Lepidoptera (/ˌlɛpɪˈdɒptərə/ lep-i-DOP-tər-ə) is an order of
insects that includes butterflies and moths (both are called
lepidopterans). About 180,000 species of the
described, in 126 families and 46 superfamilies, 10% of the
total described species of living organisms. It is one of the
most widespread and widely recognizable insect orders in the world.
Lepidoptera show many variations of the basic body structure that
have evolved to gain advantages in lifestyle and distribution. Recent
estimates suggest the order may have more species than earlier
thought, and is among the four most speciose orders, along with the
Hymenoptera, Diptera, and Coleoptera.
Lepidopteran species are characterized by more than three derived
features, some of the most apparent being the scales covering their
bodies and wings, and a proboscis. The scales are modified, flattened
"hairs", and give butterflies and moths their extraordinary variety of
colors and patterns. Almost all species have some form of membranous
wings, except for a few that have reduced wings or are wingless. Like
most other insects, butterflies and moths are holometabolous, meaning
they undergo complete metamorphosis.
Mating and the laying of eggs are
carried out by adults, normally near or on host plants for the larvae.
The larvae are commonly called caterpillars, and are completely
different from their adult moth or butterfly forms, having a
cylindrical body with a well-developed head, mandible mouth parts,
three pairs of thoracic legs and from none up to five pairs of
prolegs. As they grow, these larvae change in appearance, going
through a series of stages called instars. Once fully matured, the
larva develops into a pupa. A few butterflies and many moth species
spin a silk case or cocoon prior to pupating, while others do not,
instead going underground. A butterfly pupa, called a chrysalis,
has a hard skin, usually with no cocoon. Once the pupa has completed
its metamorphosis, a sexually mature adult emerges.
Lepidoptera have, over millions of years, evolved a wide range of
wing patterns and coloration ranging from drab moths akin to the
related order Trichoptera, to the brightly-colored and
complex-patterned butterflies. Accordingly, this is the most
recognized and popular of insect orders with many people involved in
the observation, study, collection, rearing of, and commerce in these
insects. A person who collects or studies this order is referred to as
Butterflies and moths play an important role in the natural ecosystem
as pollinators and as food in the food chain; conversely, their larvae
are considered very problematic to vegetation in agriculture, as their
main source of food is often live plant matter. In many species, the
female may produce from 200 to 600 eggs, while in others, the number
may approach 30,000 eggs in one day. The caterpillars hatching from
these eggs can cause damage to large quantities of crops. Many moth
and butterfly species are of economic interest by virtue of their role
as pollinators, the silk they produce, or as pest species.
2 Distribution and diversity
3 External morphology
4 Internal morphology
4.1 Reproductive system
4.2 Digestive system
4.3 Circulatory system
4.4 Respiratory system
6 Reproduction and development
6.1 Lepidopterans in diapause
6.3 Life cycle
8.1 Defense and predation
8.5 Other biological interactions
9 Evolution and systematics
9.1 History of study
9.2 Fossil record
10 Relationship to people
10.3 Beneficial insects
11 See also
13 Further reading
14 External links
The term was coined by Linnaeus in 1735 and is derived from Greek
λεπίς, gen. λεπίδος ("scale") and πτερόν
("wing"). Sometimes, the term
Rhopalocera is used for the clade of
all butterfly species, derived from the
Ancient Greek ῥόπαλον
(rhopalon):4150 and κέρας (keras):3993 meaning "club" and
"horn", respectively, coming from the shape of the antennae of
The origins of the common names "butterfly" and "moth" are varied and
often obscure. The English word butterfly is from Old English
buttorfleoge, with many variations in spelling. Other than that, the
origin is unknown, although it could be derived from the pale yellow
color of many species' wings suggesting the color of butter. The
Heterocera are commonly called moths. The origins of the
English word moth are more clear, deriving from the Old English
Northumbrian dialect mohðe) from Common Germanic
Old Norse motti, Dutch mot and German Motte all meaning
"moth"). Perhaps its origins are related to
Old English maða meaning
"maggot" or from the root of "midge", which until the 16th century was
used mostly to indicate the larva, usually in reference to devouring
The etymological origins of the word "caterpillar", the larval form of
butterflies and moths, are from the early 16th century, from Middle
English catirpel, catirpeller, probably an alteration of Old North
French catepelose: cate, cat (from
Latin cattus) + pelose, hairy (from
Distribution and diversity
Main article: Lists of
Lepidoptera by region
Lepidoptera are among the most successful groups of insects. They
are found on all continents, except Antarctica, and inhabit all
terrestrial habitats ranging from desert to rainforest, from lowland
grasslands to mountain plateaus, but almost always associated with
higher plants, especially angiosperms (flowering plants). Among
the most northern dwelling species of butterflies and moths is the
Arctic Apollo (Parnassius arcticus), which is found in the Arctic
Circle in northeastern Yakutia, at an altitude of 1500 m above
sea level. In the Himalayas, various Apollo species such as
Parnassius epaphus have been recorded to occur up to an altitude of
6,000 m above sea level.:221
Some lepidopteran species exhibit symbiotic, phoretic, or parasitic
lifestyles, inhabiting the bodies of organisms rather than the
environment. Coprophagous pyralid moth species, called sloth moths,
Bradipodicola hahneli and Cryptoses choloepi, are unusual in
that they are exclusively found inhabiting the fur of sloths, mammals
found in Central and South America. Two species of Tinea moths
have been recorded as feeding on horny tissue and have been bred from
the horns of cattle. The larva of
Zenodochium coccivorella is an
internal parasite of the coccid Kermes species. Many species have been
recorded as breeding in natural materials or refuse such as owl
pellets, bat caves, honeycombs or diseased fruit.
As of 2007, there was roughly 174,250 lepidopteran species described,
with butterflies and skippers estimated to comprise around 17,950, and
moths making up the rest. The vast majority of
to be found in the tropics, but substantial diversity exists on most
continents. North America has over 700 species of butterflies and over
11,000 species of moths, while about 400 species of
butterflies and 14,000 species of moths are reported from
Australia. The diversity of
Lepidoptera in each faunal region has
been estimated by John Heppner in 1991 based partly on actual counts
from the literature, partly on the card indices in the Natural History
Museum (London) and the National
Museum of Natural History
(Washington), and partly on estimates:
Lepidoptera in each faunal region
(comprising Indomalayan and Australian regions)
Estimated number of species
Main article: External morphology of Lepidoptera
Parts of an adult butterfly
A – head, B – thorax, C – abdomen, 1 – prothoracic shield, 2
– spiracle, 3 – true legs, 4 – midabdominal prolegs, 5 – anal
proleg, 6 – anal plate, 7 – tentacle, a – frontal triangle, b
– stemmata (ocelli), c – antenna, d – mandible, e – labrum.
Lepidoptera are morphologically distinguished from other orders
principally by the presence of scales on the external parts of the
body and appendages, especially the wings.
Butterflies and moths vary
in size from microlepidoptera only a few millimeters long, to
conspicuous animals with a wingspan greater than 25 centimetres, such
Queen Alexandra's birdwing
Queen Alexandra's birdwing and Atlas moth.:246
Lepidopterans undergo a four-stage life cycle: egg; larva or
caterpillar; pupa or chrysalis; and imago (plural: imagines) / adult
and show many variations of the basic body structure, which give these
animals advantages for diverse lifestyles and environments.
Face of a caterpillar with the mouthparts showing
The head is where many sensing organs and the mouth parts are found.
Like the adult, the larva also has a toughened, or sclerotized head
capsule. Here, two compound eyes, and chaetosema, raised spots or
clusters of sensory bristles unique to Lepidoptera, occur, though many
taxa have lost one or both of these spots. The antennae have a wide
variation in form among species and even between different sexes. The
antennae of butterflies are usually filiform and shaped like clubs,
those of the skippers are hooked, while those of moths have flagellar
segments variously enlarged or branched. Some moths have enlarged
antennae or ones that are tapered and hooked at the
The maxillary galeae are modified and form an elongated proboscis. The
proboscis consists of one to five segments, usually kept coiled up
under the head by small muscles when it is not being used to suck up
nectar from flowers or other liquids. Some basal moths still have
mandibles, or separate moving jaws, like their ancestors, and these
form the family Micropterigidae.:560
The larvae, called caterpillars, have a toughened head capsule.
Caterpillars lack the proboscis and have separate chewing
mouthparts. These mouthparts, called mandibles, are used to chew
up the plant matter that the larvae eat. The lower jaw, or labium, is
weak, but may carry a spinneret, an organ used to create silk. The
head is made of large lateral lobes, each having an ellipse of up to
six simple eyes.:562–563
The thorax is made of three fused segments, the prothorax, mesothorax,
and metathorax, each with a pair of legs. The first segment contains
the first pair of legs. In some males of the butterfly family
Nymphalidae, the forelegs are greatly reduced and are not used for
walking or perching.:586 The three pairs of legs are covered with
Lepidoptera also have olfactory organs on their feet, which
aid the butterfly in "tasting" or "smelling" out its food. In the
larval form there are 3 pairs of true legs, with up to 11 pairs of
abdominal legs (usually eight) and hooklets, called apical
The two pairs of wings are found on the middle and third segments, or
mesothorax and metathorax, respectively. In the more recent genera,
the wings of the second segment are much more pronounced, although
some more primitive forms have similarly sized wings of both segments.
The wings are covered in scales arranged like shingles, which form an
extraordinary variety of colors and patterns. The mesothorax has more
powerful muscles to propel the moth or butterfly through the air, with
the wing of this segment (forewing) having a stronger vein
structure.:560 The largest superfamily, the Noctuidae, has their
wings modified to act as tympanal or hearing organs.
The caterpillar has an elongated, soft body that may have hair-like or
other projections, three pairs of true legs, with none to 11 pairs of
abdominal legs (usually eight) and hooklets, called apical
crochets. The thorax usually has a pair of legs on each segment.
The thorax is also lined with many spiracles on both the mesothorax
and metathorax, except for a few aquatic species, which instead have a
form of gills.:563
Caterpillar prolegs on Papilio machaon
The abdomen, which is less sclerotized than the thorax, consists of 10
segments with membranes in between, allowing for articulated movement.
The sternum, on the first segment, is small in some families and is
completely absent in others. The last two or three segments form the
external parts of the species' sex organs. The genitalia of
Lepidoptera are highly varied and are often the only means of
differentiating between species. Male genitals include a valva, which
is usually large, as it is used to grasp the female during mating.
Female genitalia include three distinct sections.
The females of basal moths have only one sex organ, which is used for
copulation and as an ovipositor, or egg-laying organ. About 98% of
moth species have a separate organ for mating, and an external duct
that carries the sperm from the male.:561
The abdomen of the caterpillar has four pairs of prolegs, normally
located on the third to sixth segments of the abdomen, and a separate
pair of prolegs by the anus, which have a pair of tiny hooks called
crotchets. These aid in gripping and walking, especially in species
that lack many prolegs (e. g. larvae of Geometridae). In some basal
moths, these prolegs may be on every segment of the body, while
prolegs may be completely absent in other groups, which are more
adapted to boring and living in sand (e. g.,
Wing scales form the color and pattern on wings. The scales shown here
are lamellar. The pedicel can be seen attached to a few loose scales.
The wings, head, and parts of the thorax and abdomen of Lepidoptera
are covered with minute scales, a feature from which the order derives
its name. Most scales are lamellar, or blade-like and attached with a
pedicel, while other forms may be hair-like or specialized as
secondary sexual characteristics.
The lumen or surface of the lamella has a complex structure. It gives
color either by colored pigments it contains, or through structural
coloration with mechanisms that include photonic crystals and
Scales function in insulation, thermoregulation, producing pheromones
(in males only), and aiding gliding flight, but the most important
is the large diversity of vivid or indistinct patterns they provide,
which help the organism protect itself by camouflage or mimicry, and
which act as signals to other animals including rivals and potential
Electron microscopy images of scales
A patch of wing (×50)
Scales close up (×200)
A single scale (×1000)
Microstructure of a scale (×5000)
In the reproductive system of butterflies and moths, the male
genitalia are complex and unclear. In females the three types of
genitalia are based on the relating taxa: 'monotrysian', 'exoporian',
and 'ditrysian'. In the monotrysian type is an opening on the fused
segments of the sterna 9 and 10, which act as insemination and
oviposition. In the exoporian type (in Hepaloidae and Mnesarchaeoidea)
are two separate places for insemination and oviposition, both
occurring on the same sterna as the monotrysian type, i.e. 9 and
10. The ditrysian groups have an internal duct that carries sperm,
with separate openings for copulation and egg-laying. In most
species, the genitalia are flanked by two soft lobes, although they
may be specialized and sclerotized in some species for ovipositing in
area such as crevices and inside plant tissue. Hormones and the
glands that produce them run the development of butterflies and moths
as they go through their life cycles, called the endocrine system. The
first insect hormone prothoracicotropic hormone (PTTH) operates the
species life cycle and diapause. This hormone is produced by
corpora allata and corpora cardiaca, where it is also stored. Some
glands are specialized to perform certain task such as producing silk
or producing saliva in the palpi.:65, 75 While the corpora
cardiaca produce PTTH, the corpora allata also produces juvenile
hormones, and the prothorocic glands produce moulting hormones.
In the digestive system, the anterior region of the foregut has been
modified to form a pharyngeal sucking pump as they need it for the
food they eat, which are for the most part liquids. An esophagus
follows and leads to the posterior of the pharynx and in some species
forms a form of crop. The midgut is short and straight, with the
hindgut being longer and coiled. Ancestors of lepidopteran
species, stemming from Hymenoptera, had midgut ceca, although this is
lost in current butterflies and moths. Instead, all the digestive
enzymes, other than initial digestion, are immobilized at the surface
of the midgut cells. In larvae, long-necked and stalked goblet cells
are found in the anterior and posterior midgut regions, respectively.
In insects, the goblet cells excrete positive potassium ions, which
are absorbed from leaves ingested by the larvae. Most butterflies and
moths display the usual digestive cycle, but species with different
diets require adaptations to meet these new demands.:279
Internal morphology of adult male in the family Nymphalidae, showing
most of the major organ systems, with characteristic reduced forelegs
of that family: The corpora include the corpus allatum and the corpus
In the circulatory system, hemolymph, or insect blood, is used to
circulate heat in a form of thermoregulation, where muscles
contraction produces heat, which is transferred to the rest of the
body when conditions are unfavorable. In lepidopteran species,
hemolymph is circulated through the veins in the wings by some form of
pulsating organ, either by the heart or by the intake of air into the
Air is taken in through spiracles along the sides of the abdomen and
thorax supplying the trachea with oxygen as it goes through the
lepidopteran's respiratory system. Three different tracheaes supply
and diffuse oxygen throughout the species' bodies. The dorsal tracheae
supply oxygen to the dorsal musculature and vessels, while the ventral
tracheae supply the ventral musculature and nerve cord, and the
visceral tracheae supply the guts, fat bodies, and gonads.:71, 72
Main article: Polymorphism in Lepidoptera
Sexually dimorphic bagworm moths (Thyridopteryx ephemeraeformis)
mating: The female is flightless.
Heliconius butterflies from the tropics of the Western Hemisphere
are the classical model for Müllerian mimicry.
Polymorphism is the appearance of forms or "morphs", which differ in
color and number of attributes within a single species.:163 In
Lepidoptera, polymorphism can be seen not only between individuals in
a population, but also between the sexes as sexual dimorphism, between
geographically separated populations in geographical polymorphism, and
between generations flying at different seasons of the year (seasonal
polymorphism or polyphenism). In some species, the polymorphism is
limited to one sex, typically the female. This often includes the
phenomenon of mimicry when mimetic morphs fly alongside nonmimetic
morphs in a population of a particular species. Polymorphism occurs
both at specific level with heritable variation in the overall
morphological design of individuals, as well as in certain specific
morphological or physiological traits within a species.
Environmental polymorphism, in which traits are not inherited, is
often termed as polyphenism, which in
Lepidoptera is commonly seen in
the form of seasonal morphs, especially in the butterfly families of
Nymphalidae and Pieridae. An Old World pierid butterfly, the common
grass yellow (Eurema hecabe) has a darker summer adult morph,
triggered by a long day exceeding 13 hours in duration, while the
shorter diurnal period of 12 hours or less induces a paler morph in
the postmonsoon period.
Polyphenism also occurs in caterpillars,
an example being the peppered moth, Biston betularia.
Geographical isolation causes a divergence of a species into different
morphs. A good example is the Indian white admiral Limenitis procris,
which has five forms, each geographically separated from the other by
large mountain ranges.:26 An even more dramatic showcase of
geographical polymorphism is the Apollo butterfly (Parnassius apollo).
Because the Apollos live in small local populations, thus having no
contact with each other, coupled with their strong stenotopic nature
and weak migration ability, interbreeding between populations of one
species practically does not occur; by this, they form over 600
different morphs, with the size of spots on the wings of which varies
Seasonal diphenism in the common grass yellow, Eurema hecabe
Sexual dimorphism is the occurrence of differences between males and
females in a species. In Lepidoptera, it is widespread and almost
completely set by genetic determination.
Sexual dimorphism is
present in all families of the
Papilionoidea and more prominent in the
Lycaenidae, Pieridae, and certain taxa of the Nymphalidae. Apart from
color variation, which may differ from slight to completely different
color-pattern combinations, secondary sexual characteristics may also
be present.:25 Different genotypes maintained by natural selection
may also be expressed at the same time. Polymorphic and/or mimetic
females occur in the case of some taxa in the
to obtain a level of protection not available to the male of their
species. The most distinct case of sexual dimorphism is that of adult
females of many
Psychidae species which have only vestigial wings,
legs, and mouthparts as compared to the adult males that are strong
fliers with well-developed wings and feathery antennae.
Reproduction and development
Mating pair of
Laothoe populi (poplar hawk-moth) showing two different
Lepidoptera undergo holometabolism or "complete
metamorphosis". Their life cycle normally consists of an egg, a larva,
a pupa, and an imago or adult. The larvae are commonly called
caterpillars, and the pupae of moths encapsulated in silk are called
cocoons, while the uncovered pupae of butterflies are called
Lepidopterans in diapause
Part of a series on
Critical thermal maximum
Unless the species reproduces year-round, a butterfly or moth may
enter diapause, a state of dormancy that allows the insect to survive
unfavorable environmental conditions.
Males usually start eclosion (emergence) earlier than females and peak
in numbers before females. Both of the sexes are sexually mature by
the time of eclosion.:564
Butterflies and moths normally do not
associate with each other, except for migrating species, staying
Mating begins with an adult (female or male)
attracting a mate, normally using visual stimuli, especially in
diurnal species like most butterflies. However, the females of most
nocturnal species, including almost all moth species, use pheromones
to attract males, sometimes from long distances. Some species
engage in a form of acoustic courtship, or attract mates using sound
or vibration such as the polka-dot wasp moth, Syntomeida epilais.
Adaptations include undergoing one seasonal generation, two or even
more, called voltinism (Univoltism, bivoltism, and multivism,
respectively). Most lepidopterans in temperate climates are
univoltine, while in tropical climates most have two seasonal broods.
Some others may take advantage of any opportunity they can get, and
mate continuously throughout the year. These seasonal adaptations are
controlled by hormones, and these delays in reproduction are called
diapause.:567 Many lepidopteran species, after mating and laying
their eggs, die shortly afterwards, having only lived for a few days
after eclosion. Others may still be active for several weeks and then
overwinter and become sexually active again when the weather becomes
more favorable, or diapause. The sperm of the male that mated most
recently with the female is most likely to have fertilized the eggs,
but the sperm from a prior mating may still prevail.:564
The four stages of the life cycle of an anise swallowtail
Lepidoptera usually reproduce sexually and are oviparous (egg-laying),
though some species exhibit live birth in a process called
ovoviviparity. A variety of differences in egg-laying and the number
of eggs laid occur. Some species simply drop their eggs in flight
(these species normally have polyphagous larvae, meaning they eat a
variety of plants e. g., hepialids and some nymphalids) while most
lay their eggs near or on the host plant on which the larvae feed. The
number of eggs laid may vary from only a few to several thousand.
The females of both butterflies and moths select the host plant
instinctively, and primarily, by chemical cues.:564
The eggs are derived from materials ingested as a larva and in some
species, from the spermatophores received from males during
mating. An egg can only be 1/1000 the mass of the female, yet she
may lay up to her own mass in eggs. Females lay smaller eggs as they
age. Larger females lay larger eggs. The egg is covered by a
hard-ridged protective outer layer of shell, called the chorion. It is
lined with a thin coating of wax, which prevents the egg from drying
out. Each egg contains a number of micropyles, or tiny funnel-shaped
openings at one end, the purpose of which is to allow sperm to enter
and fertilize the egg.
Butterfly and moth eggs vary greatly in size
between species, but they are all either spherical or ovate.
The egg stage lasts a few weeks in most butterflies, but eggs laid
prior to winter, especially in temperate regions, go through diapause,
and hatching may be delayed until spring. Other butterflies may lay
their eggs in the spring and have them hatch in the summer. These
butterflies are usually temperate species (e. g. Nymphalis antiopa).
Main article: Caterpillar
Larval form typically lives and feeds on plants
The larvae or caterpillars are the first stage in the life cycle after
hatching. Caterpillars, are "characteristic polypod larvae with
cylindrical bodies, short thoracic legs, and abdominal prolegs
(pseudopods)". They have a toughened (sclerotised) head capsule
with an adfrontal suture formed by medial fusion of the sclerites,
mandibles (mouthparts) for chewing, and a soft tubular, segmented
body, that may have hair-like or other projections, three pairs of
true legs, and additional prolegs (up to five pairs). The body
consists of thirteen segments, of which three are thoracic and ten are
abdominal. Most larvae are herbivores, but a few are carnivores
(some eat ants or other caterpillars) and detritivores.
Different herbivorous species have adapted to feed on every part of
the plant and are normally considered pests to their host plants; some
species have been found to lay their eggs on the fruit and other
species lay their eggs on clothing or fur (e. g., Tineola
bisselliella, the common clothes moth). Some species are carnivorous
and others are even parasitic. Some lycaenid species such as Maculinea
rebeli are social parasites of
Myrmica ants nests. A species of
Hawaii has carnivorous larvae that catch and eat
flies. Some pyralid caterpillars are aquatic.
The larvae develop rapidly with several generations in a year;
however, some species may take up to 3 years to develop, and
exceptional examples like
Gynaephora groenlandica take as long as
seven years. The larval stage is where the feeding and growing
stages occur, and the larvae periodically undergo hormone-induced
ecdysis, developing further with each instar, until they undergo the
final larval-pupal molt.
The larvae of both butterflies and moths exhibit mimicry to deter
potential predators. Some caterpillars have the ability to inflate
parts of their heads to appear snake-like. Many have false eye-spots
to enhance this effect. Some caterpillars have special structures
called osmeteria (family Papilionidae), which are exposed to produce
smelly chemicals used in defense. Host plants often have toxic
substances in them and caterpillars are able to sequester these
substances and retain them into the adult stage. This helps make them
unpalatable to birds and other predators. Such unpalatability is
advertised using bright red, orange, black, or white warning colors.
The toxic chemicals in plants are often evolved specifically to
prevent them from being eaten by insects. Insects, in turn, develop
countermeasures or make use of these toxins for their own survival.
This "arms race" has led to the coevolution of insects and their host
No form of wing is externally visible on the larva, but when larvae
are dissected, developing wings can be seen as disks, which can be
found on the second and third thoracic segments, in place of the
spiracles that are apparent on abdominal segments.
Wing disks develop
in association with a trachea that runs along the base of the wing,
and are surrounded by a thin peripodial membrane, which is linked to
the outer epidermis of the larva by a tiny duct.
Wing disks are very
small until the last larval instar, when they increase dramatically in
size, are invaded by branching tracheae from the wing base that
precede the formation of the wing veins, and begin to develop patterns
associated with several landmarks of the wing.
Near pupation, the wings are forced outside the epidermis under
pressure from the hemolymph, and although they are initially quite
flexible and fragile, by the time the pupa breaks free of the larval
cuticle, they have adhered tightly to the outer cuticle of the pupa
(in obtect pupae). Within hours, the wings form a cuticle so hard and
well-joined to the body that pupae can be picked up and handled
without damage to the wings.
Eclosion of Papilio dardanus
After about five to seven instars,:26–28 or molts, certain
hormones, like PTTH, stimulate the production of ecdysone, which
initiates insect molting. Then, the larva puparium, a sclerotized or
hardened cuticle of the last larval instar, develops into the pupa.
Depending on the species, the pupa may be covered in a silk cocoon,
attached to different types of substrates, buried in the ground, or
may not be covered at all. Features of the imago are externally
recognizable in the pupa. All the appendages on the adult head and
thorax are found cased inside the cuticle (antennae, mouthparts,
etc.), with the wings wrapped around, adjacent to the
antennae.:564 The pupae of some species have functional mandibles,
while the pupal mandibles are not functional in others.
While encased, some of the lower segments are not fused, and are able
to move using small muscles found in between the membrane. Moving may
help the pupa, for example, escape the sun, which would otherwise kill
it. The pupa of the
Mexican jumping bean
Mexican jumping bean moth (Cydia deshaisiana) does
this. The larvae cut a trapdoor in the bean (species of Sebastiania)
and use the bean as a shelter. With a sudden rise in temperature, the
pupa inside twitches and jerks, pulling on the threads inside.
Wiggling may also help to deter parasitoid wasps from laying eggs on
the pupa. Other species of moths are able to make clicks to deter
The length of time before the pupa ecloses (emerges) varies greatly.
The monarch butterfly may stay in its chrysalis for two weeks, while
other species may need to stay for more than 10 months in diapause.
The adult emerges from the pupa either by using abdominal hooks or
from projections located on the head. The mandibles found in the most
primitive moth families are used to escape from their cocoon (e. g.,
Most lepidopteran species do not live long after eclosion, only
needing a few days to find a mate and then lay their eggs. Others may
remain active for a longer period (from one to several weeks), or go
through diapause and overwintering as monarch butterflies do, or
waiting out environmental stress. Some adult species of
microlepidoptera go through a stage where no reproductive-related
activity occurs, lasting through summer and winter, followed by mating
and oviposition in the early spring.:564
While most butterflies and moths are terrestrial, many species of
Pyralidae are truly aquatic with all stages except the adult occurring
in water. Many species from other families such as Arctiidae,
Nepticulidae, Cosmopterygidae, Tortricidae, Olethreutidae, Noctuidae,
Sphingidae are aquatic or semiaquatic.:22
Flight is an important aspect of the lives of butterflies and moths,
and is used for evading predators, searching for food, and finding
mates in a timely manner, as lepidopteran species do not live long
after eclosion. It is the main form of locomotion in most species. In
Lepidoptera, the forewings and hindwings are mechanically coupled and
flap in synchrony. Flight is anteromotoric, or being driven primarily
by action of the forewings. Although lepidopteran species reportedly
can still fly when their hindwings are cut off, it reduces their
linear flight and turning capabilities.
Lepidopteran species have to be warm, about 77 to 79 °F (25 to
26 °C), to fly. They depend on their body temperature being
sufficiently high and since they cannot regulate it themselves, this
is dependent on their environment.
Butterflies living in cooler
climates may use their wings to warm their bodies. They will bask in
the sun, spreading out their wings so that they get maximum exposure
to the sunlight. In hotter climates butterflies can easily overheat,
so they are usually active only during the cooler parts of the day,
early morning, late afternoon or early evening. During the heat of the
day, they rest in the shade. Some larger thick-bodied moths (e.g.
Sphingidae) can generate their own heat to a limited degree by
vibrating their wings. The heat generated by the flight muscles warms
the thorax while the temperature of the abdomen is unimportant for
flight. To avoid overheating, some moths rely on hairy scales,
internal air sacs, and other structures to separate the thorax and
abdomen and keep the abdomen cooler.
Some species of butterflies can reach fast speeds, such as the
southern dart, which can go as fast as 48.4 km/h. Sphingids are
some of the fastest flying insects, some are capable of flying at over
50 km/h (30 mi/h), having a wingspan of
35–150 mm. In some species, sometimes a gliding
component to their flight exists. Flight occurs either as hovering, or
as forward or backward motion. In butterfly and moth species, such
as hawk moths, hovering is important as they need to maintain a
certain stability over flowers when feeding on the nectar.
Timelapse of flying moths, attracted to the floodlights
Navigation is important to
Lepidoptera species, especially for those
that migrate. Butterflies, which have more species that migrate, have
been shown to navigate using time-compensated sun compasses. They can
see polarized light, so can orient even in cloudy conditions. The
polarized light in the region close to the ultraviolet spectrum is
suggested to be particularly important. Most migratory butterflies
are those that live in semiarid areas where breeding seasons are
short. The life histories of their host plants also influence the
strategies of the butterflies. Other theories include the use of
Lepidoptera may use coastal lines, mountains, and even
roads to orient themselves. Above sea, the flight direction is much
more accurate if the coast is still visible.
Many studies have also shown that moths navigate. One study showed
that many moths may use the
Earth's magnetic field
Earth's magnetic field to navigate, as a
study of the moth heart and dart suggests. Another study, of the
migratory behavior of the silver Y, showed, even at high altitudes,
the species can correct its course with changing winds, and prefers
flying with favourable winds, suggesting a great sense of
Aphrissa statira in
Panama loses its navigational
capacity when exposed to a magnetic field, suggesting it uses the
Earth's magnetic field.
Moths exhibit a tendency to circle artificial lights repeatedly. This
suggests they use a technique of celestial navigation called
transverse orientation. By maintaining a constant angular relationship
to a bright celestial light, such as the Moon, they can fly in a
straight line. Celestial objects are so far away, even after traveling
great distances, the change in angle between the moth and the light
source is negligible; further, the moon will always be in the upper
part of the visual field or on the horizon. When a moth encounters a
much closer artificial light and uses it for navigation, the angle
changes noticeably after only a short distance, in addition to being
often below the horizon. The moth instinctively attempts to correct by
turning toward the light, causing airborne moths to come plummeting
downwards, and at close range, which results in a spiral flight path
that gets closer and closer to the light source. Other
explanations have been suggested, such as the idea that moths may be
impaired with a visual distortion called a
Mach band by Henry Hsiao in
1972. He stated that they fly towards the darkest part of the sky in
pursuit of safety, thus are inclined to circle ambient objects in the
Mach band region.
Monarch butterflies, seen in a cluster in Santa Cruz, California,
where the western population migrates for the winter
Lepidopteran migration is typically seasonal, as the insects moving to
escape dry seasons or other disadvantageous conditions. Most
lepidopterans that migrate are butterflies, and the distance travelled
varies. Some butterflies that migrate include the mourning cloak,
painted lady, American lady, red admiral, and the common
buckeye.:29–30 A notable species of moth that migrates long
distances is the bogong moth. The most well-known migrations are
those of the eastern population of the monarch butterfly from Mexico
to northern United States and southern Canada, a distance of about
4,000–4,800 km (2,500–3,000 mi). Other well-known
migratory species include the painted lady and several of the danaine
butterflies. Spectacular and large-scale migrations associated with
the monsoons are seen in peninsular India. Migrations have been
studied in more recent times using wing tags and stable hydrogen
Moths also undertake migrations, an example being the uraniids. Urania
fulgens undergoes population explosions and massive migrations that
may be not surpassed by any other insect in the Neotropics. In Costa
Rica and Panama, the first population movements may begin in July and
early August and depending on the year, may be very massive,
continuing unabated for as long as five months.
Group of Melitaea athalia near Warka, Poland
Pheromones are commonly involved in mating rituals among species,
especially moths, but they are also an important aspect of other forms
of communication. Usually, the pheromones are produced by either the
male or the female and detected by members of the opposite sex with
their antennae. In many species, a gland between the eighth and
ninth segments under the abdomen in the female produces the
pheromones. Communication can also occur through stridulation, or
producing sounds by rubbing various parts of the body together.
Moths are known to engage in acoustic forms of communication, most
often as courtship, attracting mates using sound or vibration. Like
most other insects, moths pick up these sounds using tympanic
membranes in their abdomens. An example is that of the polka-dot
wasp moth (Syntomeida epilais), which produces sounds with a frequency
above that normally detectable by humans (about 20 kHz). These
sounds also function as tactile communication, or communication
through touch, as they stridulate, or vibrate a substrate like leaves
Most moths lack bright colors, as many species use coloration as
camouflage, but butterflies engage in visual communication. Female
cabbage butterflies, for example, use ultraviolet light to
communicate, with scales colored in this range on the dorsal wing
surface. When they fly, each down stroke of the wing creates a brief
flash of ultraviolet light which the males apparently recognize as the
flight signature of a potential mate. These flashes from the wings may
attract several males that engage in aerial courtship displays.
Moths and butterflies are important in the natural ecosystem. They are
integral participants in the food chain; having co-evolved with
flowering plants and predators, lepidopteran species have formed a
network of trophic relationships between autotrophs and heterotrophs,
which are included in the stages of
Lepidoptera larvae, pupae, and
Larvae and pupae are links in the diets of birds and parasitic
entomophagous insects. The adults are included in food webs in a much
broader range of consumers (including birds, small mammals, reptiles,
Defense and predation
Main article: Defense in insects
Papilio machaon caterpillar showing the osmeterium, which emits
unpleasant smells to ward off predators
Lepidopteran species are soft bodied, fragile, and almost defenseless,
while the immature stages move slowly or are immobile, hence all
stages are exposed to predation. Adult butterflies and moths are
preyed upon by birds, bats, lizards, amphibians, dragonflies, and
spiders. Caterpillars and pupae fall prey not only to birds, but also
to invertebrate predators and small mammals, as well as fungi and
Parasitoid and parasitic wasps and flies may lay eggs in the
caterpillar, which eventually kill it as they hatch inside its body
and eat its tissues. Insect-eating birds are probably the largest
predators. Lepidoptera, especially the immature stages, are an
ecologically important food to many insectivorous birds, such as the
great tit in Europe.
An "evolutionary arms race" can be seen between predator and prey
Lepidoptera have developed a number of strategies for
defense and protection, including evolution of morphological
characters and changes in ecological lifestyles and behaviors. These
include aposematism, mimicry, camouflage, and development of threat
patterns and displays. Only a few birds, such as the nightjars,
hunt nocturnal lepidopterans. Their main predators are bats. Again, an
"evolutionary race" exists, which has led to numerous evolutionary
adaptations of moths to escape from their main predators, such as the
ability to hear ultrasonic sounds, or even to emit sounds in some
cases. Lepidopteran eggs are also preyed upon. Some caterpillars, such
as the zebra swallowtail butterfly larvae, are cannibalistic.
Some species of
Lepidoptera are poisonous to predators, such as the
monarch butterfly in the Americas,
Atrophaneura species (roses,
windmills, etc.) in Asia, as well as Papilio antimachus, and the
birdwings, the largest butterflies in Africa and Asia, respectively.
They obtain their toxicity by sequestering the chemicals from the
plants they eat into their own tissues. Some
their own toxins. Predators that eat poisonous butterflies and moths
may become sick and vomit violently, learning not to eat those
species. A predator which has previously eaten a poisonous
lepidopteran may avoid other species with similar markings in the
future, thus saving many other species, as well. Toxic
butterflies and larvae tend to develop bright colors and striking
patterns as an indicator to predators about their toxicity. This
phenomenon is known as aposematism. Some caterpillars, especially
members of Papilionidae, contain an osmeterium, a Y-shaped protrusible
gland found in the prothoracic segment of the larvae. When
threatened, the caterpillar emits unpleasant smells from the organ to
ward off the predators.
Camouflage is also an important defense strategy, which involves the
use of coloration or shape to blend into the surrounding environment.
Some lepidopteran species blend with their surroundings, making them
difficult to spot by predators. Caterpillars can exhibit shades of
green that match its host plant. Others look like inedible objects,
such as twigs or leaves. For instance, the mourning cloak fades into
the backdrop of trees when it folds its wings back. The larvae of some
species, such as the common Mormon (Papilio polytes) and the western
tiger swallowtail look like bird droppings. For example, adult
Sesiidae species (also known as clearwing moths) have a general
appearance sufficiently similar to a wasp or hornet to make it likely
the moths gain a reduction in predation by Batesian mimicry.
Eyespots are a type of automimicry used by some butterflies and moths.
In butterflies, the spots are composed of concentric rings of scales
in different colors. The proposed role of the eyespots is to deflect
attention of predators. Their resemblance to eyes provokes the
predator's instinct to attack these wing patterns.
Müllerian mimicry complexes are commonly found in
Lepidoptera. Genetic polymorphism and natural selection give rise to
otherwise edible species (the mimic) gaining a survival advantage by
resembling inedible species (the model). Such a mimicry complex is
referred to as Batesian and is most commonly known in the example
between the limenitidine viceroy butterfly in relation to the inedible
danaine monarch. The viceroy is, in fact, more toxic than the monarch
and this resemblance should be considered as a case of Müllerian
mimicry. In Müllerian mimicry, inedible species, usually within a
taxonomic order, find it advantageous to resemble each other so as to
reduce the sampling rate by predators that need to learn about the
insects' inedibility. Taxa from the toxic genus
Heliconius form one of
the most well-known Müllerian complexes. The adults of the
various species now resemble each other so well, the species cannot be
distinguished without close morphological observation and, in some
cases, dissection or genetic analysis.
Moths evidently are able to hear the range emitted by bats, which in
effect causes flying moths to make evasive maneuvers because bats are
a main predator of moths. Ultrasonic frequencies trigger a reflex
action in the noctuid moth that cause it to drop a few inches in its
flight to evade attack. Tiger moths in a defense emit clicks
within the same range of the bats, which interfere with the bats and
foil their attempts to echolocate it.
A day-flying hummingbird hawk-moth drinking nectar from a species of
Most species of
Lepidoptera engage in some form of entomophily (more
specifically psychophily and phalaenophily for butterflies and moths,
respectively), or the pollination of flowers. Most adult
butterflies and moths feed on the nectar inside flowers, using their
probosces to reach the nectar hidden at the base of the petals. In the
process, the adults brush against the flowers' stamens, on which the
reproductive pollen is made and stored. The pollen is transferred on
appendages on the adults, which fly to the next flower to feed and
unwittingly deposit the pollen on the stigma of the next flower, where
the pollen germinates and fertilizes the seeds.:813–814
Flowers pollinated by butterflies tend to be large and flamboyant,
pink or lavender in color, frequently having a landing area, and
usually scented, as butterflies are typically day-flying. Since
butterflies do not digest pollen (except for heliconid species,)
more nectar is offered than pollen. The flowers have simple nectar
guides, with the nectaries usually hidden in narrow tubes or spurs,
reached by the long "tongue" of the butterflies.
Butterflies such as
Thymelicus flavus have been observed to engage in flower constancy,
which means they are more likely to transfer pollen to other
conspecific plants. This can be beneficial for the plants being
pollinated, as flower constancy prevents the loss of pollen during
different flights and the pollinators from clogging stigmas with
pollen of other flower species.
Among the more important moth pollinator groups are the hawk moths of
the family Sphingidae. Their behavior is similar to hummingbirds,
i.e., using rapid wing beats to hover in front of flowers. Most hawk
moths are nocturnal or crepuscular, so moth-pollinated flowers (e.g.,
Silene latifolia ) tend to be white, night-opening, large, and showy
with tubular corollae and a strong, sweet scent produced in the
evening, night, or early morning. A lot of nectar is produced to fuel
the high metabolic rates needed to power their flight. Other moths
(e.g., noctuids, geometrids, pyralids) fly slowly and settle on the
flower. They do not require as much nectar as the fast-flying hawk
moths, and the flowers tend to be small (though they may be aggregated
Tobacco hornworm caterpillar (Manduca sexta) parasitized by Braconidae
Mutualism is a form of biological interaction wherein each individual
involved benefits in some way. An example of a mutualistic
relationship would be that shared by yucca moths (Tegeculidae) and
their host, yucca flowers (Liliaceae). Female yucca moths enter the
host flowers, collect the pollen into a ball using specialized
maxillary palps, then move to the apex of the pistil, where pollen is
deposited on the stigma, and lay eggs into the base of the pistil
where seeds will develop. The larvae develop in the fruit pod and feed
on a portion of the seeds. Thus, both insect and plant benefit,
forming a highly mutualistic relationship.:814 Another form of
mutualism occurs between some larvae of butterflies and certain
species of ants (e. g. Lycaenidae). The larvae communicate with the
ants using vibrations transmitted through a substrate, such as the
wood of a tree or stems, as well as using chemical signals. The
ants provide some degree of protection to these larvae and they in
turn gather honeydew secretions.
Parasitoid larvae exits from the fox moth caterpillar
Only 42 species of parasitoid lepidopterans are known (1 Pyralidae; 40
Epipyropidae).:748 The larvae of the greater and lesser wax moths
feed on the honeycomb inside bee nests and may become pests; they are
also found in bumblebee and wasp nests, albeit to a lesser extent. In
northern Europe, the wax moth is regarded as the most serious
parasitoid of the bumblebee, and is found only in bumblebee nests. In
some areas in southern England, as much as 80% of nests can be
destroyed. Other parasitic larvae are known to prey upon cicadas
and leaf hoppers.
The different parasitoids affecting the gypsy moth (Lymantaria
dispar): The stage they affect and eventually kill and its duration
are denoted by arrows.
In reverse, moths and butterflies may be subject to parasitic wasps
and flies, which may lay eggs on the caterpillars, which hatch and
feed inside its body, resulting in death. Although, in a form of
parasitism called idiobiont, the adult paralyzes the host, so as not
to kill it but for it to live as long as possible, for the parasitic
larvae to benefit the most. In another form of parasitism, koinobiont,
the species live off their hosts while inside (endoparasitic). These
parasites live inside the host caterpillar throughout its life cycle,
or may affect it later on as an adult. In other orders, koinobionts
include flies, a majority of coleopteran, and many hymenopteran
parasitoids.:748–749 Some species may be subject to a variety of
parasites, such as the gypsy moth (Lymantaria dispar), which is
attacked by a series of 13 species, in six different taxa throughout
its life cycle.:750
In response to a parsitoid egg or larva in the caterpillar's body, the
plasmatocytes, or simply the host's cells can form a multilayered
capsule that eventually causes the endoparasite to asphyxiate. The
process, called encapsulation, is one of the caterpillar's only means
of defense against parasitoids.:748
Other biological interactions
A few species of
Lepidoptera are secondary consumers, or predators.
These species typically prey upon the eggs of other insects, aphids,
scale insects, or ant larvae.:567 Some caterpillars are cannibals,
and others prey on caterpillars of other species (e. g. Hawaiian
Eupithecia ). Those of the 15 species in
Eupithecia that mirror
inchworms, are the only known species of butterflies and moths that
are ambush predators. Four species are known to eat snails. For
example, the Hawaiian caterpillar, (H. molluscivora), uses silk traps,
in a manner similar to that of spiders, to capture certain species of
snails (typically Tornatellides).
Larvae of some species of moths in the Tineidae, Gelechioidae, and
Noctuidae (family/superfamily/families, respectively), besides others,
feed on detritus, or dead organic material, such as fallen leaves and
fruit, fungi, and animal products, and turn it into humus.:567
Well-known species include the cloth moths (Tineola bisselliella,
Tinea pellionella, and Trichophaga tapetzella), which feed on detritus
containing keratin, including hair, feathers, cobwebs, bird nests
(particularly of domestic pigeons, Columba livia domestica) and fruits
or vegetables. These species are important to ecosystems as they
remove substances that would otherwise take a long time to
In 2015 it was reported that wasp bracovirus
DNA was present in
Lepidoptera such as monarch butterflies, silkworms and moths.
These were described in some newspaper articles as examples of a
naturally occurring genetically engineered insects.
Evolution and systematics
History of study
Main article: Lepidopterology
Lepidoptera collection in Cherni Osam Natural Sciences Museum, Troyan,
Systema Naturae (1758) recognized three divisions of the
Lepidoptera: Papilio, Sphinx and Phalaena, with seven subgroups in
Phalaena. These persist today as 9 of the superfamilies of
Lepidoptera. Other works on classification followed including those by
Michael Denis &
Ignaz Schiffermüller (1775), Johan Christian
Fabricius (1775) and
Pierre André Latreille
Pierre André Latreille (1796). Jacob Hübner
described many genera, and the lepidopteran genera were catalogued by
Ferdinand Ochsenheimer and
Georg Friedrich Treitschke
Georg Friedrich Treitschke in a series of
volumes on the lepidopteran fauna of Europe published between 1807 and
Gottlieb August Wilhelm Herrich-Schäffer
Gottlieb August Wilhelm Herrich-Schäffer (several volumes,
Edward Meyrick (1895) based their classifications
primarily on wing venation. Sir
George Francis Hampson worked on the
microlepidoptera during this period and Philipp Christoph Zeller
published The Natural History of the Tineinae also on microlepidoptera
Among the first entomologists to study fossil insects and their
Samuel Hubbard Scudder
Samuel Hubbard Scudder (1837–1911), who worked on
butterflies. He published a study of the Florissant deposits of
Colorado, including the exceptionally preserved
Andreas V. Martynov (1879–1938) recognized the close relationship
Trichoptera in his studies on phylogeny.
Major contributions in the 20th century included the creation of the
monotrysia and ditrysia (based on female genital structure) by Borner
in 1925 and 1939.
Willi Hennig (1913–1976) developed the
cladistic methodology and applied it to insect phylogeny. Niels P.
Kristensen, E. S. Nielsen and D. R. Davis studied the relationships
among monotrysian families and Kristensen worked more generally on
insect phylogeny and higher
Lepidoptera too. While it is often
found that DNA-based phylogenies differ from those based on
morphology, this has not been the case for the Lepidoptera; DNA
phylogenies correspond to a large extent to morphology-based
Many attempts have been made to group the superfamilies of the
Lepidoptera into natural groups, most of which fail because one of the
two groups is not monophyletic:
Microlepidoptera and Macrolepidoptera,
Heterocera and Rhopalocera, Jugatae and Frenatae,
Main article: Prehistoric Lepidoptera
1887 engraving of
Prodryas persephone, a fossil lepidopteran from the
The fossil record for
Lepidoptera is lacking in comparison to other
winged species, and tends not to be as common as some other insects in
habitats that are most conducive to fossilization, such as lakes and
ponds; their juvenile stage has only the head capsule as a hard part
that might be preserved. The location and abundance of the most common
moth species are indicative that mass migrations of moths occurred
over the Palaeogene North Sea, which is why there is a serious lack of
moth fossils. Yet there are fossils, some preserved in amber and
some in very fine sediments.
Leaf mines are also seen in fossil
leaves, although the interpretation of them is tricky.
Putative fossil stem group representatives of
Trichoptera and Lepidoptera) are known from the
Triassic.:567 The earliest known lepidopteran fossils are
fossilized scales from the Triassic-
Jurassic boundary. They were found
as rare palynological elements in the sediments of the
Jurassic boundary from the cored Schandelah-1 well, drilled
Braunschweig in northern Germany. This pushes back the fossil
record and origin of glossatan lepidopterans by about 70 million
years, supporting molecular estimates of a
Norian (ca 212 million
years) divergence of glossatan and non-glossatan lepidopterans. The
findings were reported in 2018 in the journal Science Advances. The
authors of the study proposed that lepidopterans evolved a proboscis
as an adaptation to drink from droplets and thin films of water for
maintaining their fluid balance in the hot and arid climate of the
The earliest described lepidopteran taxon is
Archaeolepis mane, a
primitive moth-like species from the Jurassic, dated back to around
190 million years ago, and known only from three wings found in
Dorset, UK. The wings show scales with parallel grooves under a
scanning electron microscope and a characteristic wing venation
pattern shared with
Trichoptera (caddisflies). Only two more
Jurassic lepidopteran fossils have been found, as well as 13
sets from the Cretaceous, which all belong to primitive moth-like
Many more fossils are found from the Tertiary, and particularly the
Eocene Baltic amber. The oldest genuine butterflies of the superfamily
Papilionoidea have been found in the
of Denmark. The best preserved fossil lepidopteran is the Eocene
Prodryas persephone from the Florissant Fossil Beds.
Phylogenetic hypothesis of major lepidopteran lineages superimposed on
the geologic time scale.
Angiosperm radiation spans
130 to 95 million years ago from the earliest angiosperms,
to angiosperm domination of vegetation.
Trichoptera (caddisflies) are sister groups, sharing
many similarities that are lacking in others; for example the females
of both orders are heterogametic, meaning they have two different sex
chromosomes, whereas in most species the males are heterogametic and
the females have two identical sex chromosomes. The adults in both
orders display a particular wing venation pattern on their forewings.
The larvae in the two orders have mouth structures and glands with
which they make and manipulate silk.
Willi Hennig grouped the two
orders into the
Amphiesmenoptera superorder; they are sisters, and
together are sister to the extinct order Tarachoptera.
Lepidoptera descend from a diurnal moth-like common ancestor that
either fed on dead or living plants.
The cladogram, based on a 2008
DNA and protein analysis, shows the
order as a clade, sister to the Trichoptera, and more distantly
related to the
Diptera (true flies) and Mecoptera
part of Endopterygota
Diptera (true flies)
Boreidae (snow scorpionflies)
Lepidoptera (butterflies and moths)
Hymenoptera (sawflies, wasps, ants, bees)
Heterobathmiidae are the oldest
and most basal lineages of Lepidoptera. The adults of these families
do not have the curled tongue or proboscis, that are found in most
members order, but instead have chewing mandibles adapted for a
Micropterigidae larvae feed on leaves, fungi, or
liverworts (much like the Trichoptera). Adult
the pollen or spores of ferns. In the Agathiphagidae, larvae live
inside kauri pines and feed on seeds. In
Heterobathmiidae the larvae
feed on the leaves of Nothofagus, the southern beech tree. These
families also have mandibles in the pupal stage, which help the pupa
emerge from the seed or cocoon after metamorphosis.
Eriocraniidae have a short coiled proboscis in the adult stage,
and though they retain their pupal mandibles with which they escaped
the cocoon, their mandibles are non-functional thereafter. Most of
these non-ditrysian families, are primarily leaf miners in the larval
stage. In addition to the proboscis, there is a change in the scales
among these basal lineages, with later lineages showing more complex
With the evolution of the
Ditrysia in the mid-Cretaceous, there was a
major reproductive change. The Ditrysia, which comprise 98% of the
Lepidoptera, have two separate openings for reproduction in the
females (as well as a third opening for excretion), one for mating,
and one for laying eggs. The two are linked internally by a seminal
duct. (In more basal lineages there is one cloaca, or later, two
openings and an external sperm canal.) Of the early lineages of
Gelechioidea are mostly leaf miners, but
more recent lineages feed externally. In the Tineoidea, most species
feed on plant and animal detritus and fungi, and build shelters in the
Yponomeutoidea is the first group to have significant numbers of
species whose larvae feed on herbaceous plants, as opposed to woody
plants. They evolved about the time that flowering plants
underwent an expansive adaptive radiation in the mid-Cretaceous, and
Gelechioidea that evolved at this time also have great diversity.
Whether the processes involved coevolution or sequential evolution,
the diversity of the
Lepidoptera and the angiosperms increased
In the so-called "macrolepidoptera", which constitutes about 60% of
lepidopteran species, there was a general increase in size, better
flying ability (via changes in wing shape and linkage of the forewings
and hindwings), reduction in the adult mandibles, and a change in the
arrangement of the crochets (hooks) on the larval prolegs, perhaps to
improve the grip on the host plant. Many also have tympanal
organs, that allow them to hear. These organs evolved eight times, at
least, because they occur on different body parts and have structural
differences. The main lineages in the macrolepidoptera are the
Noctuoidea, Bombycoidea, Lasiocampidae, Mimallonoidea, Geometroidea
be a monophyletic group. The Rhopalocera, comprising the
Hesperioidea (skippers), and the
Hedyloidea (moth-butterflies), are the most recently evolved.
There is quite a good fossil record for this group, with the oldest
skipper dating from 56 million years ago.
Taxonomy of the Lepidoptera
Taxonomy of the Lepidoptera and Lepidopteran diversity
Taxonomy is the classification of species in selected taxa, the
process of naming being called nomenclature. There are over 120
families in Lepidoptera, in 45 to 48 superfamilies.
always been, historically, classified in five suborders, one of which
is of primitive moths that never lost the morphological features of
its ancestors. The rest of the moths and butterflies make up
ninety-eight percent of the other taxa, making Ditrysia. More
recently, findings of new taxa, larvae and pupa have aided in
detailing the relationships of primitive taxa, phylogenetic analysis
showing the primitive lineages to be paraphyletic compared to the rest
Lepidoptera lineages. Recently lepidopterists have abandoned clades
like suborders, and those between orders and superfamilies.:569
Zeugloptera is a clade with
Micropterigoidea being its only
superfamily, containing the single family Micropterigidae.
Micropterigoidea are practically living fossils, being one of the most
primitive lepidopteran groups, still retaining chewing mouthparts
(mandibles) in adults, unlike other clades of butterflies and moths.
About 120 species are known worldwide, with more than half the species
in the genus
Micropteryx in the Paleartic region. There are only two
known in North America (Epimartyria), with many more being found Asia
and the southwest Pacific, particularly New Zealand with about 50
Glossata contains a majority of the species, with the most obvious
difference is non-functioning mandibles, and elongated maxillary
galeae or the proboscis. The basal clades still retaining some of the
ancestral features of the wings such as similarly shaped fore- and
hindwings with relatively complete venation.
Glossata also contains
the division Ditrysia, which contains 98% of all described species in
Aglossata it is the second most primitive lineage of Lepidoptera;
being first described in 1952 by Lionel Jack Dumbleton. Agathiphagidae
Heterobathmiidae are the only families in Aglossata.
Agathiphagidae only contains about 2 species in its genus
Agathiphaga queenslandensis and Agathiphaga
vitiensis, being found along the north-eastern coast of Queensland,
Australia, and in
Vanuatu and the Solomon Islands,
Heterobathmiina was first described by Kristensen and Nielsen in 1979.
There are about 10 species, which are day-flying, metallic moths,
confined to southern South America, the adults eat the pollen of
Nothofagus or southern beech and the larvae mine the
Relationship to people
Death's-head hawkmoth (Acherontia lachesis), an old bleached specimen
still showing the classical skull pattern on the thorax
Artistic depictions of butterflies have been used in many cultures
including as early as 3500 years ago, in Egyptian hieroglyphs.
Today, butterflies are widely used in various objects of art and
jewelry: mounted in frames, embedded in resin, displayed in bottles,
laminated in paper, and in some mixed media artworks and
Butterflies have also inspired the "butterfly fairy"
as an art and fictional character.
In many cultures the soul of a dead person is associated with the
butterfly, for example in Ancient Greece, where the word for butterfly
ψυχή (psyche) also means soul and breath. In Latin, as in Ancient
Greece, the word for "butterfly" papilio was associated with the soul
of the dead. The skull-like marking on the thorax of the
death's-head hawkmoth has helped these moths, particularly A. atropos,
earn a negative reputation, such as associations with the supernatural
and evil. The moth has been prominently featured in art and movies
Un Chien Andalou
Un Chien Andalou (by Buñuel and Dalí) and The Silence of the
Lambs, and in the artwork of the Japanese metal band Sigh's album Hail
Horror Hail. According to Kwaidan: Stories and Studies of Strange
Things, by 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 guestroom and perches behind the bamboo screen, the person
whom you most love is coming to see you. However, large numbers of
butterflies are viewed as bad omens. When
Taira no Masakado
Taira no Masakado was
secretly preparing for his famous revolt, there appeared in
vast a swarm of butterflies that the people were frightened—thinking
the apparition to be a portent of coming evil.
In the ancient Mesoamerican city of Teotihuacan, the brilliantly
colored image of the butterfly was carved into many temples,
buildings, jewelry, and emblazoned on incense burners in particular.
The butterfly was sometimes depicted with the maw of a jaguar and some
species were considered to be the reincarnations of the souls of dead
warriors. The close association of butterflies to fire and warfare
persisted through to the
Aztec civilization and evidence of similar
jaguar-butterfly images has been found among the Zapotec, and Maya
Caterpillar hatchling of the grey dagger (Acronicta psi) eating leaves
from a tree
The larvae of many lepidopteran species are major pests in
agriculture. Some of the major pests include Tortricidae, Noctuidae,
and Pyralidae. The larvae of the
Noctuidae genus Spodoptera
Helicoverpa (corn earworm), or
Pieris brassicae can cause
extensive damage to certain crops.
Helicoverpa zea larvae (cotton
bollworms or tomato fruitworms) are polyphagous, meaning they eat a
variety of crops, including tomatoes and cotton. Peridroma saucia
(variegated cutworms) are described as one of the most damaging pests
to gardens, with the ability to destroy entire gardens and fields in a
matter of days.
Butterflies and moths are one of the largest taxa to solely feed and
be dependent on living plants, in terms of the number of species, and
they are in many ecosystems, making up the largest biomass to do so.
In many species, the female may produce anywhere from 200 to 600 eggs,
while in some others it may go as high as 30,000 eggs in one day. This
can create many problems for agriculture, where many caterpillars can
affect acres of vegetation. Some reports estimate that there have been
over 80,000 caterpillars of several different taxa feeding on a single
oak tree. In some cases, phytophagous larvae can lead to the
destruction of entire trees in relatively short periods of
Ecological ways of removing pest
Lepidoptera species are becoming more
economically viable, as research has shown ways like introducing
parasitic wasp and flies. For example, Sarcophaga aldrichi, a fly
which deposited larvae feed upon the pupae of the forest tent
caterpillar moth. Pesticides can affect other species other than the
species they are targeted to eliminate, damaging the natural
ecosystem. Another good biological pest control method is the use
of pheromone traps. A pheromone trap is a type of insect trap that
uses pheromones to lure insects. Sex pheromones and aggregating
pheromones are the most common types used. A pheromone-impregnated
lure is encased in a conventional trap such as a Delta trap, water-pan
trap, or funnel trap.
Species of moths that are detritivores would naturally eat detritus
containing keratin, such as hairs or feathers. Well known species are
cloth moths (T. bisselliella, T. pellionella, and T. tapetzella),
feeding on foodstuffs that people find economically important, such as
cotton, linen, silk and wool fabrics as well as furs; furthermore they
have been found on shed feathers and hair, bran, semolina and flour
(possibly preferring wheat flour), biscuits, casein, and insect
specimens in museums.
Even though most butterflies and moths affect the economy negatively,
some species are a valuable economic resource. The most prominent
example is that of the domesticated silkworm moth (Bombyx mori), the
larvae of which make their cocoons out of silk, which can be spun into
Silk is and has been an important economic resource throughout
history. The species
Bombyx mori has been domesticated to the point
where it is completely dependent on mankind for survival. A
number of wild moths such as Bombyx mandarina, and
besides others, provide commercially important silks.
The preference of the larvae of most lepidopteran species to feed on a
single species or limited range of plants is used as a mechanism for
biological control of weeds in place of herbicides. The pyralid cactus
moth was introduced from Argentina to Australia, where it successfully
suppressed millions of acres of prickly pear cactus.:567 Another
species of the Pyralidae, called the alligator weed stem borer (Arcola
malloi), was used to control the aquatic plant known as alligator weed
(Alternanthera philoxeroides) in conjunction with the alligator weed
flea beetle; in this case, the two insects work in synergy and the
weed rarely recovers.
Breeding butterflies and moths, or butterfly gardening/rearing, has
become an ecologically viable process of introducing species into the
ecosystem to benefit it.
Butterfly ranching in Papua New Guinea
permits nationals of that country to "farm" economically valuable
insect species for the collectors market in an ecologically
Beondegi, silkworm pupae steamed or boiled and seasoned for taste, for
sale by a street vendor in South Korea
Lepidoptera feature prominently in entomophagy as food items on almost
every continent. While in most cases, adults, larvae or pupae are
eaten as staples by indigenous people, beondegi or silkworm pupae are
eaten as a snack in Korean cuisine while
Maguey worm is
considered a delicacy in Mexico. In some parts of Huasteca, the
silk nests of the
Madrone butterfly are maintained on the edge of roof
tops of houses for consumption. In the
Carnia region of Italy,
children catch and eat ingluvies of the toxic Zygaena moths in early
summer. The ingluvies, despite having a very low cyanogenic content,
serve as a convenient, supplementary source of sugar to the children
who can include this resource as a seasonal delicacy at minimum
Some larvae of both moths and butterflies have a form of hair that has
been known to be a cause of human health problems.
sometimes have toxins in them and species from approximately 12
families of moths or butterflies worldwide can inflict serious human
injuries (urticarial dermatitis and atopic asthma to osteochondritis,
consumption coagulopathy, renal failure, and intracerebral
hemorrhage). Skin rashes are the most common, but there have been
Lonomia is a frequent cause of envenomation in humans
in Brazil, with 354 cases reported between 1989 and 2005. Lethality
ranging up to 20% with death caused most often by intracranial
These hairs have also been known to cause kerato-conjunctivitis. The
sharp barbs on the end of caterpillar hairs can get lodged in soft
tissues and mucous membranes such as the eyes. Once they enter such
tissues, they can be difficult to extract, often exacerbating the
problem as they migrate across the membrane. This becomes a
particular problem in an indoor setting. The hairs easily enter
buildings through ventilation systems and accumulate in indoor
environments because of their small size, which makes it difficult for
them to be vented out. This accumulation increases the risk of human
contact in indoor environments.
Butterflies and moths portal
Comparison of butterflies and moths
Lepidoptera in the 10th edition of Systema Naturae
McGuire Center for
Lepidoptera and Biodiversity, University of Florida
Societas Europaea Lepidopterologica
List of butterflies of Australia
List of butterflies of Canada
List of butterflies of Great Britain
List of butterflies of India
List of butterflies of Menorca
List of butterflies of North America
List of butterflies of Taiwan
List of butterflies of Tobago
List of fritillaries (butterflies)
List of moths
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Wikispecies has information related to Lepidoptera
The Wikibook Dichotomous Key has a page on the topic of: Lepidoptera
Wikimedia Commons has media related to Lepidoptera.
Lepidoptera at Insects (Insecta) of the World
"Lepidoptera". Integrated Taxonomic Information System.
Caught Between the Pages: Treasures from the Franclemont Collection
Online virtual exhibit featuring a selection of historic entomological
writings and images from the Comstock Library of Entomology at Cornell
Butterflies and Moths
Butterflies of Bulgaria
Butterflies of Canada
Photography of European
Butterflies and Moths
Lepidoptera of French Antilles
Butterflies of India
A Check List of
Butterflies in Indo-China
Moths of Jamaica
Japmoth Japanese moths. Access images via the numbers on the left.
Moths in the Netherlands
Moths of Northern Ireland
Butterflies of Asian Russia
Larvae and Pupae butterflies and moths. Spain
Butterflies of Asturias – Spain
Butterflies of Turkey
Archaeognatha (jumping bristletails)
Thysanura (Zygentoma) (silverfish, firebrats)
Odonata (dragonflies, damselflies)
Phasmatodea (stick and leaf insects)
Notoptera (ice-crawlers, gladiators)
Orthoptera (crickets, wetas, grasshoppers, locusts)
Zoraptera (angel insects)
Blattodea (cockroaches, termites)
Psocodea (barklice, lice)
Hemiptera (cicadas, aphids, true bugs)
Hymenoptera (sawflies, wasps, ants, bees)
Strepsiptera (twisted-winged parasites)
Megaloptera (alderflies, dobsonflies, fishflies)
Neuroptera (net-winged insects: lacewings, mantidflies, antlions)
Mecoptera (scorpionflies) +
Diptera (gnats, mosquitoes, flies)
Lepidoptera (moths, butterflies)
Four most speciose orders are marked in bold
Italic are paraphyletic groups
Based on Sasaki et al. (2013)
Extinct incertae sedis families and genera are marked in italic
Micropterigidae (mandibulate archaic moths)
Agathiphagidae (kauri moths)
Acanthopteroctetidae (archaic sun moths)
Neopseustidae (archaic bell moths)
Hepialidae (swift moths, ghost moths)
Neotheoridae (Amazonian primitive ghost moths)
Palaeosetidae (miniature ghost moths)
Prototheoridae (African primitive ghost moths)
Mnesarchaeidae (New Zealand primitive moths)
Adelidae (fairy longhorn moths)
Prodoxidae (yucca moths)
Andesianidae (Andean endemic moths)
Nepticulidae (pigmy, or midget moths)
Opostegidae (white eyecap moths)
Palaephatidae (Gondwanaland moths)
Tischeriidae (trumpet leaf miner moths)
Acrolophidae (burrowing webworm moths)
Eriocottidae (Old World spiny-winged moths)
Psychidae (bagworm moths)
Tineidae (fungus moths)
Bucculatricidae (ribbed cocoon makers)
Douglasiidae (Douglas moths)
Acrolepiidae (false diamondback moths)
Glyphipterigidae (sedge moths)
Yponomeutidae (ermine moths)
Coleophoridae (case-bearers, case moths)
Cosmopterigidae (cosmet moths)
Elachistidae (grass-miner moths)
Gelechiidae (twirler moths)
Lecithoceridae (long-horned moths)
Momphidae (mompha moths)
Oecophoridae (concealer moths)
Scythrididae (flower moths)
Xyloryctidae (timber moths)
Zygaenidae (burnet, forester, or smoky moths)
Megalopygidae (flannel moths)
Epipyropidae (planthopper parasite moths)
Dalceridae (slug caterpillars)
Limacodidae (slug, or cup moths)
Cossidae (carpenter millers, or goat moths)
Dudgeoneidae (dudgeon carpenter moths)
Brachodidae (little bear moths)
Castniidae (castniid moths: giant butterfly-moths, sun moths)
Sesiidae (clearwing moths)
Choreutidae (metalmark moths)
Tortricidae (tortrix moths)
Urodidae (false burnet moths)
Schreckensteiniidae (bristle-legged moths)
Epermeniidae (fringe-tufted moths)
Alucitidae (many-plumed moths)
Tineodidae (false plume moths)
Pterophoridae (plume moths)
Copromorphidae (tropical fruitworm moths)
Carposinidae (fruitworm moths)
Hyblaeidae (teak moths)
Pyralidae (snout moths)
Crambidae (grass moth)
Thyrididae (picture-winged leaf moths)
Mimallonidae (sack bearer moths)
Lasiocampidae (eggars, snout moths, or lappet moths)
Anthelidae (Australian lappet moth)
Bombycidae (silk moths)
Brahmaeidae (Brahmin moths)
Carthaeidae (Dryandra moth)
Endromidae (Kentish glory and relatives)
Sphingidae (hawk moths, sphinx moths and hornworms)
Erebidae (underwing, tiger, tussock, litter, snout, owlet moths)
Noctuidae (daggers, sallows, owlet moths, quakers, cutworms, darts)
Nolidae (tuft moths)
Notodontidae (prominents, kittens)
Epicopeiidae (oriental swallowtail moths)
Geometridae (geometer moths)
Cimeliidae (gold moths)
Callidulidae (Old World butterfly-moths)
Hedylidae (American moth-butterflies)
Lycaenidae (gossamer-winged butterflies: blues, coppers and relatives)
Nymphalidae (brush-footed, or four-footed butterflies)
Papilionidae (swallowtail butterflies)
Pieridae (whites, yellows, orangetips, sulphurs)
Monotrysia is not a clade.
Taxonomy of the Lepidoptera
Lists by region
Fauna Europaea: 7
BNF: cb11932240r (data)