Lepidoptera Of The Philippines
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Lepidoptera ( ) is an
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s that includes
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
and
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s (both are called lepidopterans). About 180,000
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of the Lepidoptera are described, in 126
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
and 46 superfamilies, 10 percent of the total described species of living organisms. It is one of the most widespread and widely recognizable insect orders in the world. The Lepidoptera show many variations of the basic body structure that have evolved to gain advantages in lifestyle and distribution. Recent estimates suggest the order may have more species than earlier thought, and is among the four most speciose orders, along with the
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
,
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
, and
Coleoptera Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
. Lepidopteran species are characterized by more than three derived features. The most apparent is the presence of scales that cover the
bodies Bodies may refer to: * The plural of body * ''Bodies'' (2004 TV series), BBC television programme * Bodies (upcoming TV series), an upcoming British crime thriller limited series * "Bodies" (''Law & Order''), 2003 episode of ''Law & Order'' * ...
,
wings A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expresse ...
, and a
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
. The scales are modified, flattened "hairs", and give butterflies and moths their wide 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.
Mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
and the laying of eggs is normally performed near or on host plants for the
larvae A larva (; plural larvae ) 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. The ...
. Like most other insects, butterflies and moths are
holometabolous Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygot ...
, meaning they undergo
complete metamorphosis Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. ...
. The larvae are commonly called
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
s, and are completely different from their adult moth or butterfly forms, having a cylindrical body with a well-developed head, mandible mouth parts, three pairs of thoracic legs and from none up to five pairs of
proleg A proleg is a 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 other types of in ...
s. As they grow, these larvae change in appearance, going through a series of stages called
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
s. Once fully matured, the larva develops into a
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
. A few butterflies and many moth species spin a silk case or cocoon prior to pupating, while others do not, instead going underground. A butterfly pupa, called a
chrysalis A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
, has a hard skin, usually with no cocoon. Once the pupa has completed its metamorphosis, a sexually mature adult emerges. 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 The caddisflies, or order Trichoptera, are a group of insects with aquatic larvae and terrestrial adults. There are approximately 14,500 described species, most of which can be divided into the suborders Integripalpia and Annulipalpia on the b ...
, 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 Lepidopterology ()) is a branch of entomology concerning the scientific study of moths and the three superfamilies of butterflies. Someone who studies in this field is a lepidopterist or, archaically, an aurelian. Origins Post-Renaissance, t ...
. Butterflies and moths play an important role in the natural ecosystem as
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
s and as food in the food chain; conversely, their larvae 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 from 200 to 600 eggs, while in others, the number may approach 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 is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
they produce, or as pest species.


Etymology

The term Lepidoptera was used in 1746 by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
in his ''
Fauna Svecica ''Fauna Svecica'' ("Fauna of Sweden", ed. 1, Stockholm, 1746; ed. 2 Stockholm, 1761) was written by Swedish botanist, physician, zoologist and naturalist Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after ...
''. The word is derived from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
,
gen. The Book of Genesis (from Greek language, Greek ; Hebrew language, Hebrew: בְּרֵאשִׁית ''Bəreʾšīt'', "In hebeginning") is the first book of the Hebrew Bible and the Christian Old Testament. Its Hebrew name is the same as its i ...
(" scale") and ("wing"). Sometimes, the term
Rhopalocera Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...
is used for the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of all butterfly species, derived from the Ancient Greek (') and (') meaning "club" and "horn", respectively, coming from the shape of the
antenna Antenna ( antennas or antennae) may refer to: Science and engineering * Antenna (radio), also known as an aerial, a transducer designed to transmit or receive electromagnetic (e.g., TV or radio) waves * Antennae Galaxies, the name of two collid ...
e of butterflies. The origins of the common names "butterfly" and "moth" are varied and often obscure. The
English English usually refers to: * English language * English people English may also refer to: Peoples, culture, and language * ''English'', an adjective for something of, from, or related to England ** English national ide ...
word butterfly is from
Old English Old English (, ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the early Middle Ages. It was brought to Great Britain by Anglo-Saxon settlement of Britain, Anglo ...
', 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. The species of
Heterocera Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
are commonly called
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s. The origins of the English word moth are clearer, deriving from Old English] ' (cf.
Northumbrian dialect The Northumbrian dialect refers to any of several English language varieties spoken in the traditional English region of Northumbria, which includes most of the North East England government region. The traditional Northumbrian dialect is a m ...
') from Common Germanic (compare
Old Norse Old Norse, Old Nordic, or Old Scandinavian, is a stage of development of North Germanic languages, North Germanic dialects before their final divergence into separate Nordic languages. Old Norse was spoken by inhabitants of Scandinavia and t ...
',
Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...
' and
German German(s) may refer to: * Germany (of or related to) **Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ger ...
' all meaning "moth"). Perhaps its origins are related to Old English ' meaning "
maggot A maggot is the larva of a fly (order Diptera); it is applied in particular to the larvae of Brachycera 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 A midge is any small fly, including species in several families of non-mosquito Nematoceran Diptera. Midges are found (seasonally or otherwise) on practically every land area outside permanently arid deserts and the frigid zones. Some mid ...
", 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 (abbreviated to ME) is a form of the English language that was spoken after the Norman conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English p ...
''catirpel'', ''catirpeller'', probably an alteration of Old North French ''catepelose'' (from Latin ''cattus'', "cat" + ''pilosus'', "hairy").


Distribution and diversity

The Lepidoptera are among the most successful groups of insects. They are found on all continents, except
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
, and inhabit all terrestrial habitats ranging from desert to rainforest, from lowland grasslands to mountain plateaus, but almost always associated with higher plants, especially angiosperms (
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s). Among the most northern dwelling species of butterflies and moths is the Arctic Apollo (''
Parnassius arcticus ''Parnassius arcticus'', the Siberian Apollo, is a high-altitude butterfly which is found in Northeastern Yakutia, Russia. It is a member of the snow Apollo genus (''Parnassius'') of the swallowtail family, Papilionidae. The taxonomic status of ...
''), which is found in the Arctic Circle in northeastern
Yakutia Sakha, officially the Republic of Sakha (Yakutia),, is the largest republic of Russia, located in the Russian Far East, along the Arctic Ocean, with a population of roughly 1 million. Sakha comprises half of the area of its governing Far Eas ...
, at an altitude of above sea level. In the
Himalayas The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
, various Apollo species such as ''
Parnassius epaphus ''Parnassius epaphus'', the common red Apollo, is a high altitude butterfly which is found in India and Nepal. It is a member of the snow Apollo genus (''Parnassius'') of the swallowtail family (Papilionidae). It is found from from Chitral Distr ...
'' have been recorded to occur up to an altitude of above sea level. Some lepidopteran species exhibit
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
,
phoretic Phoresis or phoresy is a non-permanent, commensalistic interaction in which one organism (a phoront or phoretic) attaches itself to another (the host) solely for the purpose of travel. Phoresis has been observed directly in ticks and mites s ...
, or
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...
lifestyles, inhabiting the bodies of organisms rather than the environment.
Coprophagous Coprophagia () or coprophagy () is the consumption of feces. The word is derived from the grc, κόπρος , "feces" and , "to eat". Coprophagy refers to many kinds of feces-eating, including eating feces of other species (heterospecifics), of ...
pyralid The Pyraloidea (pyraloid moths or snout moths) 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, and as such, they have be ...
moth species, called
sloth moth A sloth moth is a coprophagous moth which has evolved to exclusively inhabit the fur of sloths and to use sloth dung as a substrate for the early stages of reproduction. Sloth moths include '' Bradypodicola hahneli'', ''Cryptoses choloepi'', ''Cry ...
s, such as ''
Bradipodicola hahneli ''Bradypodicola hahneli'' is a sloth moth in the family Pyralidae that lives exclusively in the fur of the pale-throated three-toed sloth (''Bradypus tridactylus''), a three-toed sloth found in South America. It is the only species of the genus ...
'' and ''
Cryptoses choloepi ''Cryptoses choloepi'' is a sloth moth in the snout moth family that as an adult lives exclusively in the fur of sloths, mammals found in South and Central America. Adult female moths live in the fur of the brown three-toed sloth '' Bradypus vari ...
'', are unusual in that they are exclusively found inhabiting the fur of
sloth Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...
s, mammals found in
Central Central is an adjective usually referring to being in the center of some place or (mathematical) object. Central may also refer to: Directions and generalised locations * Central Africa, a region in the centre of Africa continent, also known as ...
and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. Two species of ''
Tinea Dermatophytosis, also known as ringworm, is a fungal infection of the skin. Typically it results in a red, itchy, scaly, circular rash. Hair loss may occur in the area affected. Symptoms begin four to fourteen days after exposure. Multiple ar ...
'' moths have been recorded as feeding on horny tissue and have been bred from the horns of cattle. The larva of ''
Zenodochium ''Zenodochium'' is a genus of moths in the family Blastodacnidae The Agonoxeninae are a subfamily of moths. History of classification Formerly, the subfamily only contained four named species – all in the type genus ''Agonoxena'' &ndash ...
coccivorella'' is an internal parasite of the
coccid Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than the ...
'' Kermes'' species. Many species have been recorded as breeding in natural materials or refuse such as owl pellets, bat caves, honeycombs or diseased fruit. As of 2007, there were roughly 174,250 lepidopteran species described, with butterflies and skippers estimated to comprise around 17,950, and 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 about 400 species of butterflies and 14,000 species of moths are 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 indices in the
Natural History Museum A natural history museum or museum of natural history is a scientific institution with natural history collections that include current and historical records of animals, plants, fungi, ecosystems, geology, paleontology, climatology, and more. ...
(London) and the
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. It has free admission and is open 364 days a year. In 2021, with 7 ...
(Washington), and partly on estimates:


External morphology

Lepidoptera are morphologically distinguished from other orders principally by the presence of scales on the external parts of the body and appendages, especially the
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expres ...
s. Butterflies and moths vary in size from
microlepidoptera Microlepidoptera (micromoths) is an artificial (i.e., unranked and not monophyletic) grouping of moth families, commonly known as the 'smaller moths' (micro, Lepidoptera). These generally have wingspans of under 20 mm, and are thus harder to ...
only a few millimeters long, to conspicuous animals with a wingspan greater than , such as the
Queen Alexandra's birdwing ''Ornithoptera alexandrae'', the Queen Alexandra's birdwing, is the largest species of butterfly in the world, with females reaching wingspans slightly in excess of 25 cm to 28 cm (9.8 inches to 11 inches). This birdwing is restricted to the ...
and
Atlas moth ''Attacus atlas'', the Atlas moth, is a large saturniid moth endemic to the forests of Asia. The species was first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''. Description The Atlas moth is one of the largest ...
. Lepidopterans undergo a four-stage
life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia *Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring *Life-cycle hypothesis, ...
:
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
;
larva A larva (; plural larvae ) 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. The ...
or
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
;
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
or
chrysalis A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
; and imago (plural: imagines) / adult and show many variations of the basic body structure, which give these animals advantages for diverse lifestyles and environments.


Head

The head is where many sensing organs and the mouth parts are found. Like the adult, the larva also has a toughened, or sclerotized head capsule.Scoble (1995). Section ''The Adult Head – Feeding and Sensation'', (pp. 4–22). Here, two
compound eye A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which disti ...
s, and ''chaetosema'', raised spots or clusters of sensory bristles unique to Lepidoptera, occur, 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 enlarged antennae or ones that are tapered and hooked at the ends. The maxillary galeae are modified and form an elongated
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
. 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 moths still have
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s, or separate moving jaws, like their ancestors, and these form the family
Micropterigidae Micropterigoidea is the superfamily of "mandibulate archaic moths", all placed in the single family Micropterigidae, containing currently about twenty living genera. They are considered the most primitive extant lineage of lepidoptera (Kristense ...
. The larvae, called
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
s, have a toughened head capsule. Caterpillars lack the proboscis and have separate chewing mouthparts. These mouthparts, called
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s, are used to chew up the plant matter that the larvae eat. The lower jaw, or labium, is weak, but may carry a
spinneret A spinneret is a silk-spinning organ of a spider or the larva of an insect. Some adult insects also have spinnerets, such as those borne on the forelegs of Embioptera. Spinnerets are usually on the underside of a spider's opisthosoma, and are ...
, 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 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 (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on ea ...
,
mesothorax The mesothorax is the middle of the three segments of the thorax of hexapods, and bears the second pair of legs. Its principal sclerites (exoskeletal plates) are the mesonotum (dorsal), the mesosternum (ventral), and the mesopleuron (lateral) on ...
, and
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 ( exoskeletal plates) are the metanotum (dorsal), the metasternum (ventral), and the metapleuron (lateral) ...
, each with a pair of legs. The first segment contains the first pair of legs. In some males of the butterfly family
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...
, the forelegs 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 up to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets. The two pairs of wings are found on the middle and third segments, or
mesothorax The mesothorax is the middle of the three segments of the thorax of hexapods, and bears the second pair of legs. Its principal sclerites (exoskeletal plates) are the mesonotum (dorsal), the mesosternum (ventral), and the mesopleuron (lateral) on ...
and
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 ( exoskeletal plates) are the metanotum (dorsal), the metasternum (ventral), and the metapleuron (lateral) ...
, 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 has 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, the
Noctuoidea Noctuoidea is the superfamily of noctuid (Latin "night owl") or "owlet" moths, and has more than 70,000 described species, the largest number of for any Lepidopteran superfamily. Its classification has not yet reached a satisfactory or stable st ...
, has their wings modified to act as tympanal or hearing organs. The caterpillar has an elongated, soft body that may have hair-like or other projections, three pairs of true legs, with none to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets. The thorax usually has 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, which instead have a form of
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
s.


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 two or three segments form the external parts of the species' sex organs. The
genitalia A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
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. The females of basal moths have only one sex organ, which is used for
copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetrat ...
and as an
ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
, or egg-laying organ. About 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 four 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 The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo'' γεω (derivative form of or "the earth"), and ''met ...
). In some basal moths, these prolegs may be on every segment of the body, while prolegs may be completely absent in other groups, which are more adapted to boring and living in sand (e. g.,
Prodoxidae The Prodoxidae are a family of moths, generally small in size and nondescript in appearance. They include species of moderate pest status, such as the Lampronia capitella, currant shoot borer, and others of considerable ecological and evolutiona ...
and
Nepticulidae Nepticulidae is a family of very small moths with a worldwide distribution. They are characterised by eyecaps over the eyes (see also Opostegidae, Bucculatricidae, Lyonetiidae). These pigmy moths or midget moths, as they are commonly known, inc ...
, respectively).


Scales

The wings, head, and parts of the thorax and abdomen of Lepidoptera are covered with minute scales, a feature from which the order derives its name. Most scales are
lamella Lamella (plural lamellae) means a small plate or flake in Latin, and in English may refer to: Biology * Lamella (mycology), a papery rib beneath a mushroom cap * Lamella (botany) * Lamella (surface anatomy), a plate-like structure in an animal * ...
r, or blade-like, and attached with a pedicel, while other forms may be hair-like or specialized as secondary sexual characteristics.Scoble (1995). Section ''Scales'', (pp. 63–66). The lumen or surface of the lamella has a complex structure. It gives color either by colored
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
s it contains, or through
structural coloration Structural coloration in animals, and a few plants, is the production of colour by microscopically structured surfaces fine enough to interfere with visible light instead of pigments, although some structural coloration occurs in combination wit ...
with mechanisms that include
photonic crystal A photonic crystal is an optical nanostructure in which the refractive index changes periodically. This affects the propagation of light in the same way that the structure of Crystal structure, natural crystals gives rise to X-ray crystallograp ...
s and
diffraction grating In optics, a diffraction grating is an optical component with a periodic structure that diffracts light into several beams travelling in different directions (i.e., different diffraction angles). The emerging coloration is a form of structura ...
s. Scales function in insulation, thermoregulation, producing
pheromones 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 like hormones outside the body of the secreting individual, to affect the behavio ...
( in males only), and aiding gliding flight, but the most important is the large diversity of vivid or indistinct
patterns A pattern is a regularity in the world, in human-made design, or in abstract ideas. As such, the elements of a pattern repeat in a predictable manner. A geometric pattern is a kind of pattern formed of geometric shapes and typically repeated l ...
they provide, which help the organism protect itself by
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
or
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
, and which act as signals to other animals including rivals and potential mates.


Internal morphology


Reproductive system

In the reproductive system of butterflies and moths, the male
genitalia A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
are complex and unclear. In females the three types of genitalia are based on the relating taxa: 'monotrysian', 'exoporian', and 'ditrysian'. In the monotrysian type is an opening on the fused segments of the sterna 9 and 10, which act as insemination and oviposition. In the exoporian type (in
Hepialoidea The Hepialoidea are the superfamily of "ghost moths" and "swift moths". Fossils Fossil Hepialoidea appear to be few. ''Prohepialus'' (possibly Hepialidae) has been described from the about 35-million-year-old Bembridge marls of Isle of Wight. ...
and
Mnesarchaeoidea Mnesarchaeoidea is a superfamily of "New Zealand primitive moths" containing one family, Mnesarchaeidae and a two genera, ''Mnesarchaea'', and ''Mnesarchella,'' both of which are endemic to New Zealand. Taxonomy and systematics Mnesarchaeoid ...
) 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 carries 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 cycles, called the
endocrine system The endocrine system is a messenger system comprising feedback loops of the hormones released by internal glands of an organism directly into the circulatory system, regulating distant target organs. In vertebrates, the hypothalamus is the neu ...
. The first insect hormone
prothoracicotropic hormone Prothoracicotropic hormone (PTTH) was the first insect hormone to be discovered.The chemical symbol for prothoracicotropic hormone is (C64H102N16O19S2). It was originally described simply as "brain hormone" by early workers such as Stefan Kopeć (1 ...
(PTTH) operates the species life cycle and
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
. This hormone is produced by
corpora allata In insect physiology and anatomy, the corpus allatum (plural: corpora allata) is an endocrine gland that generates juvenile hormone; as such, it plays a crucial role in metamorphosis. Surgical removal of the corpora allata (an allatectomy) can cau ...
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 juvenile hormones, and the prothorocic glands produce moulting hormones.


Digestive system

In the
digestive system The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver, and gallbladder). Digestion involves the breakdown of food into smaller and smaller compone ...
, 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 The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
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 is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
, 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 cells are simple columnar epithelial cells that secrete gel-forming mucins, like mucin 5AC. The goblet cells mainly use the merocrine method of secretion, secreting vesicles into a duct, but may use apocrine methods, budding off their secre ...
s are found in the anterior and posterior midgut regions, respectively. In insects, the goblet cells excrete positive
potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosphe ...
ions, which are absorbed from leaves ingested by the larvae. Most butterflies and moths display the usual digestive cycle, but species with different diets require adaptations to meet these new demands.


Circulatory system

In the
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
,
hemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
, or insect blood, is used to circulate heat in a form of
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
, 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 The trachea, also known as the windpipe, is a Cartilage, cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends ...
.


Respiratory system

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 The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies grea ...
. Three different tracheaes supply and diffuse oxygen throughout the species' bodies. 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 is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, between geographically separated populations in
geographical polymorphism Geography (from Greek: , ''geographia''. Combination of Greek words ‘Geo’ (The Earth) and ‘Graphien’ (to describe), literally "earth description") is a field of science devoted to the study of the lands, features, inhabitants, and ...
, and between generations flying at different seasons of the year (
seasonal polymorphism A season is a division of the year based on changes in weather, ecology, and the number of daylight hours in a given region. On Earth, seasons are the result of the axial parallelism of Earth's tilted orbit around the Sun. In temperate and pol ...
or
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. It is therefore a special case of phenotypic plasticity. There are several types of polyphen ...
). In some species, the polymorphism is limited to one sex, typically the female. This often includes the phenomenon of
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
when mimetic morphs fly alongside nonmimetic morphs in a population of a particular species. Polymorphism occurs both at specific level with heritable variation in the overall morphological adaptations of individuals, as well as in certain specific morphological or physiological traits within a species. Environmental polymorphism, in which traits are not inherited, is often termed as polyphenism, which in Lepidoptera is commonly seen in the form of seasonal morphs, especially in the butterfly families of
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...
and
Pieridae The Pieridae are a large family of butterflies with about 76 genera containing about 1,100 species, mostly from tropical Africa and tropical Asia with some varieties in the more northern regions of North America and Eurasia.DeVries P. J. in Levi ...
. An Old World pierid butterfly, the common grass yellow (''
Eurema hecabe ''Eurema hecabe'', the common grass yellow, is a small pierid butterfly species found in Asia, Africa and Australia. They are found flying close to the ground and are found in open grass and scrub habitats. It is simply known as "the grass yellow ...
'') 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 postmonsoon period. Polyphenism also occurs in caterpillars, an example being the peppered moth, ''
Biston betularia The peppered moth (''Biston betularia'') is a temperate species of Nocturnality, night-flying moth. It is mostly found in the northern hemisphere in places like Asia, Europe and North America. Peppered moth evolution is an example of populatio ...
''. Geographical isolation causes a divergence of a species into different morphs. A good example is the Indian white admiral ''
Limenitis procris ''Moduza procris'', the commander, sometimes included in the genus ''Limenitis'', is a medium-sized, strikingly coloured brush-footed butterfly found in South Asia and Southeast Asia. It is notable for the mode of concealment employed by its cate ...
'', 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 (''Parnassius apollo''). Because the Apollos live in small local populations, thus having no contact with each other, coupled with their strong stenotopic nature and weak migration ability, interbreeding between populations of one species practically does not occur; by this, they form over 600 different morphs, with the size of spots on the wings of which varies greatly. Sexual dimorphism is the occurrence of differences between males and females in a species. In Lepidoptera, it is widespread and almost completely set by genetic determination. Sexual dimorphism is present in all families of the Papilionoidea and more prominent in the
Lycaenidae Lycaenidae is the second-largest family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfl ...
,
Pieridae The Pieridae are a large family of butterflies with about 76 genera containing about 1,100 species, mostly from tropical Africa and tropical Asia with some varieties in the more northern regions of North America and Eurasia.DeVries P. J. in Levi ...
, and certain taxa of the
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...
. Apart from color variation, which may differ from slight to completely different color-pattern combinations, secondary sexual characteristics may also be present.Kunte, Krushnamegh (2000). ''Butterflies of Peninsular India''. Part of Project lifescape. Orient Blackswan. , . 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 Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includes the larges ...
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 The Psychidae (bagworm moths, also simply bagworms or bagmoths) are a family of the Lepidoptera (butterflies and moths). The bagworm family is fairly small, with about 1,350 species described. Bagworm species are found globally, with some, su ...
species which have only vestigial wings, legs, and mouthparts as compared to the adult males that are strong fliers with well-developed wings and feathery antennae.


Reproduction and development

Species of Lepidoptera undergo
holometabolism Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygot ...
or "complete metamorphosis". Their life cycle normally consists of an
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
, a
larva A larva (; plural larvae ) 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. The ...
, a
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
, and an
imago In biology, the imago (Latin for "image") is the last stage an insect attains during its metamorphosis, its process of growth and development; it is also called the imaginal stage, the stage in which the insect attains maturity. It follows the f ...
or adult. The larvae are commonly called
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
s, and the pupae of moths encapsulated in silk are called cocoons, while the uncovered pupae of butterflies are called
chrysalides A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
.


Lepidopterans in diapause

Unless the species reproduces year-round, a butterfly or moth may enter
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
, a state of dormancy that allows the insect to survive unfavorable environmental conditions.


Mating

Males usually start
eclosion A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
(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 do not 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. However, the females of most nocturnal species, including almost all moth species, use
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 like hormones outside the body of the secreting individual, to affect the behavio ...
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 or oleander moth, is a species of moth thought to be native to the Caribbean. Its larvae feed on the oleander plant. Like most wasp moths, these are day fliers. They prefer Neotropic areas, to wh ...
''. Adaptations include undergoing one seasonal generation, two or even more, called
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 most often applied to insects, and is particularly in use in sericulture, where silkworm varieties vary in their voltinism. ...
(Univoltism, bivoltism, and multivism, respectively). Most lepidopterans in
temperate climate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
s 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 In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
. 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. 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 Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and ...
(egg-laying), though some species exhibit live birth in a process called
ovoviviparity Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
. A variety of differences in
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
-laying and the number of eggs laid occur. Some species simply drop their eggs in flight (these species normally have polyphagous larvae, meaning they eat a variety of plants e. g., hepialids and some nymphalids) while most lay their eggs near or on the host plant on which the larvae feed. 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, and primarily, by chemical cues. The eggs are derived from materials ingested as a larva and in some species, from the spermatophores received from males during mating. An egg can only be 1/1000 the mass of the female, yet she may lay up to her own mass in eggs. Females lay smaller eggs as they age. Larger females lay larger eggs. The egg is covered by a hard-ridged protective outer layer of shell, called the
chorion The chorion is the outermost fetal membrane around the embryo in mammals, birds and reptiles (amniotes). It develops from an outer fold on the surface of the yolk sac, which lies outside the zona pellucida (in mammals), known as the vitelline ...
. It is lined with a thin coating of
wax Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to give low ...
, which prevents the egg from drying out. 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 prior to winter, especially in
temperate region In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
s, go through
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
, 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 temperate species (e. g. ''
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. The immature form of this species is sometimes known as the spiny elm caterpillar ...
'').


Larvae

The larvae or caterpillars 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 sclerotized head capsule with an adfrontal suture formed by medial fusion of the sclerites, 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 A proleg is a 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 other types of in ...
s (up to five pairs). The body consists of thirteen segments, of which three are thoracic and ten are abdominal. Most larvae are
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s, but a few are
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
s (some eat ants or other caterpillars) and
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
s. Different herbivorous species have adapted to feed on every part of the plant and are normally considered pests to their host plants; 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 ''Tineola'' is a genus of moths the family Tineidae. There are two species, including the familiar common clothes moth (''T. bisselliella'').lycaenid Lycaenidae is the second-largest family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfl ...
species such as ''
Phengaris rebeli ''Phengaris rebeli'' (formerly ''Maculinea rebeli''), common name mountain Alcon blue, is a species of butterfly in the family Lycaenidae. It was first found and described in Styria, Austria, on Mount Hochschwab around 1700. Although it was in ...
'' are social parasites of ''
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, ...
'' ants nests. A species of
Geometridae The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo'' γεω (derivative form of or "the earth"), and ''met ...
from
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
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'', the Arctic woolly bear moth, is an erebid moth native to the High Arctic in the Canadian archipelago, Greenland and Wrangel Island in Russia. It is known for its slow rate of development, as its full caterpillar li ...
'' 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 is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remna ...
, developing further with each
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
, until they undergo the final larval-pupal molt. The larvae of both butterflies and moths exhibit mimicry to deter potential predators. Some caterpillars have the ability to inflate parts of their heads to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called
osmeteria The osmeterium is a defensive organ found in all Papilionidae, papilionid larvae, in all stages. The organ is situated in the prothorax, prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy f ...
(family
Papilionidae Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includes the larges ...
), which are exposed to produce smelly chemicals 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 make 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

No form of wing is externally visible on the larva, but 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 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, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
, 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 five to seven instars, or molts, certain hormones, like PTTH, stimulate the production of
ecdysone Ecdysone is a prohormone of the major insect molting hormone 20-hydroxyecdysone, which is secreted from the prothoracic glands. It is of steroidal structure. Insect molting hormones (ecdysone and its homologues) are generally called ecdysteroids. ...
, which initiates insect molting. The larva starts to develop into the
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
: body parts specific to the larva, such as the abdominal prolegs, degenerate, while others such as the legs and wings undergo growth. After finding a suitable place, the animal sheds its last larval cuticle, revealing the pupal cuticle underneath. Depending on the species, the pupa may be covered in a silk cocoon, attached to different types of substrates, buried in the ground, or may not be covered at all. Features of the
imago In biology, the imago (Latin for "image") is the last stage an insect attains during its metamorphosis, its process of growth and development; it is also called the imaginal stage, the stage in which the insect attains maturity. It follows the f ...
are externally recognizable in the pupa. All the appendages on the adult head and thorax are found cased inside the cuticle (
antenna Antenna ( antennas or antennae) may refer to: Science and engineering * Antenna (radio), also known as an aerial, a transducer designed to transmit or receive electromagnetic (e.g., TV or radio) waves * Antennae Galaxies, the name of two collid ...
e, mouthparts, etc.), with the wings wrapped around, adjacent to the antennae. The pupae of some species have functional mandibles, while the pupal mandibles are not functional in others. Although the pupal cuticle is highly sclerotized, some of the lower abdominal 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 beans (also known as ' in Spanish) are seed pods that have been inhabited by the larva of a small moth (''Cydia saltitans'') and are native to Mexico. The "bean" is usually tan to brown. They are from the shrub '' Sebastiania pa ...
moth (''
Cydia saltitans ''Cydia saltitans'' 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 related shrubby trees, mainly '' Sebastiania pavoniana'' or '' Sapium biloculare'' ( syn ...
'') does this. The larvae cut a trapdoor in the bean (species of ''
Sebastiania ''Sebastiania'' is a genus of flowering plants in the family Euphorbiaceae first described in 1821. It is native to North and South America from Arizona and the West Indies south to Uruguay.Webster, G. L. & M.J. Huft. 1988. Revised synopsis of Pa ...
'') and use the bean as a shelter. With a sudden rise in temperature, the pupa inside twitches and jerks, pulling on the threads inside. Wiggling may also help to deter
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
wasps from laying eggs on the pupa. Other species of moths 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. The adult emerges 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 Micropterigoidea is the superfamily of "mandibulate archaic moths", all placed in the single family Micropterigidae, containing currently about twenty living genera. They are considered the most primitive extant lineage of lepidoptera (Kristense ...
).


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 a longer period (from one to several weeks), or go through diapause and overwintering as monarch butterflies do, or waiting out environmental stress. Some adult species of microlepidoptera go through a stage where no reproductive-related activity occurs, lasting through summer and winter, followed by mating and oviposition in the early spring. While most butterflies and moths are
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
, many species of
Acentropinae Acentropinae is a fairly small subfamily of the lepidopteran family Crambidae, the crambid snout moths. Species of this subfamily are exclusively found in wetlands and aquatic habitats. Systematics In modern treatments, the former subfa ...
(
Crambidae The Crambidae are the grass moth family of lepidopterans. They are variable in appearance, the nominal subfamily Crambinae (grass moths) taking up closely folded postures on grass stems where they are inconspicuous, while other subfamilies includ ...
) are truly aquatic with all stages except the adult occurring in water. Many species from other families such as
Erebidae The Erebidae are a family of moths in the superfamily Noctuoidea. The family is among the largest families of moths by species count and contains a wide variety of well-known macromoth groups. The family includes the underwings ('' Catocala'') ...
,
Nepticulidae Nepticulidae is a family of very small moths with a worldwide distribution. They are characterised by eyecaps over the eyes (see also Opostegidae, Bucculatricidae, Lyonetiidae). These pigmy moths or midget moths, as they are commonly known, inc ...
,
Cosmopterigidae The Cosmopterigidae are a family of insects (cosmet moths) in the order Lepidoptera. These are small moths with narrow wings whose tiny larvae feed internally on the leaves, seeds and stems of their host plants. About 1500 species are described. ...
,
Tortricidae The Tortricidae are a family of moths, commonly known as tortrix moths or leafroller moths, in the order Lepidoptera. This large family has over 11,000 species described, and is the sole member of the superfamily Tortricoidea, although the genu ...
,
Olethreutidae The Tortricidae are a family of moths, commonly known as tortrix moths or leafroller moths, in the order Lepidoptera. This large family has over 11,000 species described, and is the sole member of the superfamily Tortricoidea, although the genus ...
,
Noctuidae The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, along with the other f ...
,
Cossidae The Cossidae, the cossid millers or carpenter millers, make up a family (biology), family of mostly large Miller (moth), miller moths. This family contains over 110 genera with almost 700 known species, and many more species await description. ...
, and
Sphingidae The Sphingidae are a family of moths (Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as “hornworms”; it includes about 1,450 species. It is best represented in the tropics, bu ...
are aquatic or
semiaquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semiaquatic animals include: * Verte ...
.


Behavior


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 most 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 lepidopteran species reportedly can still fly when their hindwings are cut off, it reduces their linear flight and turning capabilities. Lepidopteran species have to be warm, about , to fly. They depend on their body temperature being sufficiently high and since they cannot 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) 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 butterflies can reach fast speeds, such as the southern dart, which can go as fast as . Sphingids are some of the fastest flying insects, some are capable of flying at over , having a wingspan of . In some species, sometimes a gliding component to their flight exists. Flight occurs either as hovering, or as forward or backward motion. In butterfly and moth species, such as
hawk moth The Sphingidae are a family of moths (Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as “hornworms”; it includes about 1,450 species. It is best represented in the tropics, but ...
s, hovering is important as they need to maintain a certain stability over flowers when feeding on the nectar.


Navigation

Navigation Navigation is a field of study that focuses on the process of monitoring and controlling the movement of a craft or vehicle from one place to another.Bowditch, 2003:799. The field of navigation includes four general categories: land navigation, ...
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 Polarization (also polarisation) is a property applying to transverse waves that specifies the geometrical orientation of the oscillations. In a transverse wave, the direction of the oscillation is perpendicular to the direction of motion of the ...
, so can orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is suggested to be particularly important. Most migratory butterflies are those that live in semiarid 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, 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, also known as the geomagnetic field, is the magnetic field that extends from Earth's interior out into space, where it interacts with the solar wind, a stream of charged particles emanating from the Sun. The magnetic f ...
to navigate, as a study of the
heart and dart The heart and dart (''Agrotis exclamationis'') is a moth of the family Noctuidae. The species was first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''. A familiar moth to many, it is considered one of the most commo ...
moth suggests. Another study, of the migratory behavior of the
silver Y The silver Y (''Autographa gamma'') is a migratory moth of the family Noctuidae which is named for the silvery Y-shaped mark on each of its forewings. Description The silver Y is a medium-sized moth with a wingspan of 30 to 45 mm. The wing ...
, showed, 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, is a species of Lepidoptera in the family ''Pieridae''. The species is a medium-sized yellow butterfly, with females more pale than males. They are found from southern regions of Florida and Texas throug ...
'' in
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Cos ...
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 they use a technique of
celestial navigation Celestial navigation, also known as astronavigation, is the practice of position fixing using stars and other celestial bodies that enables a navigator to accurately determine their actual current physical position in space (or on the surface of ...
called
transverse orientation Transverse orientation, keeping a fixed angle on a distant source of light for orientation, is a proprioceptive response displayed by some insects such as moth Moths are a paraphyletic group of insects that includes all members of the order Le ...
. By maintaining a constant angular relationship to a bright celestial light, such as the
Moon The Moon is Earth's only natural satellite. It is the fifth largest satellite in the Solar System and the largest and most massive relative to its parent planet, with a diameter about one-quarter that of Earth (comparable to the width of ...
, they can fly in a straight line. Celestial objects are so far away, 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 The horizon is the apparent line that separates the surface of a celestial body from its sky when viewed from the perspective of an observer on or near the surface of the relevant body. This line divides all viewing directions based on whether i ...
. 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 Mach bands is an optical illusion named after the physicist Ernst Mach. It exaggerates the contrast between edges of the slightly differing shades of gray, as soon as they contact one another, by triggering edge-detection in the human visual s ...
by Henry Hsiao in 1972. He stated that they fly towards the darkest part of the sky in pursuit of safety, thus are inclined to circle ambient objects in the Mach band region.


Migration

Lepidopteran migration is typically
season A season is a division of the year based on changes in weather, ecology, and the number of daylight hours in a given region. On Earth, seasons are the result of the axial parallelism of Earth's tilted orbit around the Sun. In temperate and pol ...
al, as the insects moving to escape dry seasons or other disadvantageous conditions. Most lepidopterans that migrate are butterflies, and the distance travelled varies. Some butterflies that migrate include the
mourning cloak ''Nymphalis antiopa'', known as the mourning cloak in North America and the Camberwell beauty in United Kingdom, Britain, is a large butterfly native to Eurasia and North America. The immature form of this species is sometimes known as the spiny ...
,
painted lady ''Vanessa cardui'' is the most widespread of all butterfly species. It is commonly called the painted lady, or formerly in North America the cosmopolitan. Description File:Vanessa cardui MHNT CUT 2013 3 14 Pontfaverger-Moronvilliers Dos. ...
, American lady, red admiral, and the
common buckeye ''Junonia coenia'', known as the common buckeye or buckeye, is a butterfly in the family Nymphalidae. It is found in the United States east of the Rocky Mountains and in Mexico. Its habitat is open areas with low vegetation and some bare ground. ...
. A notable species of moth that migrates long distances is the
bogong moth The bogong moth (''Agrotis infusa'') is a temperate species of night-flying moth, notable for its biannual long-distance seasonal migrations towards and from the Australian Alps, similar to the diurnal monarch butterfly. During the autumn a ...
. The most well-known migrations are those of the eastern population of the
monarch butterfly The monarch butterfly or simply monarch (''Danaus plexippus'') is a milkweed butterfly (subfamily Danainae) in the family Nymphalidae. Other common names, depending on region, include milkweed, common tiger, wanderer, and black-veined brown. It ...
from Mexico to northern United States and southern Canada, a distance of about . Other well-known migratory species include the painted lady and several of the danaine butterflies. Spectacular and large-scale migrations associated with the
monsoons A monsoon () is traditionally a seasonal reversing wind accompanied by corresponding changes in precipitation but is now used to describe seasonal changes in atmospheric circulation and precipitation associated with annual latitudinal oscil ...
are seen in peninsular India. Migrations have been studied in more recent times using wing tags and stable hydrogen isotopes. Moths also undertake migrations, an example being the uraniids. ''
Urania fulgens ''Urania fulgens'', the urania swallowtail moth or green page moth, is a day-flying moth of the family Uraniidae. The species was first described by Francis Walker in 1854. It is found from Veracruz, Mexico, through Central America to northwes ...
'' undergoes population explosions and massive migrations that may be not surpassed by any other insect in the
Neotropic The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
s. In
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
and
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Cos ...
, 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 antennae. In many species, a gland between the eighth and ninth segments 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 as courtship, attracting mates using sound or vibration. Like most other insects, moths pick up these sounds using tympanic membranes in their abdomens. An example is that of the polka-dot wasp moth (''Syntomeida epilais''), which produces sounds with a frequency above that normally detectable by humans (about 20 kHz). 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 is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
, 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 which the males apparently recognize as the flight signature of a potential mate. These flashes from the wings may attract several males that engage in aerial courtship displays.


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, from Ancient Greek τροφικός (''trophikos'') "pertaining to food or nourishment", may refer to: * Trophic cascade * Trophic coherence * Trophic egg * Trophic function * Trophic hormone * Trophic level index * Trophic level * Trop ...
relationships between
autotroph An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...
s and
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
s, which are included in the stages of Lepidoptera larvae, pupae, and adults. Larvae and pupae are links in the diets of birds and parasitic
entomophagous Entomophagy (, from Greek ἔντομον ', 'insect', and φαγεῖν ', 'to eat') is the practice of eating insects. An alternative term is insectivory. Terms for organisms that practice entomophagy are ''entomophage'' and ''insectivore' ...
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 is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
. Adult butterflies and moths are preyed upon by
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s,
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
s,
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s,
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s,
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
, and
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s. One spider species, ''
Argiope argentata ''Argiope argentata'', commonly known as the silver argiope due to the silvery color of its cephalothorax, is a member of the orb-weaver spider family Araneidae. This species resides in arid and warm environments in North America, Central America ...
'', eats butterflies and moths and exhibits a long bite when preying on them rather than wrapping them in silk first. This is theorized to serve as an immobilization tactic. Caterpillars and pupae fall prey not only to birds, but also to invertebrate predators and small mammals, as well as fungi and bacteria.
Parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
and
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...
wasps and
flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
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 largest predators. Lepidoptera, especially the immature stages, are an ecologically important food to many insectivorous birds, such as the
great tit The great tit (''Parus major'') is a passerine bird in the tit family Paridae. It is a widespread and common species throughout Europe, the Middle East, Central Asia and east across the Palearctic to the Amur River, south to parts of North Af ...
in Europe. An "
evolutionary arms race In evolutionary biology, an evolutionary arms race is an ongoing struggle between competing sets of co-evolving genes, phenotypic and behavioral traits that develop escalating adaptations and counter-adaptations against each other, resembling an a ...
" can be seen between predator and prey species. The Lepidoptera have developed a number of strategies for defense and protection, including evolution of morphological characters and changes in ecological lifestyles and behaviors. These include
aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
,
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
,
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
, and development of threat patterns and displays. Only a few birds, such as the
nightjar Nightjars are medium-sized nocturnal or crepuscular birds in the family Caprimulgidae and order Caprimulgiformes, characterised by long wings, short legs, and very short bills. They are sometimes called goatsuckers, due to the ancient folk ta ...
s, hunt nocturnal lepidopterans. Their main predators are
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
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. Lepidopteran eggs are also preyed upon. Some caterpillars, such as the
zebra swallowtail butterfly ''Eurytides marcellus'', the zebra swallowtail, (formerly listed under genera ''Protographium'', ''Iphiclides'', ''Graphium (butterfly), Graphium'' and ''Papilio'' by some authorities) is a swallowtail butterfly native to the eastern United Stat ...
larvae, are cannibalistic. Some species of Lepidoptera are poisonous to predators, such as the monarch butterfly in the Americas, ''
Atrophaneura Red-bodied swallowtails or ruby swallowtail ( due to the color ) are butterflies in the swallowtail family, that belong to the genera ''Atrophaneura'', ''Byasa'', ''Losaria'', or ''Pachliopta''. They are generally found in Asia (Indomalayan r ...
'' species (roses, windmills, etc.) in Asia, as well as ''
Papilio antimachus ''Papilio antimachus'', the African giant swallowtail, 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. The wings are long and narrow and th ...
'', and the
birdwing Birdwings are butterflies in the swallowtail family, that belong to the genera ''Trogonoptera'', ''Troides'', and ''Ornithoptera''. Most recent authorities recognise 36 species, however, this is debated, and some authorities include additional ...
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 species. A predator which 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 and striking patterns as an indicator to predators about their toxicity. This phenomenon is known as
aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
. (Abstract). Some caterpillars, especially members of
Papilionidae Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includes the larges ...
, contain an
osmeterium The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlik ...
, a Y-shaped protrusible
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...
found in the
prothoracic 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 (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on ea ...
segment of the larvae. When threatened, the caterpillar emits unpleasant smells from the organ to ward off the predators. Camouflage is also an important defense strategy, which involves the use of coloration or shape to blend into the surrounding environment. Some lepidopteran species blend with their surroundings, making them difficult to spot by predators. Caterpillars can exhibit shades of green that match its host plant. Others look like inedible objects, such as twigs or leaves. For instance, the
mourning cloak ''Nymphalis antiopa'', known as the mourning cloak in North America and the Camberwell beauty in United Kingdom, Britain, is a large butterfly native to Eurasia and North America. The immature form of this species is sometimes known as the spiny ...
fades into the backdrop of trees when it folds its wings back. 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 swall ...
'') and the
western tiger swallowtail ''Papilio rutulus'', the western tiger swallowtail, is a swallowtail butterfly belonging to the ''Papilionidae'' family. The species was first described by Hippolyte Lucas in 1852. Like the other tiger swallowtails, the western tiger swallowt ...
look like bird droppings. For example, adult
Sesiidae The Sesiidae or clearwing moths are a diurnal moth family in the order Lepidoptera known for their Batesian mimicry in both appearance and behaviour of various Hymenoptera. The family consists of 165 genera spread over two subfamilies, contai ...
species (also known as clearwing moths) have a general appearance sufficiently similar to a
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
or
hornet Hornets (insects in the genus ''Vespa'') are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to in length. They are distinguished from other vespine wasps by th ...
to make it likely the moths gain a reduction in predation by
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on bu ...
. Eyespots are a type of
automimicry In zoology, automimicry, Browerian mimicry, or intraspecific mimicry, is a form of mimicry in which the same species of animal is imitated. There are two different forms. In one form, first described by Lincoln Brower in 1967, weakly-defended m ...
used by some butterflies and moths. In butterflies, the spots are composed of concentric rings of scales in different colors. The proposed role of the eyespots is to deflect attention of predators. Their resemblance to eyes provokes the predator's instinct to attack these wing patterns.
Batesian Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on Bu ...
and Müllerian 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 in the example between the limenitidine
viceroy butterfly The viceroy (''Limenitis archippus'') is a North American butterfly. It was long thought to be a Batesian mimic of the monarch butterfly, but since the viceroy is also distasteful to predators, it is now considered a Müllerian mimic instead. ...
in relation to the inedible danaine monarch. 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 that need to learn about the insects' inedibility. Taxa from the toxic genus ''
Heliconius ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America a ...
'' form one of the most well-known Müllerian complexes. The adults of the various species now resemble each other so well, the species cannot be distinguished without close morphological observation and, in some cases, dissection or genetic analysis. Moths are able to hear the range emitted by bats, 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 The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, along with the other f ...
moth that cause it to drop a few inches in its flight to evade attack.
Tiger moth The de Havilland DH.82 Tiger Moth is a 1930s British biplane designed by Geoffrey de Havilland and built by the de Havilland Aircraft Company. It was operated by the Royal Air Force (RAF) and other operators as a primary trainer aircraft. ...
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 or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, some ...
(more specifically psychophily and phalaenophily for butterflies and moths, respectively), or the
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
of flowers. Most adult butterflies and moths feed on the
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
inside flowers, using their probosces to reach the nectar hidden at the base of the petals. In the process, the adults brush against the flowers'
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
s, on which the reproductive
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
is made and stored. The pollen is transferred on appendages on the adults, which fly to the next flower to feed and unwittingly deposit the pollen on the stigma of the next flower, where the pollen germinates and fertilizes the seeds. Flowers pollinated by butterflies tend to be large and flamboyant, pink or lavender in color, frequently having a landing area, and usually scented, as butterflies are typically day-flying. Since butterflies do not digest pollen (except for heliconid species,) 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 such as ''
Thymelicus flavus ''Thymelicus'' is a Palearctic genus in the skipper butterfly family, Hesperiidae. The genus is 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 or pollinator constancy is the tendency of individual pollinators to exclusively visit certain flower species or morphs within a species, bypassing other available flower species that could potentially contain more nectar. This typ ...
, which means 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 pollinator groups are the
hawk moth The Sphingidae are a family of moths (Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as “hornworms”; it includes about 1,450 species. It is best represented in the tropics, but ...
s of the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Sphingidae. Their behavior is similar to
hummingbird Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics aro ...
s, i.e., using rapid wing beats to hover in front of flowers. Most hawk moths are
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
or
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
, so moth-pollinated flowers (e.g., ''
Silene latifolia ''Silene latifolia'' subsp. ''alba'' (formerly ''Melandrium album''), the white campion is a dioecious flowering plant in the family Caryophyllaceae, native to most of Europe, Western Asia and Northern Africa. It is a herbaceous annual, occasio ...
'' ) tend to be white, night-opening, large, and showy with tubular corollae and a strong, sweet scent produced in the evening, night, or early morning. A lot of nectar is produced to fuel the high
metabolic rate Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
s needed to power their flight. Other moths (e.g., noctuids, geometrids, pyralids) 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 In ecology, a biological interaction is the effect that a pair of organisms living together in a community have on each other. They can be either of the same species (intraspecific interactions), or of different species (interspecific interaction ...
wherein each individual involved benefits in some way. An example of a mutualistic relationship would be that shared by yucca moths (Tegeculidae) and their host, yucca flowers (Asparagaceae). 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 Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s (e.g.
Lycaenidae Lycaenidae is the second-largest family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfl ...
). The larvae communicate with the ants using vibrations 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.


Parasitism

Only 42 species of
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
lepidopterans are known (1
Pyralidae The Pyralidae, commonly called pyralid moths, snout moths or grass moths, are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many (particularly older) classifications, the grass moths (Crambidae) are included in the Pyrali ...
; 40
Epipyropidae The Epipyropidae comprise 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 ...
). The larvae of the greater and
lesser wax moth The lesser wax moth (''Achroia grisella'') is a small moth of the snout moth family (Pyralidae) that belongs to the subfamily Galleriinae. The species was first described by Johan Christian Fabricius in 1794. Adults are about 0.5 inches (13  ...
s feed on the
honeycomb A honeycomb is a mass of Triangular prismatic honeycomb#Hexagonal prismatic honeycomb, hexagonal prismatic Beeswax, wax cells built by honey bees in their beehive, nests to contain their larvae and stores of honey and pollen. beekeeping, Beekee ...
inside
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
nests and may become pests; they are also found in
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
and
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
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 much as 80% of nests can be destroyed. Other parasitic larvae are known to prey upon
cicada The cicadas () are a superfamily, the Cicadoidea, of insects in the order Hemiptera (true bugs). They are in the suborder Auchenorrhyncha, along with smaller jumping bugs such as leafhoppers and froghoppers. The superfamily is divided into two ...
s and
leaf hopper A leafhopper is the common name for any species from the family Cicadellidae. These minute insects, colloquially known as hoppers, are plant feeders that suck plant sap from grass, shrubs, or trees. Their hind legs are modified for jumping, and a ...
s. File: Parasitism_in_Gypsy_moths.svg, center, 500px, The different parasitoids affecting the gypsy moth (''Lymantaria dispar''): The stage they affect and eventually kill and its duration are denoted by arrows. rect 11 106 222 135
Brachymeria intermedia ''Brachymeria'' is a genus of parasitic wasps in the family Chalcididae. Over 300 species are known worldwide, all of them parasites of insect pupae. Most species are black with limited yellow markings, and like most chalcidid wasps, they have e ...
rect 11 146 222 176 Coccygomimus instigator rect 3 515 222 535
Compsilura concinnata ''Compsilura concinnata'' (tachinid fly; order Diptera) is a parasitoid native to Europe that was introduced to North America in 1906 to control the population of an exotic forest, univoltine, spongy moth named ''Lymantria dispar''. It is an endo ...
rect 3 540 222 560
Parasetigena silvestris ''Parasetigena'' is a genus of flies in the family Tachinidae The Tachinidae are a large and variable family of true fly, flies within the insect order Fly, Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 ...
rect 185 600 385 650
Blepharipa pratensis ''Blepharipa'' is a genus of flies in the family Tachinidae. Species *'' B. albocinta'' (Mesnil, 1956) *'' B. carbonata'' (Mesnil, 1970) *'' B. chaetoparafacialis'' Chao, 1982 *'' B. fimbriata'' (Wulp, 1890) *'' B. fusiformis'' (Walker, 1849) * ...
rect 370 560 650 610
Aphantorhaphopsis samerensis ''Aphantorhaphopsis'' is a genus of flies in the family Tachinidae. Some consider this to be a subgenus of '' Siphona'', most European workers seem content that this is a genus in its own right. They are known from the Palearctic, Afrotropical ...
rect 370 612 650 630 Glyptapanteles liparidis rect 370 632 650 650
Meteorus pulchricornis ''Meteorus'' is a genus of parasitoid wasps in the family Braconidae. It comprises over 330 species worldwide. ''Meteorus'' wasps are distinguished from other braconid wasps by the presence of a second submarginal cell in the forewing and a pet ...
rect 600 100 770 150
Anastatus disparis ''Anastatus disparis'' is a species of egg parasitoid. Hosts include ''Lymantria dispar'' and ''Antheraea pernyi''. The species is sexually dimorphic, with 630 sex specific genes. Females have 11 antennal subsegments, whereas males have 8. Femal ...
rect 450 400 780 440
Cotesia melanoscelus ''Cotesia'' is a genus of braconid wasps first described by Peter Cameron in 1891. Some species parasitize caterpillars of species considered pests, and are used as biocontrol agents. '' Cotesia congregata'' parasitizes the tomato and the toba ...
rect 450 442 780 465 Glyptapanteles porthetriae rect 450 467 780 490
Hyposoter tricoloripes ''Hyposoter'' is a large cosmopolitan genus of parasitoid wasps belonging to the family Ichneumonidae. The genus has cosmopolitan distribution. Selected species * '' Hyposoter affinis'' (Cresson, 1864) * ''Hyposoter albicans'' (Brischke, 1880) ...
rect 450 492 780 520 Phobocampe disparis desc bottom-left
In reverse, moths and butterflies may be subject to
parasitic wasp Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causin ...
s and
flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
, 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, for the parasitic larvae to benefit the most. In another form of parasitism, koinobiont, the species live off their hosts while inside (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 Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describe ...
, and many
hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
n parasitoids. Some species may be subject to a variety of parasites, such as the
gypsy moth ''Lymantria dispar'', also known as the gypsy moth or the spongy moth, is a species of moth in the family Erebidae. ''Lymantria dispar'' is subdivided into several subspecies, with subspecies such as ''L. d. dispar'' and ''L. d. japonica'' bei ...
(''Lymantaria dispar''), which is attacked by a series of 13 species, in six different taxa throughout its life cycle. In response to a parasitoid egg or larva in the caterpillar's body, the plasmatocytes, or simply the host's cells can form a multilayered capsule that eventually causes the endoparasite to
asphyxiate Asphyxia or asphyxiation is a condition of deficient supply of oxygen to the body which arises from abnormal breathing. Asphyxia causes generalized hypoxia, which affects primarily the tissues and organs. There are many circumstances that can i ...
. The process, called encapsulation, 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 is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
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'' is a large genus of moths of the family Geometridae. There are hundreds of described species, found in all parts of the world (with 45 in the British Isles alone), and new species are discovered on a regular basis. ''Eupithecia'' ...
'' ). Those of the 15 species in ''Eupithecia'' that mirror inchworms, are the only known species of butterflies and moths that are ambush predators. Four species are known to eat snails. For example, the Hawaiian caterpillar (''
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. It has been called the snail-eating caterpil ...
'') uses silk traps, in a manner similar to that of spiders, to capture certain species of snails (typically
Tornatellides ''Tornatellides'' is a genus of minute, air-breathing land snails, terrestrial gastropod mollusks, or micromolluscs in the family Achatinellidae. Distribution This genus is endemic to the islands of the South Pacific. Predators Individuals of ...
). Larvae of some species of moths in the
Tineidae Tineidae is a family of moths in the order Lepidoptera described by Pierre André Latreille in 1810. Collectively, they are known as fungus moths or tineid moths. The family contains considerably more than 3,000 species in more than 300 genera. ...
,
Gelechiidae The Gelechiidae are a family of moths commonly referred to as twirler moths or gelechiid moths. They are the namesake family of the huge and little-studied superfamily Gelechioidea, and the family's taxonomy has been subject to considerable dis ...
, and
Noctuidae The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, along with the other f ...
, besides others, feed on
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
, or dead organic material, such as fallen leaves and fruit, fungi, and animal products, and turn it into
humus In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Lati ...
. Well-known species include the cloth moths (''
Tineola bisselliella ''Tineola'' is a genus of moths the family Tineidae. There are two species, including the familiar common clothes moth (''T. bisselliella'').Tinea pellionella ''Tinea pellionella'', the case-bearing clothes moth, is a species of tineoid moth in the family Tineidae, the fungus moths. This species has a cosmopolitan distribution, occurring nearly worldwide.Cheema, P. S. (1956).Studies on the bionomics o ...
'', and ''
Trichophaga tapetzella ''Trichophaga tapetzella'', the tapestry moth or carpet moth, is a moth of the family Tineidae. It is found worldwide. The wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. ...
''), which feed on detritus containing
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
, including
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
,
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier ...
s,
cobweb A spider web, spiderweb, spider's web, or cobweb (from the archaic word '' coppe'', meaning "spider") is a structure created by a spider out of proteinaceous spider silk extruded from its spinnerets, generally meant to catch its prey. Spi ...
s,
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 bl ...
s (particularly of
domestic pigeon The domestic pigeon (''Columba livia domestica'' or ''Columba livia'' ''forma'' ''domestica'') is a pigeon subspecies that was derived from the rock dove (also called the rock pigeon). The rock pigeon is the world's oldest domesticated bird. Me ...
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. In 2015 it was reported that wasp
bracovirus ''Bracovirus'' is a genus of viruses, in the family ''Polydnaviridae''. Bracoviruses are an ancient symbiotic virus contained in parasitic braconid wasps that evolved off of the ''nudivirus'' about 190 million years ago and has been evolving at ...
DNA was present in Lepidoptera such as monarch butterflies, silkworms and moths. These were described in some newspaper articles as examples of a naturally occurring
genetically engineered Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including t ...
insects.


Evolution and systematics


History of study

Linnaeus in ''
Systema Naturae ' (originally in Latin written ' with the ligature æ) is one of the major works of the Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the system, now known as binomial nomen ...
'' (1758) recognized three divisions of the Lepidoptera: ''Papilio'', ''Sphinx'' and ''
Phalaena ''Phalaena'' is an obsolete genus of 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 al ...
'', 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 Johann Nepomuk Cosmas Michael Denis, also: ''Sined the Bard'', (27 September 1729 – 29 September 1800) was an Austrian Catholic priest and Jesuit, who is best known as a poet, bibliographer, and lepidopterist. Life Denis was born at Schärding ...
&
Ignaz Schiffermüller Ignaz Schiffermüller (born 2 October 1727 in Hellmonsödt; died 21 June 1806 in Linz) was an Austrian naturalist mainly interested in Lepidoptera. Schiffermüller was a teacher at the Theresianum College in Vienna. His collection was presented ...
(1775),
Johan Christian Fabricius Johan Christian Fabricius (7 January 1745 – 3 March 1808) was a Danish zoologist, specialising in "Insecta", which at that time included all arthropods: insects, arachnids, crustaceans and others. He was a student of Carl Linnaeus, and is cons ...
(1775) and
Pierre André Latreille Pierre André Latreille (; 29 November 1762 – 6 February 1833) 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 ...
(1796).
Jacob Hübner Jacob Hübner (20 June 1761 – 13 September 1826, in Augsburg) was a German entomologist. He was the author of ''Sammlung Europäischer Schmetterlinge'' (1796–1805), a founding work of entomology. Scientific career Hübner was the author of '' ...
described many genera, and the lepidopteran genera were catalogued by
Ferdinand Ochsenheimer Ferdinand Ochsenheimer (17 March 1767 – 2 November 1822) was a German actor and entomologist (lepidopterist). Life Ochsenheimer was born and brought up in Mainz (then in the Electorate of Mainz) and began to show an interest in butterflies ...
and
Georg Friedrich Treitschke Georg Friedrich Treitschke (; 29 August 1776 – 4 June 1842) was a German libretto, librettist, translator and lepidopterist. He was born in Leipzig and died in Vienna. In 1800 he came to the Vienna Hofoper. From 1809 to 1814 he was principal ...
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 (17 December 1799 – 14 April 1874) was a German entomologist and physician. He was born, and died, in Regensburg. Herrich-Schäffer studied and collected particularly butterflies and moths (Lepidoptera). ...
(several volumes, 1843–1856), and
Edward Meyrick Edward Meyrick (25 November 1854, in Ramsbury – 31 March 1938, at Thornhanger, Marlborough) was an English schoolmaster and amateur entomologist. He was an expert on microlepidoptera and some consider him one of the founders of modern micr ...
(1895) based their classifications primarily on wing venation. Sir
George Francis Hampson Sir George Francis Hampson, 10th Baronet (14 January 1860 – 15 October 1936) was an English entomologist. Hampson studied at Charterhouse School and Exeter College, Oxford. He travelled to India to become a tea-planter in the Nilgiri Hills of ...
worked on the microlepidoptera during this period and
Philipp Christoph Zeller Philipp Christoph Zeller (8 April 1808 – 27 March 1883) was a German entomologist. Zeller was born at Steinheim an der Murr, Württemberg, two miles from Marbach, the birthplace of Schiller. The family moved to Frankfurt (Oder) where Philipp ...
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 (April 13, 1837 – May 17, 1911) was an American entomologist and paleontologist. He was a leading figure in entomology during his lifetime and the founder of insect paleontology in America. In addition to fossil insects, ...
(1837–1911), who worked on butterflies. He published a study of the Florissant deposits of Colorado, including the exceptionally preserved ''
Prodryas persephone ''Prodryas persephone'' is an extinct species of brush-footed butterfly, known from a single specimen from the Chadronian-aged Florissant Shale Lagerstatte of Late Eocene Colorado. ''P. persephone'' is the first fossil butterfly to be found in ...
''. 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 Emil Hans Willi Hennig (20 April 1913 – 5 November 1976) was a Germans, German biologist and zoologist who is considered the founder of Phylogenesis, phylogenetic systematics, otherwise known as cladistics. In 1945 as a POWs in World War II, p ...
(1913–1976) developed the
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among
monotrysia The Monotrysia are a group of moths in the lepidopteran order, not currently considered to be a natural group or clade. Apart from the recently discovered family Andesianidae, most of the group consists of small, relatively understudied species ...
n families and Kristensen worked more generally on insect
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
and higher Lepidoptera too. While it is often found that DNA-based phylogenies differ from those based on
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
, 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 In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
: Microlepidoptera 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 tends not to be as common as some other insects in habitats that are most conducive to fossilization, such as lakes and ponds; their juvenile stage has only the head capsule as a hard part that might be preserved. Lepidopteran bodies tend to come apart after death, and decompose quickly, so fossil remains are often extremely fragmentary. Of the fossils known, only an estimated 7% have been described.Sohn, JC., Labandeira, C.C. & Davis, D.R
The fossil record and taphonomy of butterflies and moths (Insecta, Lepidoptera): implications for evolutionary diversity and divergence-time estimates.
BMC Evol Biol 15, 12 (2015). https://doi.org/10.1186/s12862-015-0290-8
The location and abundance of the most common moth species are indicative that mass migrations of moths occurred over the Palaeogene
North Sea The North Sea lies between Great Britain, Norway, Denmark, Germany, the Netherlands and Belgium. An epeiric sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian S ...
, 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 are also seen in fossil leaves, although the interpretation of them is tricky. Putative fossil stem group representatives of
Amphiesmenoptera Amphiesmenoptera is an insect superorder, established by S. G. Kiriakoff, but often credited to Willi Hennig in his revision of insect taxonomy for two sister orders: Lepidoptera (butterflies and moths) and Trichoptera (caddisflies). In 2017, a t ...
(the clade comprising Trichoptera and Lepidoptera) are known from the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
. The earliest known lepidopteran fossils are fossilized scales from the Triassic-Jurassic boundary. They were found as rare
palynological Palynology is the "study of dust" (from grc-gre, παλύνω, palynō, "strew, sprinkle" and ''-logy'') or of "particles that are strewn". A classic palynologist analyses particulate samples collected from the air, from water, or from deposits ...
elements in the sediments of the Triassic-Jurassic boundary from the cored Schandelah-1 well, drilled near
Braunschweig Braunschweig () or Brunswick ( , from Low German ''Brunswiek'' , Braunschweig dialect: ''Bronswiek'') is a city in Lower Saxony, Germany, north of the Harz Mountains at the farthest navigable point of the river Oker, which connects it to the Nor ...
in northern
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. This pushes back the fossil record and origin of
glossata Glossata (Fabricius, 1775) is the suborder of the insect order Lepidoptera that contains most lepidopteran species and includes all the superfamilies of moths and butterflies that have a coilable proboscis. (See also the suborders Zeugloptera, A ...
n lepidopterans by about 70 million years, supporting molecular estimates of a
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
(ca 212 million years) divergence of glossatan and non-glossatan lepidopterans. The findings were reported in 2018 in the journal
Science Advances ''Science Advances'' is a peer-reviewed multidisciplinary open-access scientific journal established in early 2015 and published by the American Association for the Advancement of Science. The journal's scope includes all areas of science, incl ...
. The authors of the study proposed that lepidopterans evolved a proboscis as an adaptation to drink from droplets and thin films of water for maintaining their
fluid balance Fluid balance is an aspect of the homeostasis of organisms in which the amount of water in the organism needs to be controlled, via osmoregulation and behavior, such that the concentrations of electrolytes (salts in solution) in the various body ...
in the hot and arid
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologic ...
of the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
. The earliest named lepidopteran taxon is '' Archaeolepis mane'', a primitive moth-like species from the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
, dated back to around , and known only from three wings found in the
Charmouth Mudstone The Charmouth Mudstone Formation is a geological formation in England. It preserves fossils dating back to the early part of the Jurassic period (Sinemurian–Pliensbachian). It forms part of the lower Lias Group. It is most prominently exposed at ...
of
Dorset Dorset ( ; archaically: Dorsetshire , ) is a county in South West England on the English Channel coast. The ceremonial county comprises the unitary authority areas of Bournemouth, Christchurch and Poole and Dorset (unitary authority), Dors ...
, UK. The wings show scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared with
Trichoptera The caddisflies, or order Trichoptera, are a group of insects with aquatic larvae and terrestrial adults. There are approximately 14,500 described species, most of which can be divided into the suborders Integripalpia and Annulipalpia on the b ...
(caddisflies). Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
, which all belong to primitive moth-like families. Many more fossils are found from the Tertiary, and particularly the
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than 1 ...
. The oldest genuine butterflies of the superfamily Papilionoidea have been found in the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
MoClay or
Fur Formation The Fur Formation is a marine geological formation of Ypresian (Lower Eocene Epoch, c. 56.0-54.5 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 ...
of
Denmark ) , song = ( en, "King Christian stood by the lofty mast") , song_type = National and royal anthem , image_map = EU-Denmark.svg , map_caption = , subdivision_type = Sovereign state , subdivision_name = Danish Realm, Kingdom of Denmark ...
. The best preserved fossil lepidopteran is the Eocene ''
Prodryas persephone ''Prodryas persephone'' is an extinct species of brush-footed butterfly, known from a single specimen from the Chadronian-aged Florissant Shale Lagerstatte of Late Eocene Colorado. ''P. persephone'' is the first fossil butterfly to be found in ...
'' from the
Florissant Fossil Beds The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones and ...
.


Phylogeny

Lepidoptera and
Trichoptera The caddisflies, or order Trichoptera, are a group of insects with aquatic larvae and terrestrial adults. There are approximately 14,500 described species, most of which can be divided into the suborders Integripalpia and Annulipalpia on the b ...
(caddisflies) are
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
s, sharing many similarities that are lacking in others; for example the females of both orders are
heterogametic Heterogametic sex (digametic sex) refers to the individuals of a species in which the sex chromosomes are not the same. For example, in humans, males with an X and a Y sex chromosome would be referred to as the heterogametic sex, and females ...
, meaning they have two different
sex chromosomes A sex chromosome (also referred to as an allosome, heterotypical chromosome, gonosome, heterochromosome, or idiochromosome) is a chromosome that differs from an ordinary autosome in form, size, and behavior. The human sex chromosomes, a typical ...
, 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 in the two orders have mouth structures and glands with which they make and manipulate
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
.
Willi Hennig Emil Hans Willi Hennig (20 April 1913 – 5 November 1976) was a Germans, German biologist and zoologist who is considered the founder of Phylogenesis, phylogenetic systematics, otherwise known as cladistics. In 1945 as a POWs in World War II, p ...
grouped the two orders into the superorder
Amphiesmenoptera Amphiesmenoptera is an insect superorder, established by S. G. Kiriakoff, but often credited to Willi Hennig in his revision of insect taxonomy for two sister orders: Lepidoptera (butterflies and moths) and Trichoptera (caddisflies). In 2017, a t ...
; together they are sister to the extinct order
Tarachoptera Tarachoptera is an extinct order of insects, currently solely known from the mid Cretaceous aged Burmese amber. It belongs to Amphiesmenoptera alongside living Lepidopterans (butterflies and moths) and Trichoptera (caddiesflies), but is outside t ...
. Lepidoptera descend from a diurnal moth-like common ancestor that either fed on dead or living plants. The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
, based on a 2008 DNA and
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
analysis, shows the order as a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, sister to the Trichoptera, and more distantly related to the
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
(true flies) and
Mecoptera Mecoptera (from the Greek: ''mecos'' = "long", ''ptera'' = "wings") is an order of insects in the superorder Endopterygota with about six hundred species in nine families worldwide. Mecopterans are sometimes called scorpionflies after their lar ...
(scorpionflies).
Micropterigidae Micropterigoidea is the superfamily of "mandibulate archaic moths", all placed in the single family Micropterigidae, containing currently about twenty living genera. They are considered the most primitive extant lineage of lepidoptera (Kristense ...
,
Agathiphagidae ''Agathiphaga'' is a genus of moths, known as kauri moths. It is the only living in the family Agathiphagidae. This caddisfly-like lineage of primitive moths was first reported by Lionel Jack Dumbleton in 1952, as a new genus of Micropterigidae ...
and
Heterobathmiidae ''Heterobathmia'' is a genus of Lepidoptera. It is the only genus in the suborder Heterobathmiina, as well as in the superfamily Heterobathmioidea and in the family Heterobathmiidae. Primitive, day-flying, metallic moths confined to southern So ...
are the oldest and most basal lineages of Lepidoptera. The adults of these families do not have the curled tongue or
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
, that are found in most members of the order, but instead have chewing
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s adapted for a special diet. Micropterigidae larvae feed on
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
,
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
, or
liverworts The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of ge ...
(much like the
Trichoptera The caddisflies, or order Trichoptera, are a group of insects with aquatic larvae and terrestrial adults. There are approximately 14,500 described species, most of which can be divided into the suborders Integripalpia and Annulipalpia on the b ...
). 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'', also known as the southern beeches, is a genus of 43 species of trees and shrubs native to the Southern Hemisphere in southern South America (Chile, Argentina) and Australasia (east and southeast Australia, New Zealand, New Gui ...
'', 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 is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
. The
Eriocraniidae Eriocraniidae is a family of moths restricted to the Holarctic region, with six extant genera. These small, metallic moths are usually day-flying, emerging fairly early in the northern temperate spring. They have a proboscis with which they drin ...
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 A leaf miner is any one of numerous species of insects in which the larval stage lives in, and eats, the leaf tissue of plants. The vast majority of leaf-mining insects are moths (Lepidoptera), sawflies (Symphyta, the mother clade of wasps), ...
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 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 (in contra ...
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 In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
, or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia,
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 comm ...
and
Gelechioidea __NOTOC__ Gelechioidea (from the type genus ''Gelechia'', "keeping to the ground") is the superfamily of moths that contains the case-bearers, twirler moths, and relatives, also simply called curved-horn moths or gelechioid moths. It is a large a ...
are mostly leaf miners, but more recent lineages feed externally. In the
Tineoidea Tineoidea is the ditrysian superfamily of moths that includes clothes moths, Bagworm moth, bagworms and relatives. There are six families usually included within it, Eriocottidae, Arrhenophanidae, Lypusidae, Acrolophidae, Tineidae and Psychidae, ...
, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage. The
Yponomeutoidea Yponomeutoidea is a superfamily of ermine moths and relatives. There are about 1,800 species of Yponomeutoids worldwide, most of them known to come from temperate regions. This superfamily is one of the earliest groups to evolve external feeding ...
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 In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
in the mid-
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
, and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together. In the so-called "
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 artificial. However, it seems that by moving some taxa about, a monoph ...
", 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 A tympanal organ (or tympanic organ) is a hearing organ in insects, consisting of a membrane ( tympanum) stretched across a frame backed by an air sac and associated sensory neurons. Sounds vibrate the membrane, and the vibrations are sensed by ...
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 is the superfamily of noctuid (Latin "night owl") or "owlet" moths, and has more than 70,000 described species, the largest number of for any Lepidopteran superfamily. Its classification has not yet reached a satisfactory or stable st ...
,
Bombycoidea Bombycoidea is a superfamily of moths. It contains the silk moths, emperor moths, sphinx moths, and relatives. The Lasiocampoidea The Lasiocampidae are a family of moths also known as eggars, tent caterpillars, snout moths (although this also ...
,
Lasiocampidae The Lasiocampidae are a family of moths also known as eggars, tent caterpillars, snout moths (although this also refers to the Pyralidae), or lappet moths. Over 2,000 species occur worldwide, and probably not all have been named or studied. It i ...
,
Mimallonoidea Mimallonidae Burmeister (mimallonids), sometimes known as "sack-bearer" moths for the larval case-building behavior, are a family of Lepidoptera containing over 300 named species in 43 genera. These moths are found only in the New World, with mos ...
,
Geometroidea The Geometroidea are the superfamily of geometrid moths in the order Lepidoptera. It includes the families Geometridae, Uraniidae, Epicopeiidae, Sematuridae, and the recently established family Pseudobistonidae. The monotypic genus ''Apoprogones ...
and
Rhopalocera Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...
. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group. The Rhopalocera, comprising the
Papilionoidea The superfamily Papilionoidea (from the genus ''Papilio'', meaning "butterfly") contains all the butterflies except for the moth-like Hedyloidea. The members of the Papilionoidea may be distinguished by the following combination of characters: ...
(butterflies), Hesperioidea (skippers), and the
Hedyloidea Hedylidae, the "American moth-butterflies", is a Family (biology), family of insects in the order Lepidoptera, representing the superfamily Hedyloidea. They have traditionally been viewed as an Extant taxon, extant sister group of the butterfly s ...
(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 Nomenclature (, ) is a system of names or terms, or the rules for forming these terms in a particular field of arts or sciences. The principles of naming vary from the relatively informal naming conventions, conventions of everyday speech to the i ...
. 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 their ancestors. The rest of the moths and butterflies make up ninety-eight percent of the other taxa, making
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 (in contra ...
. More recently, findings of new taxa, larvae and pupa have aided in detailing the relationships of primitive taxa, phylogenetic analysis showing the primitive lineages to be
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
compared to the rest of Lepidoptera lineages. Recently, lepidopterists have abandoned clades like suborders, and those between orders and superfamilies. *
Zeugloptera Micropterigoidea is the superfamily of "mandibulate archaic moths", all placed in the single family Micropterigidae, containing currently about twenty living genera. They are considered the most primitive extant lineage of lepidoptera (Kristense ...
is a clade with
Micropterigoidea Micropterigoidea is the superfamily of "mandibulate archaic moths", all placed in the single family Micropterigidae, containing currently about twenty living genera. They are considered the most primitive extant lineage of lepidoptera (Kristense ...
being its only superfamily, containing the single family Micropterigidae. Species of Micropterigoidea are practically living fossils, being one of the most primitive lepidopteran groups, still retaining chewing mouthparts (mandibles) in adults, unlike other clades of butterflies and moths. About 120 species are known worldwide, with more than half the species in the genus ''
Micropteryx ''Micropterix'' is a genus of small primitive metallic moths, in the insect order lepidoptera within the family Micropterigidae. The name was raised by the German entomologist, Jacob Hübner in 1825 and comes from the Greek for ''mikros'', ...
'' in the Palearctic region. There are only two known in North America (''
Epimartyria ''Epimartyria'' is a genus of small primitive metallic moths in the family Micropterigidae. Species *''Epimartyria auricrinella'' Walsingham, 1898 *'' Epimartyria bimaculella'' Davis & Landry, 2012 *''Epimartyria pardella ''Epimartyria pardell ...
''), with many more being found in Asia and the southwest Pacific, particularly New Zealand with about 50 species. *
Aglossata ''Agathiphaga'' is a genus of moths, known as kauri moths. It is the only living in the family Agathiphagidae. This caddisfly-like lineage of primitive moths was first reported by Lionel Jack Dumbleton in 1952, as a new genus of Micropterigidae. ...
is the second most primitive
lineage Lineage may refer to: Science * Lineage (anthropology), a group that can demonstrate its common descent from an apical ancestor or a direct line of descent from an ancestor * Lineage (evolution), a temporal sequence of individuals, populati ...
of Lepidoptera; being first described in 1952 by
Lionel Jack Dumbleton Lionel Jack Dumbleton (1905 – 25 September 1976) was a New Zealand entomologist. He was born in Hampden, New Zealand and was a founding member of the Entomological Society of New Zealand. One of his most remarkable biological discoveries ...
.
Agathiphagidae ''Agathiphaga'' is a genus of moths, known as kauri moths. It is the only living in the family Agathiphagidae. This caddisfly-like lineage of primitive moths was first reported by Lionel Jack Dumbleton in 1952, as a new genus of Micropterigidae ...
is the only family in Aglossata and contains two species in its only genus, ''
Agathiphaga ''Agathiphaga'' is a genus of moths, known as kauri moths. It is the only living in the family Agathiphagidae. This caddisfly-like lineage of primitive moths was first reported by Lionel Jack Dumbleton in 1952, as a new genus of Micropterigidae ...
''. ''
Agathiphaga queenslandensis ''Agathiphaga queenslandensis'' is a moth of the family Agathiphagidae. 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 o ...
'' and ''
Agathiphaga vitiensis ''Agathiphaga vitiensis'', or the Fiji kauri moth, is a moth of the family Agathiphagidae. It is found from Fiji to Vanuatu and Solomon Islands Solomon Islands is an island country consisting of six major islands and over 900 smalle ...
'' are found along the north-eastern coast of
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, and in
Fiji Fiji ( , ,; fj, Viti, ; Fiji Hindi: फ़िजी, ''Fijī''), officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists ...
to
Vanuatu Vanuatu ( or ; ), officially the Republic of Vanuatu (french: link=no, République de Vanuatu; bi, Ripablik blong Vanuatu), is an island country located in the South Pacific Ocean. The archipelago, which is of volcanic origin, is east of no ...
and the
Solomon Islands Solomon Islands is an island country consisting of six major islands and over 900 smaller islands in Oceania, to the east of Papua New Guinea and north-west of Vanuatu. It has a land area of , and a population of approx. 700,000. Its capita ...
, respectively. *
Heterobathmiina ''Heterobathmia'' is a genus of Lepidoptera. It is the only genus in the suborder Heterobathmiina, as well as in the superfamily Heterobathmioidea and in the family Heterobathmiidae. Primitive, day-flying, metallic moths confined to southern So ...
was first described by Kristensen and Nielsen in 1979.
Heterobathmiidae ''Heterobathmia'' is a genus of Lepidoptera. It is the only genus in the suborder Heterobathmiina, as well as in the superfamily Heterobathmioidea and in the family Heterobathmiidae. Primitive, day-flying, metallic moths confined to southern So ...
is the only family and includes about 10 species, which are day-flying, metallic moths, confined to southern South America, the adults eat the pollen of ''
Nothofagus ''Nothofagus'', also known as the southern beeches, is a genus of 43 species of trees and shrubs native to the Southern Hemisphere in southern South America (Chile, Argentina) and Australasia (east and southeast Australia, New Zealand, New Gui ...
'' or southern beech and the larvae mine the leaves. *
Glossata Glossata (Fabricius, 1775) is the suborder of the insect order Lepidoptera that contains most lepidopteran species and includes all the superfamilies of moths and butterflies that have a coilable proboscis. (See also the suborders Zeugloptera, A ...
contains a majority of the species, with the most obvious difference being non-functioning mandibles, and elongated maxillary galeae or the proboscis. The
basal clade In phylogenetics, basal is the direction of the ''base'' (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to th ...
s 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 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 (in contra ...
, which contains 98% of all described species in Lepidoptera.


Relationship to people


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" as an art and fictional character. In many cultures the soul of a dead person is associated with the butterfly, for example in
Ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cult ...
, where the word for butterfly ψυχή (psyche) also means ''
soul In many religious and philosophical traditions, there is a belief that a soul is "the immaterial aspect or essence of a human being". Etymology The Modern English noun ''soul'' is derived from Old English ''sāwol, sāwel''. The earliest attes ...
'' and ''
breath Breathing (or ventilation) is the process of moving air into and from the lungs to facilitate gas exchange with the internal environment, mostly to flush out carbon dioxide and bring in oxygen. All aerobic creatures need oxygen for cellular ...
''. In
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
, as in Ancient Greece, the word for "butterfly"
papilio ''Papilio'' is a genus in the swallowtail butterfly family, Papilionidae, as well as the only representative of the tribe Papilionini. The word ''papilio'' is Latin for butterfly. It includes the common yellow swallowtail (''Papilio machaon''), ...
was associated with the soul of the dead. The skull-like marking on the thorax of the
death's-head hawkmoth The name death's-head hawkmoth refers to any of three moth species of the genus ''Acherontia'' (''Acherontia atropos'', '' Acherontia styx'' and '' Acherontia lachesis''). The former species is found in Europe and throughout Africa, the latter ...
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 Buñuel is a town and municipality located in the province and autonomous community of Navarre, northern Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto ...
and Dalí) and '' The Silence of the Lambs'', and in the artwork of the Japanese metal band
Sigh Paralanguage, also known as vocalics, is a component of meta-communication that may modify meaning, give nuanced meaning, or convey emotion, by using techniques such as prosody, pitch, volume, intonation, etc. It is sometimes defined as relatin ...
's album ''
Hail Horror Hail ''Hail Horror Hail'' is the third studio album by the band Sigh. It was released by Cacophonous Records originally in 1997. The album is the band's first full-length release to feature an avant-garde/experimental sound not normally associated wi ...
''. According to '' Kwaidan: Stories and Studies of Strange Things'', by
Lafcadio Hearn , born Patrick Lafcadio Hearn (; el, Πατρίκιος Λευκάδιος Χέρν, Patríkios Lefkádios Chérn, Irish language, Irish: Pádraig Lafcadio O'hEarain), was an Irish people, Irish-Greeks, Greek-Japanese people, Japanese writer, t ...
, a butterfly was seen in Japan as the
personification Personification occurs when a thing or abstraction is represented as a person, in literature or art, as a type of anthropomorphic metaphor. The type of personification discussed here excludes passing literary effects such as "Shadows hold their b ...
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 Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, bu ...
screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad
omen An omen (also called ''portent'') is a phenomenon that is believed to foretell the future, often signifying the advent of change. It was commonly believed in ancient times, and still believed by some today, that omens bring divine messages fr ...
s. When
Taira no Masakado was a Heian period provincial magnate (''gōzoku'') and samurai based in eastern Japan, notable for leading the first recorded uprising against the central government in Kyōto. Early life Masakado was one of the sons of Taira no Yoshimasa ...
was secretly preparing for his famous revolt, there appeared in
Kyoto Kyoto (; Japanese: , ''Kyōto'' ), officially , is the capital city of Kyoto Prefecture in Japan. Located in the Kansai region on the island of Honshu, Kyoto forms a part of the Keihanshin metropolitan area along with Osaka and Kobe. , the ci ...
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 is a historical region and cultural area in southern North America and most of Central America. It extends from approximately central Mexico through Belize, Guatemala, El Salvador, Honduras, Nicaragua, and northern Costa Rica. W ...
n city of
Teotihuacan Teotihuacan (Spanish language, Spanish: ''Teotihuacán'') (; ) is an ancient Mesoamerican city located in a sub-valley of the Valley of Mexico, which is located in the State of Mexico, northeast of modern-day Mexico City. Teotihuacan is ...
, the brilliantly colored image of the butterfly was carved into many temples, buildings, jewelry, and emblazoned on incense burners in particular. The butterfly was sometimes depicted with the maw of a
jaguar The jaguar (''Panthera onca'') is a large cat species and the only living member of the genus '' Panthera'' native to the Americas. With a body length of up to and a weight of up to , it is the largest cat species in the Americas and the th ...
and some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies to
fire Fire is the rapid oxidation of a material (the fuel) in the exothermic chemical process of combustion, releasing heat, light, and various reaction Product (chemistry), products. At a certain point in the combustion reaction, called the ignition ...
and warfare persisted through to the
Aztec civilization The Aztecs () were a Mesoamerican culture that flourished in central Mexico in the post-classic period from 1300 to 1521. The Aztec people included different ethnic groups of central Mexico, particularly those groups who spoke the Nahuatl l ...
and evidence of similar jaguar-butterfly images has been found among the Zapotec, and
Maya civilization The Maya civilization () of the Mesoamerican people is known by its ancient temples and glyphs. Its Maya script is the most sophisticated and highly developed writing system in the pre-Columbian Americas. It is also noted for its art, archit ...
s.


Pests

The
larva A larva (; plural larvae ) 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. The ...
e of many lepidopteran species are major pests in
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
. Some of the major pests include
Tortricidae The Tortricidae are a family of moths, commonly known as tortrix moths or leafroller moths, in the order Lepidoptera. This large family has over 11,000 species described, and is the sole member of the superfamily Tortricoidea, although the genu ...
,
Noctuidae The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, along with the other f ...
, and
Pyralidae The Pyralidae, commonly called pyralid moths, snout moths or grass moths, are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many (particularly older) classifications, the grass moths (Crambidae) are included in the Pyrali ...
. The larvae of the Noctuidae genus ''
Spodoptera ''Spodoptera'' is a genus of moths of the family Noctuidae erected by Achille Guenée in 1852. Many are known as pest insects. The larvae are sometimes called armyworms. The roughly thirty species are distributed across six continents. Descri ...
'' (armyworms), ''
Helicoverpa ''Helicoverpa'' is a genus of moths in the family Noctuidae first described by David F. Hardwick in 1965. Some species are among the worst Lepidopteran agricultural pests in the world, and three species (''H. armigera'', ''H. zea'', and ''H. pu ...
'' (corn earworm), or ''
Pieris brassicae ''Pieris brassicae'', the large white, also called cabbage butterfly, cabbage white, cabbage moth (erroneously), or in India the large cabbage white, is a butterfly in the family Pieridae. It is a close relative of the small white, ''Pieris ra ...
'' can cause extensive damage to certain crops. ''
Helicoverpa zea ''Helicoverpa zea, ''commonly known as the corn earworm, is a species (formerly in the genus ''Heliothis'') in the family Noctuidae. The larva of the moth ''Helicoverpa zea'' is a major agricultural pest. Since it is polyphagous (feeds on many ...
'' larvae (cotton bollworms or tomato fruitworms) are
polyphagous Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...
, meaning they eat a variety of crops, including
tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
es and
cotton Cotton is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus ''Gossypium'' in the mallow family Malvaceae. The fiber is almost pure cellulose, and can contain minor perce ...
. ''
Peridroma saucia ''Peridroma saucia'', the pearly underwing or variegated cutworm, is a moth of the family Noctuidae. The species was first described by Jacob Hübner in 1808. It is found in North and South America, Europe, Asia and Africa. The variegated cutwor ...
'' (variegated cutworms) are described as one of the most damaging pests to gardens, with the ability to destroy entire gardens and fields in a matter of days. 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, making 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 can create many problems for agriculture, where many caterpillars can affect 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 wasps and flies. For example, ''
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 lar ...
'', a fly which deposited larvae feed upon the pupae of the
forest tent caterpillar moth The forest tent caterpillar moth (''Malacosoma disstria'') is a moth found throughout North America, especially in the eastern regions. Unlike related tent caterpillar species, the larvae of forest tent caterpillars do not make tents, but rather ...
. 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 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, as the red rubber septa in the picture, is encased in a convent ...
s. A pheromone trap is a type of
insect trap Insect traps are used to monitor or directly reduce populations of insects or other arthropods, by trapping individuals and killing them. They typically use food, visual lures, chemical attractants and pheromones as bait and are installed so that ...
that uses
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 like hormones outside the body of the secreting individual, to affect the behavio ...
s to lure
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
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
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
s would naturally eat
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
containing
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
, such as
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
s or
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier ...
s. Well known species are cloth moths ('' T. bisselliella'', '' T. pellionella'', and '' T. tapetzella''), feeding on foodstuffs that people find economically important, such as
cotton Cotton is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus ''Gossypium'' in the mallow family Malvaceae. The fiber is almost pure cellulose, and can contain minor perce ...
,
linen Linen () is a textile made from the fibers of the flax plant. Linen is very strong, absorbent, and dries faster than cotton. Because of these properties, linen is comfortable to wear in hot weather and is valued for use in garments. It also ...
,
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
and wool
fabric Textile is an umbrella term that includes various fiber-based materials, including fibers, yarns, filaments, threads, different fabric types, etc. At first, the word "textiles" only referred to woven fabrics. However, weaving is not the ...
s as well as
fur Fur is a thick growth of hair that covers the skin of mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an insulating blanket t ...
s; furthermore they have been found on shed
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier ...
s and
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
,
bran Bran, also known as miller's bran, is the hard outer layers of Cereal, cereal grain. It consists of the combined aleurone and pericarp. Corn (maize) bran also includes the pedicel (tip cap). Along with cereal germ, germ, it is an integral pa ...
, semolina and flour (possibly preferring wheat flour), biscuits, casein, and
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
specimens in museums.


Beneficial insects

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 Bombyx mori, domesticated silkworm moth (''Bombyx mori''), the larvae of which make their cocoons out of
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
, which can be spun into cloth. Silk is and has been an important economic resource History of silk, throughout history. 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'', and ''Antheraea'' 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 noxious weed, weeds in place of herbicides. The
pyralid The Pyraloidea (pyraloid moths or snout moths) 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, and as such, they have be ...
cactus moth was introduced from Argentina to Australia, where it successfully suppressed millions of acres of Opuntia, prickly pear cactus. Another species of the Pyralidae, called the alligator weed stem borer (''Arcola malloi''), was used to control the aquatic plant known as alligator weed (''Alternanthera philoxeroides'') in conjunction with the Agasicles hygrophila, alligator weed flea beetle; in this case, the two insects work in synergy and the weed rarely recovers. Breeding butterflies and moths, or butterfly gardening/rearing, has become an ecologically viable process of introducing species into the ecosystem to benefit it. Butterfly ranching in Papua New Guinea permits nationals of that country to "farm" economically valuable insect species for the collectors market in an ecologically sustainable manner.


Food

Lepidoptera feature prominently in entomophagy as food items on almost every continent. While in most cases, adults, larvae or pupae are eaten as staples by indigenous people, beondegi or silkworm
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
e are eaten as a snack in Korean cuisineRobinson, Martin; Bartlett, Ray and Whyte, Rob (2007) ''Korea''. Lonely Planet publications, . (pg 63) while Maguey worm is considered a delicacy in Mexico. In some parts of La Huasteca, Huasteca, the silk nests of the Madrone butterfly are maintained on the edge of roof tops of houses for consumption. In the Carnia region of Italy, children catch and eat Crop (anatomy), ingluvies of the toxic ''Zygaena (genus), Zygaena'' moths in early summer. The ingluvies, despite having a very low cyanogenic content, serve as a convenient, supplementary source of sugar to the children who can include this resource as a seasonal delicacy at minimum risk. Outside of this instance, adult Lepidoptera are rarely consumed by humans, with the sole exception of the Bogong moth.


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 toxins in them and species from approximately 12 families of moths or butterflies worldwide can inflict serious human injuries (urticarial dermatitis and atopic asthma to osteochondritis, consumption coagulopathy, renal failure, and brain, intracerebral hemorrhage). Skin rashes are the most common, but there have been fatalities. ''Lonomia'' is a frequent cause of envenomation 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 keratoconjunctivitis. The sharp barbs on the end of caterpillar hairs can get lodged in soft tissues and mucous 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.. Free full text.


See also

* Comparison of butterflies and moths * Lepidoptera in the 10th edition of Systema Naturae * McGuire Center for Lepidoptera and Biodiversity, University of Florida * Societas Europaea Lepidopterologica


Lists

* Lists of Lepidoptera by region * Taxonomy of the Lepidoptera


References


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''. . ; Bibliography *


External links


Historic Moth illustrations

Lepidoptera
at Insects (Insecta) of the World *
Caught Between the Pages: Treasures from the Franclemont Collection
Online virtual exhibit featuring a selection of historic entomological writings and images from the Comstock Library of Entomology at Cornell University ; Regional sites


Butterflies of Bulgaria

Butterflies of Canada

Photography of European Butterflies and Moths



Butterflies of India





Japmoth
Japanese moths. Access images via the numbers on the left.
Butterflies and Moths in the Netherlands

Butterflies and Moths of Northern Ireland







Swedish Moths and Butterflies Lepidoptera (English)

Butterflies of Turkey
{{Authority control Lepidoptera, Articles containing video clips Extant Early Jurassic first appearances Insect orders Pliensbachian first appearances Taxa named by Carl Linnaeus Insects in culture Amphiesmenoptera