Caterpillar

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For other uses, see Caterpillar (disambiguation).

Caterpillar of Papilio machaon

A monarch butterfly (Danaus plexippus) caterpillar feeding on a leaf of the swamp milkweed

Caterpillars /ˈkætərˌpɪlər/ are the larval stage of members of the order Lepidoptera (the insect
order comprising butterflies and moths).

As with most common names, the application of the word is arbitrary and the larvae of sawflies
commonly are called caterpillars as well.[1][2] Both lepidopteran and symphytan larvae have
eruciform body shapes.

Caterpillars of most species are herbivorous (folivorous), but not all; some (about 1%) are
insectivorous, even cannibalistic. Some feed on other animal products; for example clothes
moths feed on wool, and horn moths feed on the hooves and horns of dead ungulates.

Caterpillars are typically voracious feeders and many of them are among the most serious of
agricultural pests. In fact many moth species are best known in their caterpillar stages because of
the damage they cause to fruits and other agricultural produce, whereas the moths are obscure
and do no direct harm. Conversely, various species of caterpillar are valued as sources of silk, as
human or animal food, or for biological control of pest plants.

Contents
 1 Etymology
 2 Description
 3 Defenses
o 3.1 Appearance
o 3.2 Chemical defenses
o 3.3 Defensive behaviors
o 3.4 Social behaviors and relationships with other insects
o 3.5 Predators
 4 Behavior
 5 Economic effects
 6 Human health
 7 In popular culture
 8 Gallery
 9 See also
 10 References
 11 External links

Etymology
The origins of the word "caterpillar" date from the early 16th century. They derive from Middle
English catirpel, catirpeller, probably an alteration of Old North French catepelose: cate, cat
(from Latin cattus) + pelose, hairy (from Latin pilōsus).[3]

The inchworm, or looper caterpillars from the family Geometridae are so named because of the
way they move, appearing to measure the earth (the word geometrid means earth-measurer in
Greek);[4] the primary reason for this unusual locomotion is the elimination of nearly all the
prolegs except the clasper on the terminal segment.

A geometrid caterpillar or inchworm

Description

Crochets on a caterpillar's prolegs

Larvae of Craesus septentrionalis, a sawfly showing 6 pairs of pro-legs.

Caterpillars have soft bodies that can grow rapidly between moults. Their size varies between
species and instars (moults) from as small as 1 mm up to 14 cm.[5] Some larvae of the order
Hymenoptera (ants, bees and wasps) can appear like the caterpillars of the Lepidoptera. Such
larvae are mainly seen in the sawfly suborder. However while these larvae superficially resemble
caterpillars, they can be distinguished by the presence of prolegs on every abdominal segment,
an absence of crochets or hooks on the prolegs (these are present on lepidopteran caterpillars),
one pair of prominent ocelli on the head capsule, and an absence of the upside-down Y-shaped
suture on the front of the head.[6]

Lepidopteran caterpillars can be differentiated from sawfly larvae by:

 the numbers of pairs of pro-legs; sawfly larvae have 6 or more pairs while caterpillars
have a maximum of 5 pairs.
 the number of stemmata (simple eyes); the sawfly larvae have only two,[7] while
caterpillars usually have six.
 the presence of crochets on the prolegs; these are absent in the sawflies.
 sawfly larvae have an invariably smooth head capsule with no cleavage lines, while
lepidopterous caterpillars bear an inverted "Y" or "V" (adfrontal suture).
Defenses

The saddleback caterpillar has urticating hair and aposematic colouring.

Many animals feed on caterpillars as they are rich in protein. As a result, caterpillars have
evolved various means of defense.

Caterpillars have evolved defenses against physical conditions such as cold, hot or dry
environmental conditions. Some Arctic species like Gynaephora groenlandica have special
basking and aggregation behaviours[8] apart from physiological adaptations to remain in a
dormant state.[9]

Appearance

Costa Rican hairy caterpillar. The spiny bristles are a self-defense mechanism

The appearance of a caterpillar can often repel a predator: its markings and certain body parts
can make it seem poisonous, or bigger in size and thus threatening, or non-edible. Some types of
caterpillars are indeed poisonous or distasteful and their bright coloring is aposematic. Others
may mimic dangerous caterpillars or other animals while not being dangerous themselves. Many
caterpillars are cryptically colored and resemble the plants on which they feed. An example of
caterpillars that use camouflage for defence is the species Nemoria arizonaria. If the caterpillars
hatch in the spring and feed on oak catkins they appear green. If they hatch in the summer they
appear dark colored, like oak twigs. The differential development is linked to the tannin content
in the diet.[10] Caterpillars may even have spines or growths that resemble plant parts such as
thorns. Some look like objects in the environment such as bird droppings.

Chemical defenses
More aggressive self-defense measures are taken by some caterpillars. These measures include
having spiny bristles or long fine hair-like setae with detachable tips that will irritate by lodging
in the skin or mucous membranes.[6] However some birds (such as cuckoos) will swallow even
the hairiest of caterpillars. Other caterpillars acquire toxins from their host plants that render
them unpalatable to most of their predators. For instance, ornate moth caterpillars utilize
pyrrolizidine alkaloids that they obtain from their food plants to deter predators.[11] The most
aggressive caterpillar defenses are bristles associated with venom glands. These bristles are
called urticating hairs. A venom which is among the most potent defensive chemicals in any
animal is produced by the South American silk moth genus Lonomia. Its venom is an
anticoagulant powerful enough to cause a human to hemorrhage to death (See Lonomiasis).[12]
This chemical is being investigated for potential medical applications. Most urticating hairs
range in effect from mild irritation to dermatitis. Example: Brown-tail moth.

Giant swallowtail caterpillar everting its osmeterium in defense

Plants contain toxins which protect them from herbivores, but some caterpillars have evolved
countermeasures which enable them to eat the leaves of such toxic plants. In addition to being
unaffected by the poison, the caterpillars sequester it in their body, making them highly toxic to
predators. The chemicals are also carried on into the adult stages. These toxic species, such as
the cinnabar moth (Tyria jacobaeae) and monarch (Danaus plexippus) caterpillars, usually
advertise themselves with the danger colors of red, yellow and black, often in bright stripes (see
aposematism). Any predator that attempts to eat a caterpillar with an aggressive defense
mechanism will learn and avoid future attempts.

Some caterpillars regurgitate acidic digestive juices at attacking enemies. Many papilionid larvae
produce bad smells from extrudable glands called osmeteria.

Defensive behaviors
Caterpillars linked together into a "train"

Many caterpillars display feeding behaviors which allow the caterpillar to remain hidden from
potential predators. Many feed in protected environments, such as enclosed inside silk galleries,
rolled leaves or by mining between the leaf surfaces.

Some caterpillars, like early instars of the tomato hornworm and tobacco hornworm, have long
"whip-like" organs attached to the ends of their body. The caterpillar wiggles these organs to
frighten away flies and predatory wasps.[13] Some caterpillars can evade predators by using a silk
line and dropping off from branches when disturbed. Many species thrash about violently when
disturbed to scare away potential predators. One species (Amorpha juglandis) even makes high
pitched whistles that can scare away birds.[14]

Social behaviors and relationships with other insects

Some caterpillars obtain protection by associating themselves with ants. The Lycaenid butterflies
are particularly well known for this. They communicate with their ant protectors by vibrations as
well as chemical means and typically provide food rewards.[15]

Some caterpillars are gregarious; large aggregations are believed to help in reducing the levels of
parasitization and predation.[16] Clusters amplify the signal of aposematic coloration, and
individuals may participate in group regurgitation or displays. Pine processionary
(Thaumetopoea pityocampa) caterpillars often link into a long train to move through trees and
over the ground. The head of the lead caterpillar is visible, but the other heads can appear
hidden.[17] Forest tent caterpillars cluster during periods of cold weather.

Predators

Caterpillars suffer predation from many animals. The European pied flycatcher is one species
that preys upon caterpillars. The flycatcher typically finds caterpillars among oak foliage. Paper
wasps, including those in the genus Polistes and Polybia catch caterpillars to feed their young
and themselves.

Behavior

A pasture day moth caterpillar feeding on capeweed

Caterpillars have been called "eating machines", and eat leaves voraciously. Most species shed
their skin four or five times as their bodies grow, and they eventually enter a pupal stage before
becoming adults.[18] Caterpillars grow very quickly; for instance, a tobacco hornworm will
increase its weight ten-thousandfold in less than twenty days. An adaptation that enables them to
eat so much is a mechanism in a specialized midgut that quickly transports ions to the lumen
(midgut cavity), to keep the potassium level higher in the midgut cavity than in the
hemolymph.[19]

A gypsy moth caterpillar

Most caterpillars are solely herbivorous. Many are restricted to feeding on one species of plant,
while others are polyphagous. Some, including the clothes moth, feed on detritus. Some are
predatory, and may prey on other species of caterpillars (e.g. Hawaiian Eupithecia). Others feed
on eggs of other insects, aphids, scale insects, or ant larvae. A few are parasitic on cicadas or leaf
hoppers (Epipyropidae).[20] Some Hawaiian caterpillars (Hyposmocoma molluscivora) use silk
traps to capture snails.[21]

Many caterpillars are nocturnal. For example, the "cutworms" (of the family Noctuidae) hide at
the base of plants during the day and only feed at night.[22] Others, such as gypsy moth
(Lymantria dispar) larvae, change their activity patterns depending on density and larval stage,
with more diurnal feeding in early instars and high densities.[23]

Economic effects

Hypsipyla grandela damages mahogany in Brazil

Caterpillars cause much damage, mainly by eating leaves. The propensity for damage is
enhanced by monocultural farming practices, especially where the caterpillar is specifically
adapted to the host plant under cultivation. The cotton bollworm causes enormous losses. Other
species eat food crops. Caterpillars have been the target of pest control through the use of
pesticides, biological control and agronomic practices. Many species have become resistant to
pesticides. Bacterial toxins such as those from Bacillus thuringiensis which are evolved to affect
the gut of Lepidoptera have been used in sprays of bacterial spores, toxin extracts and also by
incorporating genes to produce them within the host plants. These approaches are defeated over
time by the evolution of resistance mechanisms in the insects.[24]

Plants evolve mechanisms of resistance to being eaten by caterpillars, including the evolution of
chemical toxins and physical barriers such as hairs. Incorporating host plant resistance (HPR)
through plant breeding is another approach used in reducing the impact of caterpillars on crop
plants.[25]

Some caterpillars are used in industry. The silk industry is based on the silkworm caterpillar.

Human health

Buck moth caterpillar sting on a shin twenty-four hours after occurrence in south Louisiana. The
reddish mark covers an area about 20 mm at its widest point by about 70 mm in length.

Caterpillar hair can be a cause of human health problems. Caterpillar hairs sometimes have
venoms in them and species from approximately 12 families of moths or butterflies worldwide
can inflict serious human injuries ranging from urticarial dermatitis and atopic asthma to
osteochondritis, consumption coagulopathy, renal failure, and intracerebral hemorrhage.[26] Skin
rashes are the most common, but there have been fatalities.[27] Lonomia is a frequent cause of
envenomation in Brazil, with 354 cases reported between 1989 and 2005. Lethality ranging up to
20% with death caused most often by intracranial hemorrhage.[28]

Caterpillar hair has 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.[29]

This becomes a particular problem in an indoor setting. The hair 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.[30]
Caterpillars are a food source in some cultures. For example, in South Africa mopane worms are
eaten by the bushmen, and in China silkworms are considered a delicacy.

In popular culture

Wikimedia Commons has media related to Caterpillars in art.

William Blake's illustration of a caterpillar overlooking a child from his illustrated book For
Children The Gates of Paradise.[31]

A 1907 illustrations by Arthur Rackham of the Caterpillar talking to Alice in Alice's Adventures
in Wonderland

In the Old Testament of the Bible caterpillars are feared as pest that devour crops. They are part
of the "pestilence, blasting, mildew, locus" because of their association with the locust, thus they
are one of the plagues of Egypt. Jeremiah names them as one of the inhabitants of Babylon. The
English word caterpillar derives from the old French catepelose (hairy cat) but merged with the
piller (pillager). Caterpillars became a symbol for social dependents. Shakespeare's Bolingbroke
described King Richard's friends as "The caterpillars of the commonwealth, Which I have sworn
to weed and pluck away". In 1790 William Blake referenced this popular image in The Marriage
of Heaven and Hell when he attacked priests: "as the caterpillar chooses the fairest leaves to lay
her eggs on, so the priest lay his curse on the fairest joys".[32]
The role of caterpillars in the life stages of butterflies was badly understood. In 1679 Maria
Sibylla Merian published the first volume of The Caterpillars' Marvelous Transformation and
Strange Floral Food, which contained 50 illustrations and a description of insects, moths,
butterflies and their larvae.[33] An earlier popular publication on moths and butterflies, and their
caterpillars, by Jan Goedart had not included eggs in the life stages of European moths and
butterflies, because he had believed that caterpillars were generated from water. When Merian
published her study of caterpillars it was still widely believed that insects were spontaneously
generated. Merian's illustrations supported the findings of Francesco Redi, Marcello Malpighi
and Jan Swammerdam.[34]

Butterflies were regarded as symbol for the human soul since ancient time, and also in the
Christian tradition.[35] Goedart thus located his empirical observations on the transformation of
caterpillars into butterflies in the Christian tradition. As such he argued that the metamorphosis
from caterpillar into butterfly was a symbol, and even proof, of Christ's resurrection. He argued
"that from dead caterpillars emerge living animals; so it is equally true and miraculous, that our
dead and rotten corpses will rise from the grave."[36] Swammerdam, who in 1669 had
demonstrated that inside a caterpillar the rudiments of the future butterfly's limbs and wings
could be discerned, attacked the mystical and religious notion that the caterpillar died and the
butterfly subsequently resurrected.[37] As a militant Cartesian, Swammerdam attacked Goedart as
ridiculous, and when publishing his findings he proclaimed "here we witness the digression of
those who have tried to prove Resurrection of the Dead from these obviously natural and
comprehensible changes within the creature itself."[38]

Since then the metamorphoses of the caterpillar into a butterfly has in Western societies been
associated with countless human transformations in folktales and literature. There is no process
in the physical life of human beings that resembles this metamorphoses, and the symbol of the
caterpillar tends to depict a psychic transformation of a human. As such the caterpillar has in the
Christian tradition become a metaphor for being "born again".[39]

Famously, in Lewis Carroll's Alice's Adventures in Wonderland a caterpillar asks Alice "Who
are you?". When Alice comments on the caterpillar's inevitable transformation into a butterfly,
the caterpillar champions the position that in spite of changes it is still possible to know
something, and that Alice is the same Alice at the beginning and end of a considerable
interval.[40] When the Caterpillar asks Alice to clarify a point, the child replies "I'm afraid I can't
put it more clearly... for I can't but understand it myself, to begin with, and being so many
different sizes in a day is very confusing". Here Carroll satirizes René Descartes, the founder of
Cartesian philosophy, and his theory on innate ideas. Descartes argued that we are distracted by
urgent bodily stimuli that swamp the human mind in childhood. Descartes also theorised that
inherited preconceived opinions obstruct the human perception of the truth.[41]

More recent symbolic references to caterpillars in popular media include the Mad Men season 3
episode "The Fog", in which Betty Draper has a drug-induced dream, while in labor, that she
captures a caterpillar and holds it firmly in her hand.[42] In The Sopranos season 5 episode "The
Test Dream", Tony Soprano dreams that Ralph Cifaretto has a caterpillar on his bald head that
changes into a butterfly.
Gallery
Click left or right for a slide show.

Caterpillar of the spurge hawk-moth, near Binn, Valais, Switzerland at c. 2 km altitude.

See also
 Edible caterpillars
 Larval food plants of Lepidoptera
 Lepidopterism - caterpillar dermatitis
 List of pests and diseases of roses
 Sericulture

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42.  What's Alan Watching?: Mad Men, "The Fog"

External links

Wikimedia Commons has media related to Caterpillar.

 Caterpillars photos plus species descriptions at Lepidoptera.pro

  Images from 1837 calendar many images of catapillars
 Larva Locomotion: A Closer Look — video clips showing how monarch larvae walk
 More video clips from nature
 3-D animation Papilio polyxenes larvae walking
 UK moths.Life cycle images

Categories:

 Lepidopterology
 Insect developmental biology
 Larvae

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