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Caterpillars ( ) are the
larval stage A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
of members of the order
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
(the insect order comprising
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
and
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s). As with most common names, the application of the word is arbitrary, since the larvae of
sawflies Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
(suborder Symphyta) are commonly called caterpillars as well. Both lepidopteran and symphytan larvae have
eruciform Eruciform (literally: "caterpillar-shaped") is the entomological term describing a certain class of shapes of insect larvae. Origin and application The word ''eruciform'' literally means "caterpillar-shaped" (from Latin "eruca", a caterpilla ...
body shapes. Caterpillars of most species eat plant material ( often leaves), but not all; some (about 1%) eat insects, and some are even cannibalistic. Some feed on other animal products. For example,
clothes moth Clothes moth or clothing moth is the common name for several species of moth considered to be pests, whose larvae eat animal fibres (hairs), including clothing and other fabrics. These include: * ''Tineola bisselliella'', the common clothes moth ...
s feed on wool, and horn moths feed on the hooves and horns of dead
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...
s. Caterpillars are typically voracious feeders and many of them are among the most serious of
agricultural Agriculture or farming is the practice of cultivating Plant, plants and livestock. Agriculture was the key development in the rise of Sedentism, sedentary human civilization, whereby farming of Domestication, domesticated species created food ...
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
. In fact, many moth
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
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.


Etymology

The origins of the word "caterpillar" date from the early 16th century. They derive from
Middle English Middle English (abbreviated to ME) is a form of the English language that was spoken after the Norman conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English p ...
''catirpel'', ''catirpeller'', probably an alteration of Old North French ''catepelose'': ''cate'', cat (from Latin ''cattus'') + ''pelose'', hairy (from Latin ''pilōsus''). The inchworm, or looper caterpillars from the family
Geometridae The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo'' γεω (derivative form of or "the earth"), and ''met ...
are so named because of the way they move, appearing to measure the earth (the word ''geometrid'' means ''earth-measurer'' in
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
); the primary reason for this unusual locomotion is the elimination of nearly all the
proleg A proleg is a small, fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the order Lepidoptera, though they can also be found on other larval insects such as sawflies and a few other types of in ...
s except the clasper on the terminal segment.


Description

Caterpillars have soft bodies that can grow rapidly between moults. Their size varies between species and
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
s (moults) from as small as up to . Some larvae of the order
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
(ants, bees, and wasps) can appear like the caterpillars of the Lepidoptera. Such larvae are mainly seen in the
sawfly Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
suborder. However while these larvae superficially resemble caterpillars, they can be distinguished by the presence of
proleg A proleg is a small, fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the order Lepidoptera, though they can also be found on other larval insects such as sawflies and a few other types of in ...
s on every abdominal segment, an absence of crochets or hooks on the prolegs (these are present on lepidopteran caterpillars), one pair of prominent
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
on the head capsule, and an absence of the upside-down Y-shaped suture on the front of the head.Scoble, MJ. 1995
''The Lepidoptera: Form, Function and Diversity''
. Oxford Univ. Press.
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 A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
(simple eyes); the sawfly larvae have only two, while caterpillars usually have twelve (six each side of the head). * 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).


Fossils

In 2019, a
geometrid moth The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo'' γεω (derivative form of or "the earth"), and ''metro ...
caterpillar dating back to the
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
epoch In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...
, approximately 44
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
, was found preserved in
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than 1 ...
. It was described under ''
Eogeometer vadens ''Eogeometer'' is a prehistoric genus of Ennomine geometer moths in the tribe Boarmiini. The type and only species is ''Eogeometer vadens'', the specimen of which measured about , and was estimated to be 44 million years old, dating back to Eoce ...
''. Previously, another fossil dating back approximately 125 million years was found in Lebanese
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In Ma ...
.


Defenses

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 ''Gynaephora groenlandica'', the Arctic woolly bear moth, is an erebid moth native to the High Arctic in the Canadian archipelago, Greenland and Wrangel Island in Russia. It is known for its slow rate of development, as its full caterpillar li ...
'' have special basking and aggregation behaviours apart from physiological adaptations to remain in a dormant state.


Appearance

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 warns predators of this. Others may
mimic MIMIC, known in capitalized form only, is a former simulation computer language developed 1964 by H. E. Petersen, F. J. Sansom and L. M. Warshawsky of Systems Engineering Group within the Air Force Materiel Command at the Wright-Patterson AFB in ...
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 defense is the species ''
Nemoria arizonaria ''Nemoria arizonaria'' is a species of moth belonging to the family Geometridae. It was first described (and classified as ''Aplodes arizonaria'') by Augustus Radcliffe Grote in 1883. It is indigenous to Arizona, New Mexico and the Davis Mountains ...
''. 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. 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. Some
Geometridae The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo'' γεω (derivative form of or "the earth"), and ''met ...
cover themselves in plant parts, while bagworms construct and live in a bag covered in sand, pebbles or plant material.


Chemical defenses

More aggressive self-defense measures are taken by some caterpillars. These measures include having spiny bristles or long fine hair-like
seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
e with detachable tips that will irritate by lodging in the skin or mucous membranes. However some birds (such as
cuckoo Cuckoos are birds in the Cuculidae family, the sole taxon in the order Cuculiformes . The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals and anis. The coucals and anis are sometimes separ ...
s) 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 ''Utetheisa ornatrix'', also called the bella moth, ornate moth or rattlebox moth is a moth of the subfamily Arctiinae. It is aposematically colored ranging from pink, red, orange and yellow to white coloration with black markings arranged in va ...
caterpillars utilize
pyrrolizidine alkaloids Pyrrolizidine alkaloids (PAs), sometimes referred to as necine bases, are a group of naturally occurring alkaloids based on the structure of pyrrolizidine. Pyrrolizidine alkaloids are produced by plants as a defense mechanism against insect he ...
that they obtain from their food plants to deter predators. The most aggressive caterpillar defenses are bristles associated with
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
glands. These bristles are called
urticating hair Urticating hairs or urticating bristles are one of the primary defense mechanisms used by numerous plants, almost all New World tarantulas, and various lepidopteran caterpillars. ''Urtica'' is Latin for "nettle" (stinging nettles are in the genu ...
s. A venom which is among the most potent defensive chemicals in any animal is produced by the South American
silk moth The domestic silk moth (''Bombyx mori''), is an insect from the moth family Bombycidae. It is the closest relative of ''Bombyx mandarina'', the wild silk moth. The silkworm is the larva or caterpillar of a silk moth. It is an economically imp ...
genus ''
Lonomia The genus ''Lonomia'' is a moderate-sized group of fairly cryptic saturniid moths from South America, famous not for the adults, but for their highly venomous caterpillars, which are responsible for a few deaths each year, especially in southern ...
''. Its venom is an
anticoagulant Anticoagulants, commonly known as blood thinners, are chemical substances that prevent or reduce coagulation of blood, prolonging the clotting time. Some of them occur naturally in blood-eating animals such as leeches and mosquitoes, where the ...
powerful enough to cause a human to
hemorrhage Bleeding, hemorrhage, haemorrhage or blood loss, is blood escaping from the circulatory system from damaged blood vessels. Bleeding can occur internally, or externally either through a natural opening such as the mouth, nose, ear, urethra, vag ...
to death (See
Lonomia The genus ''Lonomia'' is a moderate-sized group of fairly cryptic saturniid moths from South America, famous not for the adults, but for their highly venomous caterpillars, which are responsible for a few deaths each year, especially in southern ...
sis). This chemical is being investigated for potential medical applications. Most urticating hairs range in effect from mild irritation to
dermatitis Dermatitis is inflammation of the skin, typically characterized by itchiness, redness and a rash. In cases of short duration, there may be small blisters, while in long-term cases the skin may become thickened. The area of skin involved can v ...
. Example:
brown-tail moth The brown-tail moth (''Euproctis chrysorrhoea'') is a moth of the family Erebidae. It is native to Europe, neighboring countries in Asia, and the north coast of Africa. Descriptions of outbreaks, i.e., large population increases of several year ...
. 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 Sequestration or sequester may refer to: Law and government * Sequestration (law), the seizure of property for creditors or the state. See also Committee for Compounding with Delinquents * Jury sequestration, the isolation of a jury * Bankruptcy, ...
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 The cinnabar moth (''Tyria jacobaeae'') is a brightly coloured arctiid moth found as a native species in Europe and western and central Asia then east across the Palearctic to Siberia to China. It has been introduced into New Zealand, Australia a ...
(''Tyria jacobaeae'') and monarch (''
Danaus plexippus The monarch butterfly or simply monarch (''Danaus plexippus'') is a milkweed butterfly (subfamily Danainae) in the family Nymphalidae. Other common names, depending on region, include milkweed, common tiger, wanderer, and black-veined brown. I ...
'') caterpillars, usually advertise themselves with the danger colors of red, yellow and black, often in bright stripes (see
aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
). 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 The osmeterium is a defensive organ found in all Papilionidae, papilionid larvae, in all stages. The organ is situated in the prothorax, prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy f ...
.


Defensive behaviors

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 Mining is the extraction of valuable minerals or other geological materials from the Earth, usually from an ore body, lode, vein, seam, reef, or placer deposit. The exploitation of these deposits for raw material is based on the economic via ...
between the leaf surfaces. Some caterpillars, like early instars of the
tomato hornworm ''Manduca quinquemaculata'', the five-spotted hawkmoth, is a brown and gray hawk moth of the family Sphingidae. The caterpillar, often referred to as the tomato hornworm, can be a major pest in gardens; they get their name from a dark projectio ...
and
tobacco hornworm ''Manduca sexta'' is a moth of the family Sphingidae present through much of the Americas. The species was first described by Carl Linnaeus in his 1763 '' Centuria Insectorum''. Commonly known as the Carolina sphinx moth and the tobacco hawk m ...
, have long "whip-like" organs attached to the ends of their body. The caterpillar wiggles these organs to frighten away flies and predatory wasps. 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 ''Amorpha juglandis'', the walnut sphinx, is the only species in the monotypic moth genus ''Amorpha'', which is in the family Sphingidae, erected by Jacob Hübner in 1809. The species was first described by James Edward Smith in 1797. Distribu ...
'') even makes high pitched whistles that can scare away birds.


Social behaviors and relationships with other insects

Some caterpillars obtain protection by associating themselves with
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s. 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. Some caterpillars are
gregarious Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
; large aggregations are believed to help in reducing the levels of parasitization and predation. Clusters amplify the signal of aposematic coloration, and individuals may participate in group regurgitation or displays.
Pine processionary The pine processionary (''Thaumetopoea pityocampa'') is a moth of the subfamily Thaumetopoeinae in the family Notodontidae, known for the irritating hairs of its caterpillars, their processions, and the economic damage they cause in coniferous ...
(''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. Forest tent caterpillars cluster during periods of cold weather.


Predators

Caterpillars are eaten by many animals. The
European pied flycatcher The European pied flycatcher (''Ficedula hypoleuca'') is a small passerine bird in the Old World flycatcher family. One of the four species of Western Palearctic black-and-white flycatchers, it hybridizes to a limited extent with the collared ...
is one species that preys upon caterpillars. The flycatcher typically finds caterpillars among oak foliage.
Paper wasp Paper wasps are Vespidae, vespid wasps that gather fibers from dead wood and plant stems, which they mix with saliva, and use to construct nests made of gray or brown papery material. Some types of paper wasps are also sometimes called umbrell ...
s, including those in the genus''
Polistes Wasps of the cosmopolitan genus ''Polistes'' (the only genus in the tribe Polistini) are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. Walter Ebeling coined the vernacular name "umbrella ...
'' and '' Polybia'' catch caterpillars to feed their young and themselves.


Behavior

Caterpillars have been called "eating machines", and eat leaves voraciously. Most species shed their
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...
four or five times as their bodies grow, and they eventually enter a
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
l stage before becoming adults. Caterpillars grow very quickly; for instance, a
tobacco hornworm ''Manduca sexta'' is a moth of the family Sphingidae present through much of the Americas. The species was first described by Carl Linnaeus in his 1763 '' Centuria Insectorum''. Commonly known as the Carolina sphinx moth and the tobacco hawk m ...
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 Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
. Most caterpillars are solely
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
. Many are restricted to feeding on one species of plant, while others are polyphagous. Some, including the
clothes moth Clothes moth or clothing moth is the common name for several species of moth considered to be pests, whose larvae eat animal fibres (hairs), including clothing and other fabrics. These include: * ''Tineola bisselliella'', the common clothes moth ...
, feed on
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
. Some are predatory, and may prey on other species of caterpillars (e.g. Hawaiian ''
Eupithecia ''Eupithecia'' is a large genus of moths of the family Geometridae. There are hundreds of described species, found in all parts of the world (with 45 in the British Isles alone), and new species are discovered on a regular basis. ''Eupithecia'' ...
''). Others feed on eggs of other insects, aphids, scale insects, or ant larvae. A few are parasitic on cicadas or leaf hoppers (
Epipyropidae The Epipyropidae comprise a small family of moths. This family and the closely related Cyclotornidae are unique among the Lepidoptera in that the larvae are ectoparasites, the hosts typically being fulgoroid planthoppers, thus the common name ...
). Some Hawaiian caterpillars ('' Hyposmocoma molluscivora'') use silk traps to capture snails. Many caterpillars are nocturnal. For example, the "cutworms" (of the family
Noctuidae The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, along with the other f ...
) hide at the base of plants during the day and only feed at night. Others, such as gypsy moth (
Lymantria dispar ''Lymantria dispar'', also known as the gypsy moth or the spongy moth, is a species of moth in the family Erebidae. ''Lymantria dispar'' is subdivided into several subspecies, with subspecies such as ''L. d. dispar'' and ''L. d. japonica'' be ...
) larvae, change their activity patterns depending on density and larval stage, with more diurnal feeding in early instars and high densities.


Economic effects

Caterpillars cause much damage, mainly by eating leaves. The propensity for damage is enhanced by
monocultural Monoculturalism is the policy or process of supporting, advocating, or allowing the expression of the culture of a single social or ethnic group. It generally stems from beliefs within the dominant group that their cultural practices are superior ...
farming practices, especially where the caterpillar is specifically adapted to the host plant under cultivation. The
cotton bollworm Cotton bollworm is a problem in growing cotton. "A major pest in hot countries of irrigated crops. Enters into a summer diapause when irrigated crops are not present and the soil and air temperatures are high. When the end of the dry season comes, t ...
causes enormous losses. Other species eat food crops. Caterpillars have been the target of
pest control Pest control is the regulation or management of a species defined as a pest; any animal, plant or fungus that impacts adversely on human activities or environment. The human response depends on the importance of the damage done and will range ...
through the use of
pesticide Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lampri ...
s,
biological control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
and
agronomic Agricultural economics is an applied field of economics concerned with the application of economic theory in optimizing the production and distribution of food and fiber products. Agricultural economics began as a branch of economics that spe ...
practices. Many species have become resistant to
pesticide Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lampri ...
s. 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. 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 Plant defense against herbivory or host-plant resistance (HPR) describes a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants can sense being touched, and they can ...
(HPR) through plant breeding is another approach used in reducing the impact of caterpillars on crop plants. Some caterpillars are used in industry. The
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
industry is based on the
silkworm The domestic silk moth (''Bombyx mori''), is an insect from the moth family Bombycidae. It is the closest relative of ''Bombyx mandarina'', the wild silk moth. The silkworm is the larva or caterpillar of a silk moth. It is an economically imp ...
caterpillar.


Human health

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
urticaria Hives, also known as urticaria, is a kind of skin rash with red, raised, itchy bumps. Hives may burn or sting. The patches of rash may appear on different body parts, with variable duration from minutes to days, and does not leave any long-lasti ...
l dermatitis and
atopic Atopy is the tendency to produce an exaggerated immunoglobulin E (IgE) immune response to otherwise harmless substances in the environment. Allergic diseases are clinical manifestations of such inappropriate, atopic responses. Atopy may have a ...
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, cou ...
to
osteochondritis Osteochondritis is a painful type of osteochondrosis where the cartilage or bone in a joint is inflamed. It often refers to osteochondritis dissecans (OCD). The term ''dissecans'' refers to the "creation of a flap of cartilage that further disse ...
, consumption
coagulopathy Coagulopathy (also called a bleeding disorder) is a condition in which the blood's ability to coagulate (form clots) is impaired. This condition can cause a tendency toward prolonged or excessive bleeding (bleeding diathesis), which may occur spo ...
,
kidney failure Kidney failure, also known as end-stage kidney disease, is a medical condition in which the kidneys can no longer adequately filter waste products from the blood, functioning at less than 15% of normal levels. Kidney failure is classified as eit ...
, and
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
bleeding Bleeding, hemorrhage, haemorrhage or blood loss, is blood escaping from the circulatory system from damaged blood vessels. Bleeding can occur internally, or externally either through a natural opening such as the mouth, nose, ear, urethra, vag ...
. Skin rashes are the most common, but there have been fatalities. ''
Lonomia The genus ''Lonomia'' is a moderate-sized group of fairly cryptic saturniid moths from South America, famous not for the adults, but for their highly venomous caterpillars, which are responsible for a few deaths each year, especially in southern ...
'' 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. Caterpillar hair has also been known to cause kerato-
conjunctivitis Conjunctivitis, also known as pink eye, is inflammation of the conjunctiva, outermost layer of the white part of the eye and the inner surface of the eyelid. It makes the eye appear pink or reddish. Pain, burning, scratchiness, or itchiness may ...
. The sharp barbs on the end of caterpillar hairs can get lodged in soft tissues and
mucous membrane A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It is ...
s 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 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. Caterpillars are a food source in some cultures. For example, in South Africa
mopane worms ''Gonimbrasia belina'' is a species of emperor moth which is native to the warmer parts of southern Africa. Its large edible caterpillar, known as the mopane worm, madora, amacimbi or masontja, feeds primarily but not exclusively on mopane tree ...
are eaten by the
bushmen The San peoples (also Saan), or Bushmen, are members of various Khoe, Tuu, or Kxʼa-speaking indigenous hunter-gatherer cultures that are the first cultures of Southern Africa, and whose territories span Botswana, Namibia, Angola, Zambia, ...
, and in China
silkworms The domestic silk moth (''Bombyx mori''), is an insect from the moth family Bombycidae. It is the closest relative of ''Bombyx mandarina'', the wild silk moth. The silkworm is the larva or caterpillar of a silk moth. It is an economically im ...
are considered a delicacy.


In popular culture

In the
Old Testament The Old Testament (often abbreviated OT) is the first division of the Christian biblical canon, which is based primarily upon the 24 books of the Hebrew Bible or Tanakh, a collection of ancient religious Hebrew writings by the Israelites. The ...
of the
Bible The Bible (from Koine Greek , , 'the books') is a collection of religious texts or scriptures that are held to be sacred in Christianity, Judaism, Samaritanism, and many other religions. The Bible is an anthologya compilation of texts of a ...
caterpillars are feared as pest that devour crops. They are part of the "pestilence, blasting, mildew, locust" because of their association with the
locust Locusts (derived from the Vulgar Latin ''locusta'', meaning grasshopper) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumstanc ...
, thus they are one of the plagues of Egypt.
Jeremiah Jeremiah, Modern:   , Tiberian: ; el, Ἰερεμίας, Ieremíās; meaning " Yah shall raise" (c. 650 – c. 570 BC), also called Jeremias or the "weeping prophet", was one of the major prophets of the Hebrew Bible. According to Jewish ...
names them as one of the inhabitants of
Babylon ''Bābili(m)'' * sux, 𒆍𒀭𒊏𒆠 * arc, 𐡁𐡁𐡋 ''Bāḇel'' * syc, ܒܒܠ ''Bāḇel'' * grc-gre, Βαβυλών ''Babylṓn'' * he, בָּבֶל ''Bāvel'' * peo, 𐎲𐎠𐎲𐎡𐎽𐎢 ''Bābiru'' * elx, 𒀸𒁀𒉿𒇷 ''Babi ...
. 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 William Shakespeare ( 26 April 1564 – 23 April 1616) was an English playwright, poet and actor. He is widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's nation ...
'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 William Blake (28 November 1757 – 12 August 1827) was an English poet, painter, and printmaker. Largely unrecognised during his life, Blake is now considered a seminal figure in the history of the poetry and visual art of the Romantic Age. ...
referenced this popular image in
The Marriage of Heaven and Hell ''The Marriage of Heaven and Hell'' is a book by the English poet and printmaker William Blake. It is a series of texts written in imitation of biblical prophecy but expressing Blake's own intensely personal Romantic and revolutionary beliefs ...
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". The role of caterpillars in the life stages of butterflies was badly understood. In 1679
Maria Sibylla Merian Maria Sibylla Merian (2 April 164713 January 1717) was a German naturalist and scientific illustrator. She was one of the earliest European naturalists to observe insects directly. Merian was a descendant of the Frankfurt branch of the Swiss M ...
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 A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
. An earlier popular publication on moths and butterflies, and their caterpillars, by
Jan Goedart Johannes Goedaert (also spelled Goetaart, Goedhart, Goedaard or Jean Goedart in French) (19 March 1617 (baptized) – 15 January 1668 (buried)) was a Dutch Natural history, natur ...
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 Francesco Redi (18 February 1626 – 1 March 1697) was an Italian physician, naturalist, biologist, and poet. He is referred to as the "founder of experimental biology", and as the "father of modern parasitology". He was the first person to cha ...
,
Marcello Malpighi Marcello Malpighi (10 March 1628 – 30 November 1694) was an Italian biologist and physician, who is referred to as the "Founder of microscopical anatomy, histology & Father of physiology and embryology". Malpighi's name is borne by several phy ...
and
Jan Swammerdam Jan Swammerdam (February 12, 1637 – February 17, 1680) was a Dutch biologist and microscopist. His work on insects demonstrated that the various phases during the life of an insect—egg, larva, pupa, and adult—are different forms of the ...
. Butterflies were regarded as symbol for the human soul since ancient time, and also in the Christian tradition. 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." 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. As a militant
Cartesian Cartesian means of or relating to the French philosopher René Descartes—from his Latinized name ''Cartesius''. It may refer to: Mathematics *Cartesian closed category, a closed category in category theory *Cartesian coordinate system, modern ...
, 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." 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". Famously, in
Lewis Carroll Charles Lutwidge Dodgson (; 27 January 1832 – 14 January 1898), better known by his pen name Lewis Carroll, was an English author, poet and mathematician. His most notable works are ''Alice's Adventures in Wonderland'' (1865) and its sequel ...
's
Alice's Adventures in Wonderland ''Alice's Adventures in Wonderland'' (commonly ''Alice in Wonderland'') is an 1865 English novel by Lewis Carroll. It details the story of a young girl named Alice (Alice's Adventures in Wonderland), Alice who falls through a rabbit hole into a ...
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. 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 René Descartes ( or ; ; Latinized: Renatus Cartesius; 31 March 1596 – 11 February 1650) was a French philosopher, scientist, and mathematician, widely considered a seminal figure in the emergence of modern philosophy and science. Mathem ...
, 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 Perception () is the organization, identification, and interpretation of sensory information in order to represent and understand the presented information or environment. All perception involves signals that go through the nervous system ...
of the
truth Truth is the property of being in accord with fact or reality.Merriam-Webster's Online Dictionarytruth 2005 In everyday language, truth is typically ascribed to things that aim to represent reality or otherwise correspond to it, such as beliefs ...
. More recent symbolic references to caterpillars in popular media include the ''Mad Men'' season 3 episode "The Fog", in which
Betty Draper Elizabeth "Betty" Hofstadt Francis (formerly Draper) is a fictional character on AMC's television series '' Mad Men'', wife of Don Draper (Jon Hamm) and mother of his three children. Blonde and beautiful but emotionally distant and immature, she s ...
has a drug-induced dream, while in labor, that she captures a caterpillar and holds it firmly in her hand.What's Alan Watching?: ''Mad Men'', "The Fog" In ''The Sopranos'' season 5 episode " The Test Dream",
Tony Soprano Anthony John Soprano is a fictional character and the antihero protagonist in the HBO television drama series ''The Sopranos'' (1999–2007), portrayed by James Gandolfini. Soprano is a member of the American Mafia, Italian-American Mafia and, ...
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. File:CH Caterpillar.jpg, Caterpillar of the
spurge hawk-moth ''Hyles euphorbiae'', the spurge hawk-moth, is a European moth of the family Sphingidae. This hawk moth is used as an agent of biological pest control against the noxious weed leafy spurge (''Euphorbia virgata''), but usually only in conjunction ...
, near
Binn Binn (Walser German: ''Bìi'') is a municipality in the district of Goms in the canton of Valais in Switzerland. The Binn Valley is known for its rich mineral deposits, some types of which are unique to the area. The notable Lengenbach Quar ...
,
Valais Valais ( , , ; frp, Valês; german: Wallis ), more formally the Canton of Valais,; german: Kanton Wallis; in other official Swiss languages outside Valais: it, (Canton) Vallese ; rm, (Chantun) Vallais. is one of the cantons of Switzerland, 26 ...
, Switzerland at c. 2 km altitude. File:Phalaenoides_glyciae_caterpillar.jpg, '' Phalaenoides glycinae'' (Australian grapevine moth) feeding on a
grapevine ''Vitis'' (grapevine) is a genus of 79 accepted species of vining plants in the flowering plant family Vitaceae. The genus is made up of species predominantly from the Northern Hemisphere. It is economically important as the source of grapes, b ...
. File:Sphinx ligustri, the privet hawk moth caterpillar-IMG 4927.jpg, ''
Sphinx ligustri ''Sphinx ligustri'', the privet hawk moth, is a moth found in most of the Palearctic realm. The species was first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''. Description It has a wingspan (generally deflexed ...
'', the privet hawk moth caterpillar File:Caterpillar at 5th stage.jpg, Caterpillar of the
emperor gum moth ''Opodiphthera eucalypti'', the emperor gum moth, is a species of moth in the family Saturniidae native to Australia. This species was formerly placed in the genus ''Antheraea''. Distribution The emperor gum moth may inhabit all states of Aust ...
. File:Poplar hawk-moth.jpg, A poplar hawk-moth caterpillar (a common species of caterpillar in the UK). File:Fuzzy-Orange-Caterpillar.jpg, Fuzzy orange caterpillar File:Lycaenid ant sec.jpg, Ant tending a lycaenid caterpillar. File:Caterpillar on leaf (1) 01.jpg, Caterpillar feeding on leaf File:Snodgrass Schizura concinna.jpg, Life cycle of the red-humped caterpillar ('' Schizura concinna ''). File:Forest-tent-caterpillar-malacosoma-disstria.jpg,
Forest tent caterpillar The forest tent caterpillar moth (''Malacosoma disstria'') is a moth found throughout North America, especially in the eastern regions. Unlike related tent caterpillar species, the larvae of forest tent caterpillars do not make tents, but rathe ...
(Malacosoma disstria) File:Eighteyescatterpilar.jpg, Camouflage: apparently with eight eyes, only two of them are real. Photo in a
eucalyptus ''Eucalyptus'' () is a genus of over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as euca ...
tree,
São Paulo São Paulo (, ; Portuguese for 'Saint Paul') is the most populous city in Brazil, and is the capital of the state of São Paulo, the most populous and wealthiest Brazilian state, located in the country's Southeast Region. Listed by the GaWC a ...
, Brazil File:AntheraeaPolyphemusCaterpillar.jpg, Caterpillar of the
Polyphemus moth ''Antheraea polyphemus'', the Polyphemus moth, is a North American member of the family Saturniidae, the giant silk moths. It is a tan-colored moth, with an average wingspan of 15 cm (6 in). The most notable feature of the moth is its ...
(''Antheraea polyphemus''),
Virginia Virginia, officially the Commonwealth of Virginia, is a state in the Mid-Atlantic and Southeastern regions of the United States, between the Atlantic Coast and the Appalachian Mountains. The geography and climate of the Commonwealth ar ...
, United States File:Baby Caterpillars crop2.jpg, Caterpillars on an apple tree in
Victoria, British Columbia Victoria is the capital city of the Canadian province of British Columbia, on the southern tip of Vancouver Island off Canada's Pacific coast. The city has a population of 91,867, and the Greater Victoria area has a population of 397,237. Th ...
, Canada File:Caterpiller on Leaf.jpg, Caterpillar on a leaf File:Automeris illustris Automeris (caterpillar) - Laslovarga.JPG, Caterpillar of Belize File:Dryas julia caterpillar.JPG, ''
Dryas iulia ''Dryas iulia'' (often incorrectly spelled ''julia''),Lamas, G. (editor) (2004). Atlas of Neotropical Lepidoptera. Checklist: Part 4A. Hesperioidea – Papilionoidea. commonly called the Julia butterfly, Julia heliconian, the flame, or flambeau ...
'' File:Hebomoea glaucippe cat sec.jpg, Caterpillar of great orange tip resembling the common green vine snake ('' Ahaetulla nasuta'') File:Cabbage looper in cocoon.jpg, Prepupa of
cabbage looper The cabbage looper (''Trichoplusia ni'') is a medium-sized moth in the family Noctuidae, a family commonly referred to as owlet moths. Its common name comes from its preferred host plants and distinctive crawling behavior. Cruciferous vegetable ...
in its cocoon File:Caterpillar_(locomotion)_04.jpg, Locomotion of a small
Geometrid The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo'' γεω (derivative form of or "the earth"), and ''metro ...
caterpillar.


See also

* Edible caterpillars *
Larval food plants of Lepidoptera Caterpillars (larvae) of Lepidoptera species (i.e. of butterflies and moths) are mostly (though not exclusively) herbivores, often oligophagous, i.e. feeding on a narrow variety of plant species (mostly on their leaves, but sometimes on fruit or o ...
*
Lepidopterism __NOTOC__ Lepidopterism is an irritant contact dermatitis caused by irritating caterpillar or moth hairs coming into contact with the skin or mucosa. When referring to the cause, moth dermatitis and caterpillar dermatitis are commonly used; Caripi ...
- caterpillar dermatitis *
List of pests and diseases of roses Roses (''Rosa'' species) are susceptible to a number of pests, diseases and disorders. Many of the problems affecting roses are seasonal and climatic.Ross, D.,''Rose-growing for Pleasure'', Lothian Publishing, Melbourne, 1985, pp. 27 ...
*
Sericulture Sericulture, or silk farming, is the cultivation of silkworms to produce silk. Although there are several commercial species of silkworms, ''Bombyx mori'' (the caterpillar of the domestic silkmoth) is the most widely used and intensively studie ...


References


External links


Photos of caterpillars
at ''Insecta.pro''
Images from 1837 calendar
many images of catapillars

* ttps://web.archive.org/web/20130910094053/http://www.learner.org/jnorth/tm/monarch/jr/KidsJourneyNorth.html More video clips from nature
3-D animation ''Papilio polyxenes'' larvae walking


{{Authority control Lepidopterology Insect developmental biology Larvae Biological anti-agriculture weapons