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Butterflies are
insect Insects (from Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication Communication (from Latin ''communicare'', meaning "to share" or "to be in ...

insect
s in the
macrolepidoptera Macrolepidoptera is a group within the insect Insects or Insecta (from Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the are ...
n
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyly, monophyletic – that is, composed of a common ancestor and all its lineage (evolution), lineal descendants - on a phylogenetic tree. R ...

clade
Rhopalocera from the
order Order, ORDER or Orders may refer to: * Orderliness Orderliness is a quality that is characterized by a person’s interest in keeping their surroundings and themselves well organized, and is associated with other qualities such as cleanliness a ...
Lepidoptera Lepidoptera ( ; ) is an order Order or ORDER or Orders may refer to: * Orderliness Orderliness is associated with other qualities such as cleanliness Cleanliness is both the abstract state of being clean and free from germs, dirt, trash, or ...

Lepidoptera
, which also includes
moth Moths are a paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and all descendants of that ancestor excluding a few—typically only one or two—Monophyly, monophyletic subgroups. The g ...

moth
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 ...

Papilionoidea
, 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 In chronology 222px, Joseph Scaliger's ''De emendatione temporum'' (1583) began the modern science of chronology Chronology (from Latin Latin (, or , ) is a classical language bel ...
, about 56 million years ago. Butterflies have the typical four-stage insect life cycle. Winged adults lay eggs on the food plant on which their
larva A larva (plural larvae ) is a distinct juvenile form many animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that f ...
e, known as
caterpillar Caterpillars ( ) are the larval stage A larva (plural larvae ) is a distinct juvenile form many animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Anim ...

caterpillar
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 insect Insects (from Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communicat ...

pupa
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 Holometabolism, holometabolous: they go through four distinc ...

chrysalis
. When
metamorphosis Metamorphosis is a biological process Biological processes are those processes that are vital for an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are ...
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 The leopard (''Pan ...

camouflage
,
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 ...

mimicry
and
aposematism Aposematism is the advertising by an animal to potential predation , 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, f ...
to evade their predators. Some, like the
monarch A monarch is a head of state A head of state (or chief of state) is the public persona A persona (plural personae or personas), depending on the context, can refer to either the public image of one's personality, or the social role tha ...

monarch
and the
painted lady ''Vanessa cardui'' is a well-known colourful butterfly, known as the painted lady, or formerly in North America as the cosmopolitan. Description File:Vanessa cardui MHNT CUT 2013 3 14 Pontfaverger-Moronvilliers Dos.jpg, Dorsal side File:V ...

painted lady
,
migrate Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum len ...
over long distances. Many butterflies are attacked by
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is adaptation (biology), adapted structurally to this w ...
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 Insects (from Latin ') are pancrustacean Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-p ...
,
protozoa Protozoa (singular protozoon or protozoan, plural protozoa or protozoans) is an informal term for a group of single-celled eukaryote Eukaryotes () are organism In biology, an organism () is any organic, life, living system that ...

protozoa
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 ...

flies
, and other invertebrates, or are
preyed upon
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 Pollen is a powdery substance consisting of pollen grains which are Sporophyte, microsporophytes of spermatophyta, seed plants, which produce male gametes (sperm cells). Pollen grains have a hard coat ...

pollination
of some plants. Larvae of a few butterflies (e.g.,
harvesters Harvester may refer to: Agriculture and forestry * Combine harvester The modern combined harvester, or simply combine, is a versatile machine designed to efficiently harvest a variety of grain crops. The name derives from its combining four ...
) eat harmful insects, and a few are predators of
ant Ants are eusocial Eusociality (from Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population i ...

ant
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 engage ...
in association with ants. Culturally, butterflies are a popular motif in the visual and literary arts.


Etymology

The ''
Oxford English Dictionary The ''Oxford English Dictionary'' (''OED'') is the principal historical dictionary A historical dictionary or dictionary on historical principles is a dictionary which deals not only with the latterday meanings of words but also the historica ...
'' derives the word straightforwardly from
Old English Old English (, ), or Anglo-Saxon, is the earliest recorded form of the English language English is a West Germanic language of the Indo-European language family The Indo-European languages are a language family A language ...
''butorflēoge'', butter-fly; similar names in
Old Dutch In linguistics, Old Dutch or Old Low Franconian is the set of Franconian dialects (i.e. dialects that evolved from Frankish Frankish may refer to: * Franks The Franks ( la, Franci or ) were a group of Germanic peoples The historical ...

Old Dutch
and
Old High German Old High German (OHG, german: Althochdeutsch, German abbr. ) is the earliest stage of the German language German ( Standard High German: , ) is a West Germanic language mainly spoken in Central Europe Central Europe is an area of Euro ...
show that the name is ancient. 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 ...

Gonepteryx rhamni
''); another is that butterflies were on the wing in meadows during the spring and summer butter season while the grass was growing.


Taxonomy and phylogeny

The earliest
Lepidoptera Lepidoptera ( ; ) is an order Order or ORDER or Orders may refer to: * Orderliness Orderliness is associated with other qualities such as cleanliness Cleanliness is both the abstract state of being clean and free from germs, dirt, trash, or ...

Lepidoptera
fossils are of a small moth, '' Archaeolepis mane'', of
Jurassic The Jurassic ( ) is a geologic period The geologic time scale (GTS) is a system of chronological dating that classifies Geology, geological strata (stratigraphy) in time. It is used by geologists, paleontology, paleontologists, and other earth ...
age, around 190 million years ago (mya). 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 grou ...

monophyletic
(forming a single
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyly, monophyletic – that is, composed of a common ancestor and all its lineage (evolution), lineal descendants - on a phylogenetic tree. R ...

clade
), the moths are not. The oldest butterflies are from the
Palaeocene The Paleocene, ( ) or Palaeocene, is a geological epoch In chronology 222px, Joseph Scaliger's ''De emendatione temporum'' (1583) began the modern science of chronology Chronology (from Latin Latin (, or , ) is a classical language bel ...
MoClay or
Fur Formation
Fur Formation
of Denmark, approximately 55 million years old. The oldest American butterfly is the
Late Eocene The Eocene ( ) Epoch is a geological epoch In chronology 222px, Joseph Scaliger's ''De emendatione temporum'' (1583) began the modern science of chronology Chronology (from Latin Latin (, or , ) is a classical language belonging to the ...
''
Prodryas persephone ''Prodryas persephone'' is an extinct species of Nymphalidae, brush-footed butterfly, known from a single specimen from the Chadronian-aged Florissant Fossil Beds National Monument, Florissant Shale Lagerstatte of Late Eocene Colorado. ''P. persep ...

Prodryas persephone
'' from the Florissant Fossil Beds, approximately 34 million years old. 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 ...

Papilionoidea
excluding the smaller groups of the
Hesperiidae Skippers are a family (biology), family of the Lepidoptera (moths and butterflies) named the Hesperiidae. Being Diurnality, diurnal, they are generally called butterflies. They were previously placed in a separate superfamily, Hesperioidea; howev ...

Hesperiidae
(skippers) and the more moth-like
Hedylidae Hedylidae, the "American moth-butterflies", is a family In , family (from la, familia) is a of people related either by (by recognized birth) or (by marriage or other relationship). The purpose of families is to maintain the well-being ...
of America.
Phylogenetic In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanism ...

Phylogenetic
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 all descendants of that ancestor excluding a few—typically only one or two—Monophyly, monophyletic subgroups. The group is said to be paraphyleti ...
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 Greek ( el, label=Modern Greek Modern Greek (, , or , ''Kiní Neoellinikí Glóssa''), generally referred to by speakers simply as Greek (, ), refers collectively to the diale ...
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. Melanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the ami ...

melanin
s that give them blacks and browns, as well as
uric acid Uric acid is a heterocyclic compound A heterocyclic compound or ring structure is a cyclic compound that has atoms of at least two different chemical element, elements as members of its ring(s). Heterocyclic chemistry is the branch of orga ...

uric acid
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, and so ...
that give them yellows, but many of the blues, greens, reds and iridescent colours are created by
structural coloration In living creatures, structural coloration is the production of colour by microscopically structured surfaces fine enough to interfere with visible light, sometimes in combination with pigments. For example, peacock tail feathers are pigmented ...
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 Anatomy (Greek ''anatomē'', 'dissection') is the branch of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Bioc ...
, and
abdomen The abdomen (colloquially called the belly, tummy, midriff 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 Trunk (anatomy) ...

abdomen
. 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 In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and all descendants of that ancestor excluding a few—typically only one or two—Monophyly, monophyletic subgroups. The g ...

moth
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 Diurnal ("daily Daily or The Daily may refer to: Journalism * Daily newspaper A newspaper is a Periodical literature, periodical publication containing written News, information about current events and is often typed in black ink with a ...
, 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 The Sphingidae are a family of Moth, moths (Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as ...

hummingbird hawk-moth
, are exceptions to these rules. Butterfly
larva A larva (plural larvae ) is a distinct juvenile form many animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that f ...
e,
caterpillar Caterpillars ( ) are the larval stage A larva (plural larvae ) is a distinct juvenile form many animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Anim ...

caterpillar
s, have a hard ( sclerotised) 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 insect Insects (from Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communicat ...

pupa
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 species exhibit different characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most animals and some plants. Differences may in ...
. Most butterflies have the
ZW sex-determination system The ZW sex-determination system is a chromosomal system that determines the sex of offspring in bird Birds are a group of warm-blooded vertebrates constituting the class Class or The Class may refer to: Common uses not otherwise c ...
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. Image:Ecozone Nearctic.svg, 400px, The Nearctic realm The Nearctic realm covers most of North America, including Greenland, Central Florida, and t ...
; 7,700
Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropics, tropical Ecoregion#Terrestrial, terrestrial ecoregions of the Americas and the entire South American temperat ...
; 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-Siberi ...
; 3,650
Afrotropical The Afrotropical realm is one of Earth's eight biogeographic realms. It includes Africa south of the Sahara, Sahara Desert, the majority of the Arabian Peninsula, the island of Madagascar, southern Iran and extreme southwestern Pakistan, and the ...
; and 4,800 are distributed across the combined
Oriental The Orient is a term for the East East is one of the four cardinal direction The four cardinal directions, or cardinal points, are the directions north North is one of the four compass points or cardinal directions. It is the opposite ...
and
Australian Australians, colloquially referred to as "Aussies", are the citizens Citizenship is a relationship between an individual and a state to which the individual owes allegiance and in turn is entitled to its protection. Each state determines ...
/
Oceania Oceania (, , ) is a geographic region In geography Geography (from Greek: , ''geographia'', literally "earth description") is a field of science devoted to the study of the lands, features, inhabitants, and phenomena of the Eart ...

Oceania
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. It ...
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 Aragonese or Aragones may refer to: * Something related to Aragon, an autonomous community and former kingdom in Spain * the Aragonese people, those originating from or living in the historical region ...

Iberian Peninsula
. 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 a well-known colourful butterfly, known as the painted lady, or formerly in North America as the cosmopolitan. Description File:Vanessa cardui MHNT CUT 2013 3 14 Pontfaverger-Moronvilliers Dos.jpg, Dorsal side File:V ...

painted lady
, monarch, and several 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 In meteorology Meteorology is a branch of the (which include and ), with a major focus on . The study of meteorology ...

monsoon
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'') 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 Geography (from Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its populati ...
to
tropical regions The tropics are the region of Earth Earth is the third planet from the Sun and the only astronomical object known to harbor life. About 29% of Earth's surface is land consisting of continent A continent is one of several larg ...

tropical regions
with tropical regions showing a trend towards multivoltinism.
Courtship Courtship is the period of development towards a sexual relationship An intimate relationship is an interpersonal relationship that involves Physical intimacy, physical or emotional intimacy. Although an intimate relationship is commonly a ...

Courtship
is often aerial and often involves
pheromone A pheromone (from Ancient Greek ' "to bear" and hormone) 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 s ...

pheromone
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 A spermatozoon (pronounced , alternate spelling spermatozoön; plural spermatozoa; from grc, σπέρμα ("seed") and grc, ζῷον ("living being")) is a motile spe ...
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 ...
'') to prevent her from mating again. The vast majority of butterflies have a four-stage life cycle;
egg An egg is the organic vessel containing the in which an develops until it can survive on its own, at which point the animal hatches. An egg results from of an . Most s, (excluding s), and lay eggs, although some, such as s, do not. eg ...

egg
,
larva A larva (plural larvae ) is a distinct juvenile form many animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that f ...
(caterpillar),
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insect Insects (from Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communicat ...

pupa
(chrysalis) and
imago In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanisms, ...

imago
(adult). In the genera ''
Colias ''Colias'' is a genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), circumsc ...

Colias
'', ''
Erebia ''Erebia'' is a Holarctic genus of brush-footed butterflies, family (biology), family Nymphalidae. Most of the about 90–100 species (see also #Taxonomy and systematics, below) are dark brown or black in color, with reddish-brown to orange or ...
'', ''
Euchloe ''Euchloe'' is a genus of Pieridae, pierid butterfly, butterflies from the Anthocharini, orangetip tribe (biology), tribe (Anthocharini). They are Holarctic in distribution, with most species in Europe, Central Asia, and North America. Like other ...
'', 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 Butterflies are insect Insects or Insecta (from Latin Latin (, ...

Pieris brassicae
'', 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 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 regularly and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press ...
(resting) stage, and the hatching may take place only in spring. Some temperate region butterflies, such as the , 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 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 List of feeding behaviours, feeding behaviours that includes parasitism and micropredation (which ...

predators
: '''' eats
scale insect Scale insects are small insect Insects (from Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as L ...

scale insect
s, while lycaenids such as ''
Liphyra brassolis ''Liphyra brassolis'', the moth butterfly, is a butterfly found in South Asia, Southeast Asia and Australia that belongs to the Lycaenidae, lycaenid family. The larvae are predatory and feed on ant larvae. This is one of the largest species of lyca ...

Liphyra brassolis
'' are
myrmecophilous Myrmecophily ( ; literally "ant-love") is the term applied to positive interspecies Association (ecology), associations between ants and a variety of other organisms, such as plants, other arthropods, and fungi. Myrmecophily refers to Mutualism (b ...
, eating ant larvae. Some larvae, especially those of the
Lycaenidae Lycaenidae is the second-largest family In , family (from la, familia) is a of people related either by (by recognized birth) or (by marriage or other relationship). The purpose of families is to maintain the well-being of its members ...

Lycaenidae
, form mutual associations with ants. They communicate with the ants using vibrations that are transmitted through the
substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached **Substrate (locomotion), the surface over which an organism loco ...
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 Lycaenidae is the second-largest family (biology), family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide ...
(''Phengaris arion'') caterpillars trick ''
Myrmica ''Myrmica'' is a genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a ...
'' ants into taking them back to the
ant colony An ant colony is the basic unit around which ants organize their lifecycle. Ant colonies are eusocial, communal, and division of labor, efficiently organized and are very much like those found in other social Hymenoptera, though the various grou ...

ant colony
where they feed on the ant eggs and larvae in a parasitic relationship. Caterpillars mature through a series of developmental stages known as instars. 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 tough outer layer made of a mixture of chitin and specialized proteins, is released from the softer Squamous epithelium, epidermis beneath, and the epidermis begins to form a new cuticle. At the end of each instar, the larva ecdysis, 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 eye in invertebrates, simple eyes. The Insect mouthparts, mouthparts 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 prolegs 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 setae, 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 gonads start development in the egg stage.


Pupa

When the larva is fully grown, hormones such as prothoracicotropic hormone (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 Pupa#Cocoon, 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 Biological processes are those processes that are vital for an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are ...
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 Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanisms, ...

imago
. The surface of both butterflies and moths is covered by scales, each of which is an outgrowth from a single Epidermis, epidermal cell. The head is small and dominated by the two large Eye#Compound eyes, compound eyes. 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 Antenna (biology), 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 mandibles are usually reduced in size or absent. The first maxillae are elongated into a tubular proboscis 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'' 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 Nymphalidae, 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 (Wing coupling (Lepidoptera anatomy), as they are in moths) but are coordinated by the friction of their overlapping parts. The front two segments have a pair of Spiracle (arthropods), 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 A spermatozoon (pronounced , alternate spelling spermatozoön; plural spermatozoa; from grc, σπέρμα ("seed") and grc, ζῷον ("living being")) is a motile spe ...
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 and letting them dry, during which time it is extremely vulnerable to predators.


Behaviour

Butterflies feed primarily on nectar from flowers. Some also derive nourishment from pollen, 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 bees, but they are capable of moving pollen over greater distances. Flower constancy 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 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 behaviour is restricted to the males, and studies have suggested that the nutrients collected may be provided as a nuptial gift, along with the spermatophore, during mating. In Hill-topping (biology), 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 sensillum, sensillae. A butterfly's sense of taste is coordinated by chemoreceptors on the Arthropod leg#Insects, 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 (from Ancient Greek ' "to bear" and hormone) 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 s ...

pheromone
s; some have specialized scent scales (androconia) or other structures (coremata 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 Stridulation, stridulatory 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 (force), lift generated by butterflies is more than can be accounted for by steady-state, non-transitory aerodynamics. Studies using ''Vanessa atalanta'' in a wind tunnel 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 Torkel Weis-Fogh, Weis-Fogh 'clap-and-fling' 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. Braconidae, Braconid 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 Pieris brassicae, large white butterfly. When the Pieris rapae, 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'' has been used in sprays to reduce damage to crops by the caterpillars of the large white butterfly, and the entomopathogenic fungus ''Beauveria bassiana'' has proved effective for the same purpose.


Endangered species

Queen Alexandra's birdwing is the largest butterfly in the world. The species is List of endangered insects#Other Lepidoptera species, endangered, and is one of only three insects (the other two being butterflies as well) to be listed on CITES#Appendix I, Appendix I of CITES, making international trade illegal.CITES appendices I, II and III
official website
Ocybadistes knightorum, Black grass-dart butterfly ''(Ocybadistes knightorum)'' is a butterfly of the family
Hesperiidae Skippers are a family (biology), family of the Lepidoptera (moths and butterflies) named the Hesperiidae. Being Diurnality, diurnal, they are generally called butterflies. They were previously placed in a separate superfamily, Hesperioidea; howev ...

Hesperiidae
. It is endemic to New South Wales. It has a very limited distribution in the Boambee, New South Wales, 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 plant defense against herbivory, protection 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 predation , 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, f ...
). This signal is commonly mimicry, mimicked by other butterflies, usually only females. A Batesian mimicry, Batesian mimic imitates another species to enjoy the protection of that species' aposematism. The Papilio polytes, common Mormon of India has female morphs which imitate the unpalatable red-bodied swallowtails, the Pachliopta aristolochiae, common rose and the Pachliopta hector, crimson rose. Müllerian mimicry occurs when aposematic species evolve to resemble each other, presumably to reduce predator sampling rates; ''Heliconius'' butterflies from the Americas are a good example. Camouflage is found in many butterflies. Some like the oakleaf butterfly and Doleschallia bisaltide, autumn leaf are remarkable imitations of leaves. As caterpillars, many defend themselves by freezing and appearing like sticks or branches. Others have deimatic 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'') 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 myrmecophiles, forming Symbiosis, mutualistic associations with
ant Ants are eusocial Eusociality (from Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population i ...

ant
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 wasps. 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 Eyespot (mimicry), 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 refers to vibrations generated by the butterfly upon expanding its wings in an attempt to communicate with ant predators. Many tropical butterflies have seasonal polyphenism, seasonal forms for dry and wet seasons. These are switched by the hormone ecdysone. 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 created by the small whirlpools formed by the wings during flight.


Declining numbers

Declining butterfly populations have been observed in many areas of the world, and this phenomenon is consistent with the decline in insect populations , 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 Mesoamerican city of Teotihuacan, the brilliantly coloured image of the butterfly was carved into many temples, buildings, jewellery, and emblazoned on Censer, incense burners. The butterfly was sometimes depicted with the maw of a jaguar, 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; evidence of similar jaguar-butterfly images has been found among the Zapotec peoples, 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 Norwegians, Norwegian naturalist Kjell B. Sandved, 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 drew a famous illustration of Alice (Alice's Adventures in Wonderland), Alice meeting a Caterpillar (Alice's Adventures in Wonderland), caterpillar for Lewis Carroll's ''Alice's Adventures in Wonderland, Alice in Wonderland'', c. 1865. The caterpillar is seated on a toadstool and is smoking a hookah; 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's children's book ''The Very Hungry Caterpillar'' portrays the larva as an extraordinarily hungry animal, while also teaching children how to count (to five) and the days of the week. One of the most popular, and most often recorded, songs by Sweden's eighteenth-century bard, Carl Michael Bellman, is "Fjäriln vingad syns på Haga" (The butterfly wingèd is seen in Haga), one of his ''Fredman's Songs''. ''Madam Butterfly'' is a 1904 opera by Giacomo Puccini 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's short story written in 1898.


In mythology and folklore

According to Lafcadio Hearn, 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 omens. When Taira no Masakado was secretly preparing for his famous revolt, there appeared in Kyoto so vast a swarm of butterflies that the people were frightened—thinking the apparition to be a portent of coming evil. Diderot's ''Encyclopédie'' 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, some of the Naga people, Nagas of Manipur claim ancestry from a butterfly.Rabuzzi, M. 1997. Butterfly etymology. Cultural Entomology November 1997 Fourth issu
online
In some cultures, butterflies symbolise Reincarnation, rebirth. The butterfly is a symbol of being transgender, because of the transformation from caterpillar to winged adult. In the English county of Devon, people once hurried to kill the first butterfly of the year, to avoid a year of bad luck. In the Philippines, a lingering black butterfly or moth in the house is taken to mean a death in the family. Several American states have chosen an List of U.S. state butterflies, 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 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 In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and all descendants of that ancestor excluding a few—typically only one or two—Monophyly, monophyletic subgroups. The g ...

moth
s can be reared for recreation or for release.


In technology

Study of the
structural coloration In living creatures, structural coloration is the production of colour by microscopically structured surfaces fine enough to interfere with visible light, sometimes in combination with pigments. For example, peacock tail feathers are pigmented ...
of the wing scales of swallowtail butterflies has led to the development of more efficient light-emitting diodes, and is inspiring nanotechnology 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
*
Rhopalocera
at insectoid.info


Regional lists


North America

AfricaGhana
* Asia
Indo-ChinaSulawesiTurkey
{{Use dmy dates, date=October 2020 Butterflies, Articles containing video clips Extant Lutetian first appearances Insects in culture