Butterflies Of Oceania
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Butterflies are
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 pa ...
s in the
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 monophy ...
n
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 ter ...
Rhopalocera from the
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 ...
Lepidoptera Lepidoptera ( ) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families and 46 superfamilies, 10 percent of the total described speci ...
, which also includes
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. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises the large
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
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 character ...
, which contains at least one former group, the skippers (formerly the superfamily "Hesperioidea"), and the most recent analyses suggest it also contains the moth-butterflies (formerly the superfamily "Hedyloidea"). Butterfly fossils date to the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
, about 56 million years ago. Butterflies have a four-stage life cycle, as like most insects 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. ...
. Winged adults lay eggs on the food plant on which their
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. ...
e, known as
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 Sy ...
s, will feed. The caterpillars grow, sometimes very rapidly, and when fully developed,
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 thei ...
te in 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 thei ...
. When
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 ...
is complete, the pupal skin splits, the adult insect climbs out, and after its wings have expanded and dried, it flies off. Some butterflies, especially in the tropics, have several generations in a year, while others have a single generation, and a few in cold locations may take several years to pass through their entire life cycle. Butterflies are often polymorphic, and many species make use of
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 b ...
,
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
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 ...
to evade their predators. Some, like the
monarch A monarch is a head of stateWebster's II New College DictionarMonarch Houghton Mifflin. Boston. 2001. p. 707. Life tenure, for life or until abdication, and therefore the head of state of a monarchy. A monarch may exercise the highest authority ...
and the
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. ...
, migrate over long distances. Many butterflies are attacked by
parasite 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 h ...
s or
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 ...
s, including
wasps 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. T ...
,
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
ns,
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 m ...
, and other invertebrates, or are preyed upon by other organisms. Some species are pests because in their larval stages they can damage domestic crops or trees; other species are agents of
pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
of some plants. Larvae of a few butterflies (e.g.,
harvesters Harvester may refer to: Agriculture and forestry * Combine harvester, a machine commonly used to harvest grain crops * Forage harvester, a machine used to harvest forage * Harvester (forestry), a type of heavy vehicle employed in cut-to-length log ...
) eat harmful insects, and a few are predators 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 ...
s, while others live as
mutualists Mutualism describes the ecological Biological interaction, interaction between two or more species where each species has a net benefit. Mutualism is a common type of ecological interaction. Prominent examples include most vascular plants engag ...
in association with ants. Culturally, butterflies are a popular motif in the visual and literary arts. The
Smithsonian Institution The Smithsonian Institution ( ), or simply the Smithsonian, is a group of museums and education and research centers, the largest such complex in the world, created by the U.S. government "for the increase and diffusion of knowledge". Found ...
says "butterflies are certainly one of the most appealing creatures in nature".


Etymology

The ''
Oxford English Dictionary The ''Oxford English Dictionary'' (''OED'') is the first and foundational historical dictionary of the English language, published by Oxford University Press (OUP). It traces the historical development of the English language, providing a c ...
'' derives the word straightforwardly 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 settlers in the mid-5th ...
''butorflēoge'', butter-fly; similar names in
Old Dutch In linguistics, Old Dutch (Dutch: Oudnederlands) or Old Low Franconian (Dutch: Oudnederfrankisch) is the set of Franconian dialects (i.e. dialects that evolved from Frankish) spoken in the Low Countries during the Early Middle Ages, from aro ...
and
Old High German Old High German (OHG; german: Althochdeutsch (Ahd.)) is the earliest stage of the German language, conventionally covering the period from around 750 to 1050. There is no standardised or supra-regional form of German at this period, and Old Hig ...
show that the name is ancient, but modern Dutch and German use different words (''vlinder'' and ''Schmetterling'') and the common name often varies substantially between otherwise closely-related languages. A possible source of the name is the bright yellow male of the brimstone (''
Gonepteryx rhamni ''Gonepteryx rhamni'' (known as the common brimstone) is a butterfly of the family Pieridae. It lives throughout the Palearctic zone and is commonly found across Europe, Asia, and North Africa. Across much of its range, it is the only species ...
''); another is that butterflies were on the wing in meadows during the spring and summer butter season while the grass was growing.


Paleontology

The earliest
Lepidoptera Lepidoptera ( ) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families and 46 superfamilies, 10 percent of the total described speci ...
fossils date to 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 per ...
-
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
boundary, around 200 million years ago. Butterflies evolved from moths, so while the butterflies are
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 gr ...
(forming a single
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 ter ...
), the moths are not. The oldest known butterfly is '' Protocoeliades kristenseni'' from the
Palaeocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
aged
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 ...
of Denmark, approximately 55 million years old, which belongs to the family
Hesperiidae Skippers are a family of the Lepidoptera (moths and butterflies) named the Hesperiidae. Being diurnal, they are generally called butterflies. They were previously placed in a separate superfamily, Hesperioidea; however, the most recent taxonomy ...
(skippers).
Molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleo ...
estimates suggest that butterflies originated sometime 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 ...
, but only significantly diversified during the Cenozoic. The oldest American butterfly is the
Late Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
''
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, approximately 34 million years old. File:Prodryas.png, ''
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 ...
'', a
Late Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
butterfly from the Florissant Fossil Beds, 1887 engraving File:PZSL1889Plate31 Fossil Papilionid Butterfly Lithopsyche antiqua from Early Oligocene Bembridge Marls.png, ''
Lithopsyche antiqua ''Lithopsyche'' is a fossil butterfly known from Oligocene-aged strata of the Isle of Wight, England. The sole specimen is too incomplete to allow a certain assignment of a family (biology), family, but it was placed on its description as a geom ...
'', an
Early Oligocene The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/ Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chattian ...
butterfly from the Bembridge Marls,
Isle of Wight The Isle of Wight ( ) is a county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the largest and second-most populous island of England. Referred to as 'The Island' by residents, the Is ...
, 1889 engraving


Taxonomy and phylogeny

Traditionally, butterflies have been divided into the
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
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 character ...
excluding the smaller groups of the
Hesperiidae Skippers are a family of the Lepidoptera (moths and butterflies) named the Hesperiidae. Being diurnal, they are generally called butterflies. They were previously placed in a separate superfamily, Hesperioidea; however, the most recent taxonomy ...
(skippers) and the more moth-like
Hedylidae Hedylidae, the "American moth-butterflies", is a family of insects in the order Lepidoptera, representing the superfamily Hedyloidea. They have traditionally been viewed as an extant sister group of the butterfly superfamily Papilionoidea. In 19 ...
of America.
Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
analysis suggests that the traditional Papilionoidea is
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
with respect to the other two groups, so they should both be included within Papilionoidea, to form a single butterfly group, thereby synonymous with the clade Rhopalocera.


Biology


General description

Butterfly adults are characterized by their four scale-covered wings, which give the Lepidoptera their name (
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
λεπίς lepís, scale + πτερόν pterón, wing). These scales give butterfly wings their colour: they are pigmented with
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
s that give them blacks and browns, as well as
uric acid Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates, such as ammonium acid urate. Uric acid is a product of the metabolic breakdown ...
derivatives and
flavones Flavones (from Latin ''flavus'' "yellow") are a class of flavonoids based on the backbone of 2-phenylchromen-4-one (2-phenyl-1- benzopyran-4-one) (as shown in the first image of this article). Flavones are common in foods, mainly from spices, ...
that give them yellows, but many of the blues, greens, reds and iridescent colours are created by
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 wi ...
produced by the micro-structures of the scales and hairs. As in all insects, the body is divided into three sections: the head,
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the c ...
, and
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
. The thorax is composed of three segments, each with a pair of legs. In most families of butterfly the antennae are clubbed, unlike those of
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 which may be threadlike or feathery. The long proboscis can be coiled when not in use for sipping nectar from flowers. Nearly all butterflies are diurnal, have relatively bright colours, and hold their wings vertically above their bodies when at rest, unlike the majority of moths which fly by night, are often cryptically coloured (well camouflaged), and either hold their wings flat (touching the surface on which the moth is standing) or fold them closely over their bodies. Some day-flying moths, such as the
hummingbird hawk-moth The hummingbird hawk-moth (''Macroglossum stellatarum'') is a species of hawk moth found across temperate regions of Eurasia. The species is named for its similarity to hummingbirds, as they feed on the nectar of tube-shaped flowers using thei ...
, are exceptions to these rules. Butterfly
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. ...
e,
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 Sy ...
s, have a hard (
sclerotised Sclerotin is a component of the cuticle of various Arthropoda, most familiarly insects. It is formed by cross-linking members of particular classes of protein molecules, a biochemical process called sclerotization, a form of tanning in which qu ...
) head with strong mandibles used for cutting their food, most often leaves. They have cylindrical bodies, with ten segments to the abdomen, generally with short prolegs on segments 3–6 and 10; the three pairs of true legs on the thorax have five segments each. Many are well camouflaged; others are aposematic with bright colours and bristly projections containing toxic chemicals obtained from their food plants. 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 thei ...
or chrysalis, unlike that of moths, is not wrapped in a cocoon. Many butterflies are
sexually dimorphic 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 ...
. Most butterflies have the
ZW sex-determination system The ZW sex-determination system is a chromosomal system that determines the sex of offspring in birds, some fish and crustaceans such as the giant river prawn, some insects (including butterflies and moths), the schistosome family of flatworms ...
where females are the heterogametic sex (ZW) and males homogametic (ZZ).


Distribution and migration

Butterflies are distributed worldwide except Antarctica, totalling some 18,500 species. Of these, 775 are
Nearctic The Nearctic realm is one of the eight biogeographic realms constituting the Earth's land surface. The Nearctic realm covers most of North America, including Greenland, Central Florida, and the highlands of Mexico. The parts of North America ...
; 7,700
Neotropical 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 ...
; 1,575
Palearctic The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Sib ...
; 3,650
Afrotropical The Afrotropical realm is one of Earth's eight biogeographic realms. It includes Africa south of the Sahara Desert, the majority of the Arabian Peninsula, the island of Madagascar, southern Iran and extreme southwestern Pakistan, and the island ...
; and 4,800 are distributed across the combined
Oriental The Orient is a term for the East in relation to Europe, traditionally comprising anything belonging to the Eastern world. It is the antonym of '' Occident'', the Western World. In English, it is largely a metonym for, and coterminous with, the ...
and
Australian Australian(s) may refer to: Australia * Australia, a country * Australians, citizens of the Commonwealth of Australia ** European Australians ** Anglo-Celtic Australians, Australians descended principally from British colonists ** Aboriginal A ...
/
Oceania Oceania (, , ) is a geographical region that includes Australasia, Melanesia, Micronesia, and Polynesia. Spanning the Eastern and Western hemispheres, Oceania is estimated to have a land area of and a population of around 44.5 million ...
regions. 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. ...
is native to the Americas, but in the nineteenth century or before, spread across the world, and is now found in Australia, New Zealand, other parts of Oceania, and the
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, def ...
. It is not clear how it dispersed; adults may have been blown by the wind or larvae or pupae may have been accidentally transported by humans, but the presence of suitable host plants in their new environment was a necessity for their successful establishment. Many butterflies, such as the
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. ...
, monarch, and several danaine migrate for long distances. These migrations take place over a number of generations and no single individual completes the whole trip. The eastern North American population of monarchs can travel thousands of miles south-west to overwintering sites in Mexico. There is a reverse migration in the spring. It has recently been shown that the British painted lady undertakes a 9,000-mile round trip in a series of steps by up to six successive generations, from tropical Africa to the Arctic Circle — almost double the length of the famous migrations undertaken by monarch. Spectacular large-scale migrations associated with the
monsoon 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 also using stable hydrogen isotopes. Butterflies navigate using a time-compensated sun compass. They can see polarized light and therefore orient even in cloudy conditions. The polarized light near the ultraviolet spectrum appears to be particularly important. Many migratory butterflies live in semi-arid areas where breeding seasons are short. The life histories of their host plants also influence butterfly behaviour.


Life cycle

Butterflies in their adult stage can live from a week to nearly a year depending on the species. Many species have long larval life stages while others can remain dormant in their pupal or egg stages and thereby survive winters. The
Melissa Arctic ''Oeneis melissa'', the Melissa Arctic, is a species of butterfly in the family Nymphalidae. The wingspan is 42–51 mm. The larvae feed on various sedges, including ''Carex bigelowii'' and ''Carex rupestris''. Subspecies *''Oeneis mel ...
(''Oeneis melissa'') overwinters twice as a caterpillar. Butterflies may have one or more broods per year. The number of generations per year varies from
temperate 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 ...
to tropical regions with tropical regions showing a trend towards multivoltinism.
Courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...
is often aerial and often involves
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. Butterflies then land on the ground or on a perch to mate. Copulation takes place tail-to-tail and may last from minutes to hours. Simple photoreceptor cells located at the genitals are important for this and other adult behaviours. The male passes a
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophore ...
to the female; to reduce sperm competition, he may cover her with his scent, or in some species such as the Apollos (''
Parnassius ''Parnassius'' is a genus of northern circumpolar and montane (alpine and Himalayan) butterflies usually known as Apollos or snow Apollos. They can vary in colour and form significantly based on their altitude. They also show an adaptation to h ...
'') plugs her genital opening to prevent her from mating again. The vast majority of butterflies have a four-stage life cycle; egg,
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. ...
(caterpillar),
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 thei ...
(chrysalis) and
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 ...
(adult). In the genera ''
Colias ''Colias'' is a genus of butterflies in the family Pieridae. They are often called clouded yellows; the North American name "sulphurs" is elsewhere used for Coliadinae in general. The closest living relative is the genus '' Zerene'', which is so ...
'', ''
Erebia ''Erebia'' is a Holarctic genus of brush-footed butterflies, family Nymphalidae. Most of the about 90–100 species (see also below) are dark brown or black in color, with reddish-brown to orange or more rarely yellowish wing blotches or ban ...
'', ''
Euchloe ''Euchloe'' is a genus of pierid butterflies from the orangetip tribe (Anthocharini). They are Holarctic in distribution, with most species in Europe, Central Asia, and North America. Like other Anthocharini, the American species are usually c ...
'', and ''Parnassius'', a small number of species are known that reproduce semi-parthenogenetically; when the female dies, a partially developed larva emerges from her abdomen.


Egg

Butterfly eggs are protected by a hard-ridged outer layer of shell, called the ''chorion''. This is lined with a thin coating of wax which prevents the egg from drying out before the larva has had time to fully develop. Each egg contains a number of tiny funnel-shaped openings at one end, called ''micropyles''; the purpose of these holes is to allow sperm to enter and fertilize the egg. Butterfly eggs vary greatly in size and shape between species, but are usually upright and finely sculptured. Some species lay eggs singly, others in batches. Many females produce between one hundred and two hundred eggs. Butterfly eggs are fixed to a leaf with a special glue which hardens rapidly. As it hardens it contracts, deforming the shape of the egg. This glue is easily seen surrounding the base of every egg forming a meniscus. The nature of the glue has been little researched but in the case of ''
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 ...
'', it begins as a pale yellow granular secretion containing acidophilic proteins. This is viscous and darkens when exposed to air, becoming a water-insoluble, rubbery material which soon sets solid. Butterflies in the genus '' Agathymus'' do not fix their eggs to a leaf, instead the newly laid eggs fall to the base of the plant. Eggs are almost invariably laid on plants. Each species of butterfly has its own host plant range and while some species of butterfly are restricted to just one species of plant, others use a range of plant species, often including members of a common family. In some species, such as the
great spangled fritillary The great spangled fritillary (''Speyeria cybele'') is a North American butterfly of the family Nymphalidae. Description Its wingspan ranges from . It is characterized by its orange color above with five black dashes near forewing base and sev ...
, the eggs are deposited close to but not on the food plant. This most likely happens when the egg overwinters before hatching and where the host plant loses its leaves in winter, as do violets in this example. The egg stage lasts a few weeks in most butterflies, but eggs laid close to winter, especially in temperate regions, go through a
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 I ...
(resting) stage, and the hatching may take place only in spring. Some temperate region butterflies, such as the
Camberwell beauty ''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 ...
, lay their eggs in the spring and have them hatch in the summer.


Caterpillar larva

Butterfly larvae, or caterpillars, consume plant leaves and spend practically all of their time searching for and eating food. Although most caterpillars are herbivorous, a few species are predators: ''
Spalgis epius ''Spalgis epius'', the apefly, is a small butterfly found in the Indomalayan realm that belongs to the lycaenids or blues family. It gets its name from the supposed resemblance of its pupa to the face of an ape. Description Male Upperside: du ...
'' eats
scale insect 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 th ...
s, while lycaenids such as '' Liphyra brassolis'' are
myrmecophilous Myrmecophily ( , ) is the term applied to positive interspecies associations between ants and a variety of other organisms, such as plants, other arthropods, and fungi. Myrmecophily refers to mutualistic associations with ants, though in its ...
, eating ant larvae. Some larvae, especially those of 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 butterf ...
, form mutual associations with ants. They communicate with the ants using vibrations that are transmitted through the substrate as well as using chemical signals. The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions.
Large blue The large blue (''Phengaris arion'') is a species of butterfly in the family Lycaenidae. The species was first defined in 1758 and first recorded in Britain in 1795. In 1979 the species became mostly extinct in Britain but has been successfully ...
(''Phengaris arion'') caterpillars trick ''
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, a ...
'' ants into taking them back to the
ant colony An ant colony is a population of a single ant species capable to maintain its complete lifecycle. Ant colonies are eusocial, communal, and efficiently organized and are very much like those found in other social Hymenoptera, though the vario ...
where they feed on the ant eggs and larvae in a parasitic relationship. Caterpillars mature through a series of developmental stages known as
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 ...
s. Near the end of each stage, the larva undergoes a process called apolysis, mediated by the release of a series of neurohormones. During this phase, the
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
, a tough outer layer made of a mixture of
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
and specialized
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, res ...
s, is released from the softer
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rel ...
beneath, and the epidermis begins to form a new cuticle. At the end of each instar, the larva moults, the old cuticle splits and the new cuticle expands, rapidly hardening and developing pigment. Development of butterfly wing patterns begins by the last larval instar. Caterpillars have short antennae and several
simple eyes A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
. The
mouthparts Mouthparts may refer to: * The parts of a mouth ** Arthropod mouthparts The mouthparts of arthropods have evolved into a number of forms, each adapted to a different style or mode of feeding. Most mouthparts represent modified, paired append ...
are adapted for chewing with powerful mandibles and a pair of maxillae, each with a segmented palp. Adjoining these is the labium-hypopharynx which houses a tubular spinneret which is able to extrude silk. Caterpillars such as those in the genus '' Calpodes'' (family Hesperiidae) have a specialized tracheal system on the 8th segment that function as a primitive lung. Butterfly caterpillars have three pairs of true legs on the thoracic segments and up to six 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 arising from the abdominal segments. These prolegs have rings of tiny hooks called crochets that are engaged hydrostatically and help the caterpillar grip the substrate. The epidermis bears tufts of
seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
e, the position and number of which help in identifying the species. There is also decoration in the form of hairs, wart-like protuberances, horn-like protuberances and spines. Internally, most of the body cavity is taken up by the gut, but there may also be large silk glands, and special glands which secrete distasteful or toxic substances. The developing wings are present in later stage instars and the
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sp ...
s start development in the egg stage.


Pupa

When the larva is fully grown, hormones such as
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) are produced. At this point the larva stops feeding, and begins "wandering" in the quest for a suitable pupation site, often the underside of a leaf or other concealed location. There it spins a button of silk which it uses to fasten its body to the surface and moults for a final time. While some caterpillars spin a cocoon to protect the pupa, most species do not. The naked pupa, often known as a chrysalis, usually hangs head down from the cremaster, a spiny pad at the posterior end, but in some species a silken girdle may be spun to keep the pupa in a head-up position. Most of the tissues and cells of the larva are broken down inside the pupa, as the constituent material is rebuilt into the imago. The structure of the transforming insect is visible from the exterior, with the wings folded flat on the ventral surface and the two halves of the proboscis, with the antennae and the legs between them. The pupal transformation into a butterfly through
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 ...
has held great appeal to mankind. To transform from the miniature wings visible on the outside of the pupa into large structures usable for flight, the pupal wings undergo rapid mitosis and absorb a great deal of nutrients. If one wing is surgically removed early on, the other three will grow to a larger size. In the pupa, the wing forms a structure that becomes compressed from top to bottom and pleated from proximal to distal ends as it grows, so that it can rapidly be unfolded to its full adult size. Several boundaries seen in the adult colour pattern are marked by changes in the expression of particular transcription factors in the early pupa.


Adult

The reproductive stage of the insect is the winged adult or
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 ...
. The surface of both butterflies and moths is covered by scales, each of which is an outgrowth from a single
epidermal The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
cell. The head is small and dominated by the two large
compound eyes 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 distin ...
. These are capable of distinguishing flower shapes or motion but cannot view distant objects clearly. Colour perception is good, especially in some species in the blue/violet range. The antennae are composed of many segments and have clubbed tips (unlike moths that have tapering or feathery antennae). The sensory receptors are concentrated in the tips and can detect odours. Taste receptors are located on the palps and on the feet. The mouthparts are adapted to sucking and the
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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
s are usually reduced in size or absent. The first maxillae are elongated into a tubular
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 ...
which is curled up at rest and expanded when needed to feed. The first and second maxillae bear palps which function as sensory organs. Some species have a reduced proboscis or maxillary palps and do not feed as adults. Many ''
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 ...
'' butterflies also use their proboscis to feed on pollen; in these species only 20% of the amino acids used in reproduction come from larval feeding, which allow them to develop more quickly as caterpillars, and gives them a longer lifespan of several months as adults. The thorax of the butterfly is devoted to locomotion. Each of the three thoracic segments has two legs (among nymphalids, the first pair is reduced and the insects walk on four legs). The second and third segments of the thorax bear the wings. The leading edges of the forewings have thick veins to strengthen them, and the hindwings are smaller and more rounded and have fewer stiffening veins. The forewings and hindwings are not hooked together ( as they are in moths) but are coordinated by the friction of their overlapping parts. The front two segments have a pair of spiracles which are used in respiration. The abdomen consists of ten segments and contains the gut and genital organs. The front eight segments have spiracles and the terminal segment is modified for reproduction. The male has a pair of clasping organs attached to a ring structure, and during copulation, a tubular structure is extruded and inserted into the female's vagina. A
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophore ...
is deposited in the female, following which the sperm make their way to a seminal receptacle where they are stored for later use. In both sexes, the genitalia are adorned with various spines, teeth, scales and bristles, which act to prevent the butterfly from mating with an insect of another species. After it emerges from its pupal stage, a butterfly cannot fly until the wings are unfolded. A newly emerged butterfly needs to spend some time inflating its wings with
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 letting them dry, during which time it is extremely vulnerable to predators.


Pattern formation

The colorful patterns on many butterfly wings tell potential predators that they are toxic. Hence, the genetic basis of wing
pattern formation The science of pattern formation deals with the visible, ( statistically) orderly outcomes of self-organization and the common principles behind similar patterns in nature. In developmental biology, pattern formation refers to the generation of ...
can illuminate both the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of butterflies as well as their
developmental biology Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of st ...
. The color of butterfly wings is derived from tiny structures called scales, each of which have their own
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 compou ...
s. In ''
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 ...
'' butterflies, there are three types of scales: yellow/white, black, and red/orange/brown scales. Some mechanism of wing pattern formation are now being solved using genetic techniques. For instance, a
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
called ''cortex'' determines the color of scales: deleting ''cortex'' turned black and red scales yellow. Mutations, e.g. transposon insertions of the non-coding DNA around the ''cortex'' gene can turn a black-winged butterfly into a butterfly with a yellow wing band.


Behaviour

Butterflies feed primarily on
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 mutualist ...
from flowers. Some also derive nourishment from
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 gametop ...
, tree sap, rotting fruit, dung, decaying flesh, and dissolved minerals in wet sand or dirt. Butterflies are important as pollinators for some species of plants. In general, they do not carry as much pollen load as
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 monophyletic lineage within the superfamil ...
s, but they are capable of moving pollen over greater distances.
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 ...
has been observed for at least one species of butterfly. Adult butterflies consume only liquids, ingested through the proboscis. They sip water from damp patches for hydration and feed on nectar from flowers, from which they obtain sugars for energy, and
sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable ...
and other minerals vital for reproduction. Several species of butterflies need more sodium than that provided by nectar and are attracted by sodium in salt; they sometimes land on people, attracted by the salt in human sweat. Some butterflies also visit dung and scavenge rotting fruit or carcasses to obtain minerals and nutrients. In many species, this
mud-puddling Mud-puddling, or simply puddling, is a behaviour most conspicuous in butterflies, but occurs in other animals as well, mainly insects; they seek out nutrients in certain moist substances such as rotting plant matter, mud and carrion and they suck ...
behaviour is restricted to the males, and studies have suggested that the nutrients collected may be provided as a
nuptial gift A nuptial gift is a nutritional gift given by one partner in some animals' sexual reproduction practices. Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simpl ...
, along with the spermatophore, during mating. In hilltopping, males of some species seek hilltops and ridge tops, which they patrol in search for females. Since it usually occurs in species with low population density, it is assumed these landscape points are used as meeting places to find mates. Butterflies use their antennae to sense the air for wind and scents. The antennae come in various shapes and colours; the hesperiids have a pointed angle or hook to the antennae, while most other families show knobbed antennae. The antennae are richly covered with sensory organs known as sensillae. A butterfly's sense of taste is coordinated by chemoreceptors on the tarsi, or feet, which work only on contact, and are used to determine whether an egg-laying insect's offspring will be able to feed on a leaf before eggs are laid on it. Many butterflies use chemical signals,
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; some have specialized scent scales (
androconia The external morphology of Lepidoptera is the physiological structure of the bodies of insects belonging to the order Lepidoptera, also known as butterflies and moths. Lepidoptera are distinguished from other orders by the presence ...
) or other structures (
coremata ''Coremata'' is a monotypic moth genus of the family Crambidae described by Hans Georg Amsel in 1956. It contains only one species, ''Coremata stigmatalis'', described by George Hampson in 1899, which is found in São Paulo, Brazil. The wingspan ...
or "hair pencils" in the Danaidae). Vision is well developed in butterflies and most species are sensitive to the ultraviolet spectrum. Many species show sexual dimorphism in the patterns of UV reflective patches. Colour vision may be widespread but has been demonstrated in only a few species. Some butterflies have organs of hearing and some species make
stridulatory Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fish, snakes and spiders. The mech ...
and clicking sounds. Many species of butterfly maintain territories and actively chase other species or individuals that may stray into them. Some species will bask or perch on chosen perches. The flight styles of butterflies are often characteristic and some species have courtship flight displays. Butterflies can only fly when their temperature is above ; when it is cool, they can position themselves to expose the underside of the wings to the sunlight to heat themselves up. If their body temperature reaches , they can orientate themselves with the folded wings edgewise to the sun. Basking is an activity which is more common in the cooler hours of the morning. Some species have evolved dark wingbases to help in gathering more heat and this is especially evident in alpine forms. As in many other insects, the
lift Lift or LIFT may refer to: Physical devices * Elevator, or lift, a device used for raising and lowering people or goods ** Paternoster lift, a type of lift using a continuous chain of cars which do not stop ** Patient lift, or Hoyer lift, mobil ...
generated by butterflies is more than can be accounted for by steady-state, non-transitory
aerodynamics Aerodynamics, from grc, ἀήρ ''aero'' (air) + grc, δυναμική (dynamics), is the study of the motion of air, particularly when affected by a solid object, such as an airplane wing. It involves topics covered in the field of fluid dy ...
. Studies using ''
Vanessa atalanta ''Vanessa atalanta'', the red admiral or, previously, the red admirable, is a well-characterized, medium-sized butterfly with black wings, red bands, and white spots. It has a wingspan of about . It was first described by Carl Linnaeus in his ...
'' in a
wind tunnel Wind tunnels are large tubes with air blowing through them which are used to replicate the interaction between air and an object flying through the air or moving along the ground. Researchers use wind tunnels to learn more about how an aircraft ...
show that they use a wide variety of aerodynamic mechanisms to generate force. These include wake capture, vortices at the wing edge, rotational mechanisms and the Weis-Fogh '
clap-and-fling Insects are the only group of invertebrates that have evolved wings and flight. Insects first flew in the Carboniferous, some 350 to 400 million years ago, making them the first animals to evolve flight. Wings may have evolved from appenda ...
' mechanism. Butterflies are able to change from one mode to another rapidly.


Ecology


Parasitoids, predators, and pathogens

Butterflies are threatened in their early stages by
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 ...
s and in all stages by predators, diseases and environmental factors.
Braconid The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis ...
and other parasitic wasps lay their eggs in lepidopteran eggs or larvae and the wasps' parasitoid larvae devour their hosts, usually pupating inside or outside the desiccated husk. Most wasps are very specific about their host species and some have been used as biological controls of pest butterflies like the large white butterfly. When the small cabbage white was accidentally introduced to New Zealand, it had no natural enemies. In order to control it, some pupae that had been parasitised by a chalcid wasp were imported, and natural control was thus regained. Some flies lay their eggs on the outside of caterpillars and the newly hatched fly larvae bore their way through the skin and feed in a similar way to the parasitoid wasp larvae. Predators of butterflies include ants, spiders, wasps, and birds. Caterpillars are also affected by a range of bacterial, viral and fungal diseases, and only a small percentage of the butterfly eggs laid ever reach adulthood. The bacterium ''
Bacillus thuringiensis ''Bacillus thuringiensis'' (or Bt) is a gram-positive, soil-dwelling bacterium, the most commonly used biological pesticide worldwide. ''B. thuringiensis'' also occurs naturally in the gut of caterpillars of various types of moths and butterf ...
'' has been used in sprays to reduce damage to crops by the caterpillars of the large white butterfly, and the
entomopathogenic fungus An entomopathogenic fungus is a fungus that can kill or seriously disable insects. Typical life cycle These fungi usually attach to the external body surface of insects in the form of microscopic spores (usually asexual, mitosporic spores also ...
''
Beauveria bassiana ''Beauveria bassiana'' is a fungus that grows naturally in soils throughout the world and acts as a parasite on various arthropod species, causing white muscardine disease; it thus belongs to the entomopathogenic fungi. It is used as a biologic ...
'' has proved effective for the same purpose.


Endangered species

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 ...
is the largest butterfly in the world. The species is
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and in ...
, and is one of only three insects (the other two being butterflies as well) to be listed on Appendix I of
CITES CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of intern ...
, making international trade illegal.CITES appendices I, II and III
, official website
Black grass-dart butterfly ''(Ocybadistes knightorum)'' is a butterfly of the family
Hesperiidae Skippers are a family of the Lepidoptera (moths and butterflies) named the Hesperiidae. Being diurnal, they are generally called butterflies. They were previously placed in a separate superfamily, Hesperioidea; however, the most recent taxonomy ...
. It is endemic to
New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...
. It has a very limited distribution in the Boambee area.


Defences

Butterflies protect themselves from predators by a variety of means. Chemical defences are widespread and are mostly based on chemicals of plant origin. In many cases the plants themselves evolved these toxic substances as
protection Protection is any measure taken to guard a thing against damage caused by outside forces. Protection can be provided to physical objects, including organisms, to systems, and to intangible things like civil and political rights. Although th ...
against herbivores. Butterflies have evolved mechanisms to sequester these plant toxins and use them instead in their own defence. These defence mechanisms are effective only if they are well advertised; this has led to the evolution of bright colours in unpalatable butterflies (
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 ...
). This signal is commonly mimicked by other butterflies, usually only females. A Batesian mimic imitates another species to enjoy the protection of that species' aposematism. The
common Mormon ''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 swallo ...
of India has female morphs which imitate the unpalatable red-bodied swallowtails, the
common rose ''Pachliopta aristolochiae'', the common rose, is a swallowtail butterfly belonging to the genus ''Pachliopta'', the roses, or red-bodied swallowtails. It is a common butterfly which is extensively distributed across south and southeast Asia. ...
and the crimson rose.
Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Mülleria ...
occurs when aposematic species evolve to resemble each other, presumably to reduce predator sampling rates; ''
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 ...
'' butterflies from the Americas are a good example.
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 b ...
is found in many butterflies. Some like the oakleaf butterfly and autumn leaf are remarkable imitations of leaves. As caterpillars, many defend themselves by freezing and appearing like sticks or branches. Others have
deimatic Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey anima ...
behaviours, such as rearing up and waving their front ends which are marked with eyespots as if they were snakes. Some papilionid caterpillars such as the giant swallowtail (''
Papilio cresphontes The giant swallowtail (''Papilio cresphontes'') is the largest butterfly in North America. It is abundant through many parts of eastern North America; populations from western North America and down into Panama are now (as of 2014) considered t ...
'') resemble bird droppings so as to be passed over by predators. Some caterpillars have hairs and bristly structures that provide protection while others are gregarious and form dense aggregations. Some species are
myrmecophile Myrmecophily ( , ) is the term applied to positive interspecies associations between ants and a variety of other organisms, such as plants, other arthropods, and fungi. Myrmecophily refers to mutualistic associations with ants, though in its m ...
s, forming mutualistic associations with
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 ...
s and gaining their protection. Behavioural defences include perching and angling the wings to reduce shadow and avoid being conspicuous. Some female Nymphalid butterflies guard their eggs from parasitoidal
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 ...
s. The Lycaenidae have a false head consisting of eyespots and small tails (false antennae) to deflect attack from the more vital head region. These may also cause ambush predators such as spiders to approach from the wrong end, enabling the butterflies to detect attacks promptly. Many butterflies have eyespots on the wings; these too may deflect attacks, or may serve to attract mates. Auditory defences can also be used, which in the case of the
grizzled skipper ''Pyrgus'' is a genus in the skippers butterfly family, Hesperiidae, known as the grizzled skippers. The name "checkered" or "chequered skipper" may also be applied to some species, but also refers to species in the genera '' Burnsius'' and '' ...
refers to vibrations generated by the butterfly upon expanding its wings in an attempt to communicate with ant predators. Many tropical butterflies have seasonal forms for dry and wet seasons. These are switched by the hormone
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. E ...
. The dry-season forms are usually more cryptic, perhaps offering better camouflage when vegetation is scarce. Dark colours in wet-season forms may help to absorb solar radiation. Butterflies without defences such as toxins or mimicry protect themselves through a flight that is more bumpy and unpredictable than in other species. It is assumed this behavior makes it more difficult for predators to catch them, and is caused by the
turbulence In fluid dynamics, turbulence or turbulent flow is fluid motion characterized by chaotic changes in pressure and flow velocity. It is in contrast to a laminar flow, which occurs when a fluid flows in parallel layers, with no disruption between ...
created by the small whirlpools formed by the wings during flight.


Declining numbers

Declining butterfly populations have been noticed in many areas of the world, and this phenomenon is consistent with the rapidly decreasing insect populations around the world. At least in the Western United States, this collapse in the number of most species of butterflies has been determined to be driven by global climate change, specifically, by warmer autumns.


In culture


In art and literature

Butterflies have appeared in art from 3500 years ago in ancient Egypt. 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. Wit ...
n city of
Teotihuacan Teotihuacan ( 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 known today as ...
, the brilliantly coloured image of the butterfly was carved into many temples, buildings, jewellery, and emblazoned on incense burners. 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 with fire and warfare persisted into the
Aztec civilisation 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 ...
; evidence of similar jaguar-butterfly images has been found among the Zapotec and Maya civilisations. Butterflies are widely used in objects of art and jewellery: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and used in some mixed media artworks and furnishings. The
Norwegian Norwegian, Norwayan, or Norsk may refer to: *Something of, from, or related to Norway, a country in northwestern Europe * Norwegians, both a nation and an ethnic group native to Norway * Demographics of Norway *The Norwegian language, including ...
naturalist Kjell Sandved compiled a photographic '' Butterfly Alphabet'' containing all 26 letters and the numerals 0 to 9 from the wings of butterflies. Sir
John Tenniel Sir John Tenniel (; 28 February 182025 February 1914)Johnson, Lewis (2003), "Tenniel, John", ''Grove Art Online, Oxford Art Online'', Oxford University Press. Web. Retrieved 12 December 2016. was an English illustrator, graphic humorist and poli ...
drew a famous illustration of
Alice Alice may refer to: * Alice (name), most often a feminine given name, but also used as a surname Literature * Alice (''Alice's Adventures in Wonderland''), a character in books by Lewis Carroll * ''Alice'' series, children's and teen books by ...
meeting a
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 Sy ...
for
Lewis Carroll Charles Lutwidge Dodgson (; 27 January 1832 – 14 January 1898), better known by his pen name Lewis Carroll, was an English author, poet and mathematician. His most notable works are '' Alice's Adventures in Wonderland'' (1865) and its sequ ...
's ''
Alice in Wonderland ''Alice's Adventures in Wonderland'' (commonly ''Alice in Wonderland'') is an 1865 English novel by Lewis Carroll. It details the story of a young girl named Alice who falls through a rabbit hole into a fantasy world of anthropomorphic creatur ...
'', c. 1865. The caterpillar is seated on a toadstool and is smoking a
hookah A hookah ( Hindustani: ( Nastaleeq), (Devanagari), IPA: ; also see other names), shisha, or waterpipe is a single- or multi-stemmed instrument for heating or vaporizing and then smoking either tobacco, flavored tobacco (often '' muʽassel ...
; the image can be read as showing either the forelegs of the larva, or as suggesting a face with protruding nose and chin.
Eric Carle Eric Carle (June 25, 1929 – May 23, 2021) was an American author, designer and illustrator of children's books. His picture book '' The Very Hungry Caterpillar'', first published in 1969, has been translated into more than 66 languages and sol ...
's children's book ''
The Very Hungry Caterpillar ''The Very Hungry Caterpillar'' is a 1969 children’s picture book designed, illustrated, and written by Eric Carle. The book features a hungry caterpillar that eats a variety of foods before pupating and emerging as a butterfly. It has won m ...
'' portrays the larva as an extraordinarily hungry animal, while also teaching children how to count (to five) and the days of the week. A butterfly appeared in one of
Rudyard Kipling Joseph Rudyard Kipling ( ; 30 December 1865 – 18 January 1936)'' The Times'', (London) 18 January 1936, p. 12. was an English novelist, short-story writer, poet, and journalist. He was born in British India, which inspired much of his work. ...
's ''
Just So Stories ''Just So Stories for Little Children'' is a 1902 collection of origin stories by the British author Rudyard Kipling. Considered a classic of children's literature, the book is among Kipling's best known works. Kipling began working on the ...
'', "
The Butterfly that Stamped "The Butterfly that Stamped" is part of a series of books known as ''Just So Stories'' by Rudyard Kipling. These are collected short stories. The stories explain how things supposedly came to be. Plot “The Butterfly that Stamped” is one of t ...
". One of the most popular, and most often recorded, songs by
Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic countries, Nordic c ...
's eighteenth-century bard,
Carl Michael Bellman Carl Michael Bellman (; 4 February 1740 – 11 February 1795) was a Swedish songwriter, composer, musician, poet and entertainer. He is a central figure in the Swedish song tradition and remains a powerful influence in Swedish music, as well ...
, is "
Fjäriln vingad syns på Haga Fjäril'n vingad syns på Haga (The butterfly wingèd's seen in Hagaparken, Haga) is one of Carl Michael Bellman's collection of songs called ''Fredmans sånger'', published in 1791, where it is No. 64. The song describes Hagaparken, Haga Park, t ...
" (The butterfly wingèd is seen in Haga), one of his '' Fredman's Songs''. ''
Madam Butterfly ''Madama Butterfly'' (; ''Madame Butterfly'') is an opera in three acts (originally two) by Giacomo Puccini, with an Italian libretto by Luigi Illica and Giuseppe Giacosa. It is based on the short story "Madame Butterfly" (1898) by John Luther ...
'' is a 1904
opera Opera is a form of theatre in which music is a fundamental component and dramatic roles are taken by singers. Such a "work" (the literal translation of the Italian word "opera") is typically a collaboration between a composer and a librett ...
by
Giacomo Puccini Giacomo Puccini (Lucca, 22 December 1858Bruxelles, 29 November 1924) was an Italian composer known primarily for his operas. Regarded as the greatest and most successful proponent of Italian opera after Verdi, he was descended from a long li ...
about a romantic young Japanese bride who is deserted by her American officer husband soon after they are married. It was based on
John Luther Long John Luther Long (January 1, 1861 – October 31, 1927) was an American lawyer and writer best known for his short story "Madame Butterfly", which was based on the recollections of his sister, Jennie Correll, who had been to Japan with her husba ...
's short story written in 1898.


In mythology and folklore

According to
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 of a person's soul; whether they be living, dying, or already dead. One Japanese superstition says that if a butterfly enters your guest room and perches behind the bamboo screen, the person whom you most love is coming to see you. 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. Diderot's ''
Encyclopédie ''Encyclopédie, ou dictionnaire raisonné des sciences, des arts et des métiers'' (English: ''Encyclopedia, or a Systematic Dictionary of the Sciences, Arts, and Crafts''), better known as ''Encyclopédie'', was a general encyclopedia publis ...
'' cites butterflies as a symbol for the soul. A Roman sculpture depicts a butterfly exiting the mouth of a dead man, representing the Roman belief that the soul leaves through the mouth. In line with this, the ancient Greek word for "butterfly" is ψυχή (''psȳchē''), which primarily means "soul" or "mind". According to
Mircea Eliade Mircea Eliade (; – April 22, 1986) was a Romanians, Romanian History of religion, historian of religion, fiction writer, philosopher, and professor at the University of Chicago. He was a leading interpreter of religious experience, who establ ...
, some of the Nagas of
Manipur Manipur () ( mni, Kangleipak) is a state in Northeast India, with the city of Imphal as its capital. It is bounded by the Indian states of Nagaland to the north, Mizoram to the south and Assam to the west. It also borders two regions of Myanm ...
claim ancestry from a butterfly.Rabuzzi, M. 1997. Butterfly etymology. Cultural Entomology November 1997 Fourth issu
online
In some cultures, butterflies symbolise
rebirth Rebirth may refer to: Arts, entertainment, and media Film * ''Rebirth'' (2011 film), a 2011 Japanese drama film * ''Rebirth'' (2016 film), a 2016 American thriller film * ''Rebirth'', a documentary film produced by Project Rebirth * ''The Re ...
. The butterfly is a symbol of being
transgender A transgender (often abbreviated as trans) person is someone whose gender identity or gender expression does not correspond with their sex assigned at birth. Many transgender people experience dysphoria, which they seek to alleviate through tr ...
, because of the transformation from caterpillar to winged adult. In the English county of
Devon Devon ( , historically known as Devonshire , ) is a ceremonial and non-metropolitan county in South West England. The most populous settlement in Devon is the city of Plymouth, followed by Devon's county town, the city of Exeter. Devon is ...
, people once hurried to kill the first butterfly of the year, to avoid a year of bad luck. In the Philippines, a lingering black or dark butterfly or moth in the house is taken to mean an impending or recent death in the family. Several American states have chosen an official state butterfly.


Collecting, recording, and rearing

"Collecting" means preserving dead specimens, not keeping butterflies as pets. Collecting butterflies was once a popular hobby; it has now largely been replaced by photography, recording, and rearing butterflies for release into the wild. The zoological illustrator
Frederick William Frohawk Frederick William Frohawk (16 July 1861 – 10 December 1946) was an England, English zoological artist and lepidopterist. Frohawk was the author of ''Natural History of British Butterflies'' (1914), ''The Complete Book of British Butterflies' ...
succeeded in rearing all the butterfly species found in Britain, at a rate of four per year, to enable him to draw every stage of each species. He published the results in the folio sized handbook ''The Natural History of British Butterflies'' in 1924. Butterflies 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 can be reared for recreation or for release.


In technology

Study of the
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 wi ...
of the wing scales of swallowtail butterflies has led to the development of more efficient
light-emitting diode A light-emitting diode (LED) is a semiconductor device that emits light when current flows through it. Electrons in the semiconductor recombine with electron holes, releasing energy in the form of photons. The color of the light (cor ...
s, and is inspiring
nanotechnology Nanotechnology, also shortened to nanotech, is the use of matter on an atomic, molecular, and supramolecular scale for industrial purposes. The earliest, widespread description of nanotechnology referred to the particular technological goal o ...
research to produce paints that do not use toxic pigments and the development of new display technologies.


References


External links


Papilionoidea on the Tree of Life

Butterfly species and observations on iNaturalist
*
Rhopalocera
at insectoid.info


Regional lists


North America

AfricaGhana
* Asia
Indo-ChinaSulawesiTurkey
{{Authority control Articles containing video clips Extant Lutetian first appearances Insects in culture