeroides, commonly known as Alligator weed, is an immersed aquatic plant. It originated in South America, but has spread to many parts of the world and is considered an invasive species in Australia, China, New Zealand, Thailand and the United States.Alligator weed can grow in a...

 (Alternanthera philoxeroides) in conjunction with the alligator weed flea beetle
Agasicles hygrophila
Agasicles hygrophila is a species of leaf beetle known by the common name alligator weed flea beetle. It has been used successfully as an agent of biological pest control against the noxious aquatic plant known as alligator weed .This beetle is native to South America but has been imported to areas...

; in this case, the two insects work in synergy
Synergy
Synergy may be defined as two or more things functioning together to produce a result not independently obtainable.The term synergy comes from the Greek word from , , meaning "working together".-Definitions and usages:...

 and the weed rarely recovers.

Breeding butterflies and moths, or butterfly gardening
Butterfly gardening
Butterfly gardening is a growing school of gardening, specifically wildlife gardening, that is aimed at creating an environment that attracts butterflies, as well as certain moths, such as those in the Hemaris genus. Butterfly gardening is often aimed at inviting those butterflies and moths to lay...

, has become an ecologically viable process of introducing species into the ecosystem for the better of benefiting it. Butterfly ranching in Papua New Guinea
Butterfly ranching in Papua New Guinea
Butterfly ranching in Papua New Guinea is a method for sustainable use of insect biodiversity endorsed and supported by the national government. The trade is controlled by the Insect Farming and Trading Agency, an organ of the PNG government...

 permits nationals of that country to 'farm' economically valuable insect species for the collectors market in an ecologically sustainable manner.

As food


Lepidoptera feature prominently in entomophagy
Entomophagy
Entomophagy is the consumption of insects as food. Insects are eaten by many animals, but the term is generally used to refer to human consumption of insects; animals that eat insects are known as insectivores...

 as food items on almost every continent. While in most cases, adults, larvae or pupae are eaten as staples by indigenous people, beondegi
Beondegi
Beondegi are a popular snack food in Korean cuisine. Literally meaning "chrysalis" or "pupa" in Korean, Beondegi are steamed or boiled silkworm pupae which are seasoned and eaten as a snack. Beondegi are often served by street vendors, as well as in restaurants and drinking establishments...

 or silkworm pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

e are eaten as a snack in −Korean cuisine while Maguey worm
Maguey worm
A maguey worm is one of two varieties of edible caterpillars that infest maguey and Agave tequilana plants. The white maguey worms, known as meocuiles, are caterpillars of a butterfly commonly named "tequila giant skipper," Aegiale hesperiaris...

 is considered a delicacy in Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...

. In the Carnia
Carnia
Carnia is a historical-geographic region of Friuli, whose municipalities all belong to the province of Udine, which is part of the Friuli-Venezia Giulia Region.It covers the western and central part of the mountainous region of the Province of Udine...

 region of Italy, children catch and eat Zygaena
Zygaena (genus)
Zygaena is a genus of moths in the Zygaenidae family. These brightly coloured, day-flying moths are native to the West Palearctic.-Species:* Zygaena afghana* Zygaena algira* Zygaena alluaudi* Zygaena angelicae Ochsenheimer, 1808...

moths in early summer. The ingluvies, despite having a very low cyanogenic content, serves as a convenient, supplementary source of sugar to the children who can include this resource as a seasonal delicacy at minimum risk.

Health


Some larvae of both moths and butterflies have a form of hair that has been known to be a cause of human health problems. Caterpillar hairs sometimes have venomous toxins in them and species from approximately 12 families of moths or butterflies worldwide can inflict serious human injuries (Urticaria
Urticaria
Urticaria is a kind of skin rash notable for pale red, raised, itchy bumps. Hives is frequently caused by allergic reactions; however, there are many non-allergic causes...

l dermatitis and atopic asthma
Asthma
Asthma is the common chronic inflammatory disease of the airways characterized by variable and recurring symptoms, reversible airflow obstruction, and bronchospasm. Symptoms include wheezing, coughing, chest tightness, and shortness of breath...

 to osteochondritis
Osteochondritis
Osteochondritis is a painful type of osteochondrosis where the cartilage or bone in a joint is inflamed.It often refers to osteochondritis dissecans...

, consumption coagulopathy
Coagulopathy
Coagulopathy is a condition in which the blood’s ability to clot is impaired. This condition can cause prolonged or excessive bleeding, which may occur spontaneously or following an injury or medical and dental procedures.The normal clotting process depends on the interplay of various proteins in...

, renal failure, and intracerebral
Brain
The brain is the center of the nervous system in all vertebrate and most invertebrate animals—only a few primitive invertebrates such as sponges, jellyfish, sea squirts and starfishes do not have one. It is located in the head, usually close to primary sensory apparatus such as vision, hearing,...

 hemorrhage). Skin rashes are the most common, but there have been fatalities. Lonomia
Lonomia
The genus Lonomia is a moderate-sized group of fairly cryptic saturniid moths from South America, famous not for the adults, but for their highly venomous caterpillars, which are responsible for a few deaths each year , especially in southern Brazil, and the subject of hundreds of published medical...

is a frequent cause of economization in humans in Brazil, with 354 cases reported between 1989 and 2005. Lethality ranging up to 20% with death caused most often by intracranial hemorrhage.

These hairs have also been known to cause kerato
Keratin
Keratin refers to a family of fibrous structural proteins. Keratin is the key of structural material making up the outer layer of human skin. It is also the key structural component of hair and nails...

-conjunctivitis
Conjunctivitis
Conjunctivitis refers to inflammation of the conjunctiva...

. The sharp barbs on the end of caterpillar hairs can get lodged in soft tissues and mucus membranes such as the eyes. Once they enter such tissues, they can be difficult to extract, often exacerbating the problem as they migrate across the membrane. This becomes a particular problem in an indoor setting. The hairs easily enter buildings through ventilation systems and accumulate in indoor environments because of their small size, which makes it difficult for them to be vented out. This accumulation increases the risk of human contact in indoor environments.

See also



  • Taxonomy of the Lepidoptera
    Taxonomy of the Lepidoptera
    This is a taxonomy of the order Lepidoptera down to family level.Lepidoptera is made up of moths; and butterflies; . Traditionally, all groups up until Glossata have been regarded as 'archaic', and groups up to Heteroneura as 'primitive'...

  • Lepidoptera in the 10th edition of Systema Naturae
    Lepidoptera in the 10th edition of Systema Naturae
    In the 10th edition of Systema Naturae, Carl Linnaeus classified the arthropods, including insects, arachnids and crustaceans, among his class "Insecta". Butterflies and moths were brought together under the name Lepidoptera. Linnaeus divided the group into three genera – Papilio, Sphinx and Phalaena...

  • Differences between butterflies and moths
    Differences between butterflies and moths
    A common classification of the Lepidoptera involves their differentiation into butterflies and moths. Butterflies are a natural monophyletic group, often given the sub-order Rhopalocera, which includes Papilionoidea , Hesperiidae , and Hedylidae . In this taxonomic scheme moths belong to the...

  • Societas Europaea Lepidopterologica
    Societas Europaea Lepidopterologica
    Societas Europaea Lepidopterologica is the European society for the study of moths and butterflies and for the conservation of these insects and their natural habitats...

  • McGuire Center for Lepidoptera and Biodiversity, University of Florida
    University of Florida
    The University of Florida is an American public land-grant, sea-grant, and space-grant research university located on a campus in Gainesville, Florida. The university traces its historical origins to 1853, and has operated continuously on its present Gainesville campus since September 1906...


Lists


Further reading

  • Kristensen, N. P. (Ed.) 1999. Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography. Handbuch der Zoologie. Eine Naturgeschichte der Stämme des Tierreiches / Handbook of Zoology. A Natural History of the phyla of the Animal Kingdom. Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York.
  • Nye, I. W. B. & Fletcher, D. S. 1991. Generic Names of Moths of the World. Volume 6: xxix + 368 pp. Trustees of the British Museum (Natural History), London.
  • O'Toole, Christopher. 2002. Firefly Encyclopedia of Insects and Spiders. ISBN 1-55297-612-2.

External links