Butterflies

The title of this article is ambiguous. Further meanings are listed under butterflies (disambiguation) .

Butterflies
Colored engraving with forest-damaging butterflies by Tieffenbach from Ratzeburg's "Die Waldverderbnis" (1866)
Systematics
Sub-stem : Trachea (Tracheata) Superclass : Six-footed (Hexapoda) Class : Insects (Insecta) Subclass : Flying insects (Pterygota) Superordinate : New winged wing (Neoptera) Order : Butterflies
Scientific name
Lepidoptera
Linnaeus , 1758
Submissions
Zeugloptera (Protolepidoptera) Aglossata Heterobathmiina Glossata complete systematics of butterflies

The butterflies (Lepidoptera, from ancient Greek λέπος lépos "scales" and πτερόν pterón "wings") form with almost 160,000 described species , (as of 2011) about 130 families and 46 superfamilies after the beetles (Coleoptera) the most species-rich insect order. Around 700 species are discovered every year. Butterflies are common on all continents except Antarctica . In Central Europe they are represented with about 4,000 species; For the whole of Europe, Ole Karsholt's catalog lists over 10,600 species. In Germany there are around 3,700 species.

origin of the name

Cethosia cyane

The German name "butterfly", first documented in 1501, comes from the Slavic-derived East Central German word Schmetten (that is, Schmand , cream), to which some species are often attracted. In superstition , butterflies were even considered to be the embodiment of witches who were after the cream, which is also indicated by earlier landscape names for butterflies such as milk thief, whey stealer or the like. The English name butterfly points in the same direction and corresponds to the regionally common names Buttervogel, Bottervagel, Botterlicker, as the animals were attracted by the butter whipping . Various other names also existed locally; In addition to the above-mentioned, for example, Westphalian Schmandlecker (from Schmand), Bavarian Müllermaler, Hessian Lattichvogel (from Lattich ), Silesian, Transylvanian and in parts of Switzerland Sommervogel (equivalent to the Danish sommerfugl ), in other parts of Switzerland also (P) Fifalter .

The word butterfly only became generally accepted in the second half of the 18th century. Until then, this order of insects was still called "day birds" (for butterflies) or "night birds" (for moths) according to Rösel von Rosenhof (1749). The term butterfly has nothing to do with folding (the wing) or with fluttering . The Germanic word - Middle High German  vīvalter , Old High German  fīfalt (a) ra , Old English fīff (e) alde , Old Norse fífrildi  - is probably related to the Latin pāpilio , from which Italian farfalla or French papillon are derived. In addition, the Indo-European derivation is unclear.

The scientific name Lepidoptera (emphasis on the o) means "flake wing". It is one of Linné embossed composition of ancient Greek λεπίς Lepiš "scale" (genitive lepídos ) and πτερόν pteron "wings" (plural Ptera ). The ancient Greek word for butterfly was ψυχή ( psuche or psyche ), breath, breath, soul, because the animals were regarded as the embodiment of the human soul. This expression was mainly used for moths and was only found for butterflies in the Hellenistic period. A seldom used expression was φάλαινα phalaina (later spelling φάλλαινα phallaina , adopted as phalaena in Latin).

Development cycle

During their life, butterflies go through a cycle with unusual changes in shape: Eggs develop into flightless " caterpillars ", which move by changing their shape and grow considerably with intensive food intake. In doing so, they usually change their skin several times for a larger one ("moult"). At the end of the caterpillar stage, with the formation of a firmer shell, they change into a state that externally appears as a state of rest, the so-called “ pupa ”. They change significantly in the pupae: They develop wings and transform into the form capable of flying, the so-called "butterfly", also known as the imago. The moths take in liquid food through proboscis and one female and one male moths mate with fertilization of the female's eggs. The females then lay the fertilized eggs from which caterpillars hatch.

Adults

The body structure of the adults in butterflies corresponds to the basic plan of practically all other insects: They have an exoskeleton made of chitin and proteins, several plates called sclerites are arranged in segmental rings and flexibly connected by joint membranes. The legs and antennae are also made of such rings. The body is divided into head ( caput ), chest ( thorax ) and abdomen ( abdomen ). In the head of the butterflies the antennae, the eyes and in most species the mouthparts with the proboscis are very conspicuous, on the thorax the mostly large and very delicate wings are suspended, which dominate the entire shape of the butterfly.

Butterflies reach a body length (measured without wings) of 1.5 to 100 millimeters. The largest butterfly is the owl butterfly Thysania agrippina from South America . These moths reach wingspans of 25 to 30 centimeters. The Queen Alexandra bird butterfly ( Ornithoptera alexandrae ) is the largest butterfly with wingspan of 20 to 28 centimeters. The smallest moths are members of the crested-forehead moths (Tischeriidae), which contain species with a wingspan of only 1.5 to 2 millimeters. As a butterfly with the largest wing surface is regarded Atlas Moth ( Attacus atlas ) from Southeast Asia.

sensor

Different antennae types of butterflies

The structure of the antennae can be very different and is often a characteristic of the respective butterfly family. There are thread-shaped, clubbed (which are thread-shaped and have a thickening at the end), sawn (which have protrusions on one side) and combed antennae (which carry these on both sides), along with all sorts of transitions. The antennae are often built differently in the sexes, in these cases they are much more pronounced in the males. With their antennae, the butterflies can smell, some also touch, taste and perceive temperatures . The stimulus is picked up by small hairs that are distributed on the feelers. Sawn or combed feelers greatly increase the surface area, which considerably improves the sense of smell. This enables males to perceive the pheromones released by females ready to mate from a great distance. This is especially important in the case of butterflies that live in very dispersed populations or in forests and therefore do not collide by accident. The females sense the right host plants with their antennae.

Mouthparts

SEM image of a butterfly's head with a curled trunk

The mouthparts of butterflies are very specialized and modified compared to other insects. Your mandibles (upper jaw) are severely stunted. Only in the family of the ancient moths (Micropterigidae) are these still used as biting tools. In most butterflies, the lower jaws ( maxillae ) form two flexible half-tubes that are connected by folded seams. This creates the suction tube between the two tubes, with which the moths can suck up their food. This can only be liquid. Almost all butterflies feed on flower nectar , plant juices and other nutrient-rich liquids. At rest is proboscis rolled under the head. In addition to the changed maxillae, the butterflies have maxillary palps that have receded, as well as lip palpals ( labial palps ), which are elongated and large in some species (e.g. in the subfamily Libytheinae of the noble butterfly ). The palps have organs of touch and smell.

The length of the trunk is very different depending on the species. The swarmers (Sphingidae) have the longest proboscis. In a hawkmoth species Amphimoea walkeri living in the subtropics , the trunk length is 280 millimeters; up to now no other butterfly species has been discovered that exceeds this length. This allows them to penetrate the particularly narrow flower necks of orchids . The trunk of the skull hawk ( Acherontia atropos ), on the other hand, is very short but very strong. With it, the animals already capped can honeycomb pierce and suck, in addition they can produce with it beeps.

In some butterfly species such as the peacock moth (Saturniidae) or the mother hen (Lasiocampidae), the proboscis is completely regressed. Your only mouth parts are the unpaired lower lip ( labium ) with the labial palps. But they cannot take in any food with it. These animals die soon after mating. Their real life takes place in the caterpillar stage.

The primeval moths (Micropterigidae) do not have a proboscis, but they can chew with mandibles and feed on pollen .

On the lower lip of the caterpillar, on a cone, there is the opening of the spinneret, in which silk is produced in the form of a liquid that solidifies in the air after it escapes.

eyes

Eyes and proboscis of the little cabbage white butterfly

As with other insects, the eyes are designed as compound eyes . These consist of up to 6,000 small individual eyes ( ommatidia ). In addition to these, many butterfly species also have a pair of single eyes ( ocelles ) with which they control their day-night rhythm. In contrast to butterflies, moths, which are exposed to large differences in brightness, have pigment cells in their eyes with which they can regulate the intensity of the incident light. They are nearsighted because they cannot accommodate due to the compound eyes, i.e. they cannot adapt their visual acuity to the object distance. In addition, they only see “pixelated” through the facets. However, they have a large field of vision and respond well to movements. The moths also have a different color sensitivity than humans. They do not recognize red colors, but they are sensitive in the ultraviolet range . The moths in particular are attracted to UV lamps. Butterflies can see about 200 m with their eyes and move towards a target within this range.

Chest section (thorax)

The thorax consists of three ring-shaped parts ( prothorax , mesothorax and metathorax ), which encompass all of the animals' locomotor organs. A pair of legs is placed on each segment. In many butterflies, the first pair of legs has receded and their cleaning spurs are only used for cleaning. The legs consist of the hip ( coxa ), thigh ring ( trochanter ), thigh ( femur ), splint ( tibia ) and foot ( tarsus ). The foot again consists of five links, the last link has claws to hold on to. With some butterflies there are sensory organs on the tarsi with which they can taste. The legs of the olive borer owl ( Zanclognatha tarsipennalis ) have thorns and large tufts of hair, which are thought to be used to distribute scents. The two pairs of wings sit on the two posterior segments of the thorax.

Butterflies are able to hear. Your ears ( tympanic organs ) are located either in the back of the thorax or on the abdomen in a pit covered by a thin membrane. This membrane works in a similar way to the human eardrum . Moths, such as owl butterflies (Noctuidae) or bear moths (Arctiidae), are evolutionarily well developed , are also sensitive in the ultrasound range, since their main enemies, the bats , use these signals for localization. If a locating tone is received, the moths drop in flight to avoid being captured. Bear moths can even make ultrasonic sounds. Since many of them are poisonous, bats associate these noises with inedible and let go of the butterflies.

Wing scales under the microscope

Black moth ( Biston betularia ); dark shape

Wing of Heliconius ismenius

wing

With a few exceptions, the wings are the actual locomotor apparatus of the moths. The front and rear wings are suspended individually, but are sometimes coupled with one another by special mechanisms in flight. Most butterflies lack such a connection. The veins run over the wings, between an upper and a lower membrane . After hatching, when the wings are still limp and immobile, these are filled with a blood fluid. The wings can then dry out and these veins lose their function. The wings are covered with scales on the top and bottom . In addition, the entire body of most butterflies is scaled. These scales are flattened, species-specific hairs that lie on the wings like roof tiles and thus cover the wing veins.

Since the wings are particularly important for identifying butterflies, and there are often only slight differences between different species, the wings are divided into regions and the veins of the wings and the cells formed from them are numbered. The wings of a butterfly are described as an example: The regions run from the base of the wing to the tip, with each of the fore and hind wings being divided into four regions. The veins are numbered from one to twelve on the forewings, starting from the rear with one, which runs parallel to the inner edge. There are only nine continuous veins on the hind wings, but sometimes there is also a tenth. The areas that are bounded by the wing veins are called cells or central or discoidal cells. You can find more information about the wings in the article Wings (butterfly) .

Butterflies are often brightly colored. The coloring is created on the one hand by pigments , on the other hand by special surface structures (also called structural colors ) that cause light refraction effects. The individual scales are always only one color. Their shape, however, varies greatly. The most common form is the shield-shaped with three to five points and a narrow stem anchored in a recess at the end. Others are lance-shaped or circular. Some scales, which are usually in fields next to each other and are provided with tufts of hair, allow odors to be emitted through pores. These scent scales ( androconia ) make it easier for sexual partners to find each other. The scales are not necessary for flying. In the case of the glass winged birds (Sesiidae), large areas of the wings are initially still loosely scaled, but on the first flight they become transparent and crystal clear due to the loss of the scales.

In the females of some species, and very rarely also in the males, the wings are completely regressed. You can only move around continuously. This is the case , for example, with the real bag carriers (Psychidae). The spanners (Geometridae) also have species with shortened wings.

In addition to the "normal" colored ones, there may also be specimens of the same species that are colored completely differently. They are called aberrations or morphs . Mostly these are darker to completely black specimens. In earlier times, separate names were coined for the different color variants of a species, but according to today's view, these names have no nomenclature relevance.

Abdomen (abdomen)

The abdomen consists of ten uniformly shaped segments that are covered with species-specific colored scales. The abdomen contains the life-sustaining organs, a tubular heart , the nervous system , the digestive tract and the sexual organs, as well as various glands that primarily produce fragrances. The sex organs are built specifically for the species and are therefore very important for their determination. On the outside, the male wears a clipper to hold the female in place during mating. The female is equipped with an ovipositor . In some species, the females have tufts of hair at the end of the abdomen, which can be stripped off over the egg clutches for camouflage. Other species have thorns that release poison when touched.

internal structure

The tubular heart pumps the blood ( hemolymph ), which flows around the organs, around the body in a simple circuit . The blood transports nutrients in the body, but no oxygen or carbon dioxide . The gas exchange takes place via tracheas , which with their branched pipe system supply all organs with oxygen, which is pumped into the body through lateral openings (spiracles). The maximum transport path is limited in this respiratory system, which is also the reason why butterflies and insects are generally limited in their growth in size.

The nervous system is on the underside below the intestine . It consists of two parallel nerve cords that are connected by ganglia like a rope ladder. At the anterior end of the abdomen, the cords run around the intestine and connect with the head ganglia of the brain , consisting of the lower and upper pharyngeal ganglions . These two nerve segments are independent of each other. This means that the body can still work even though the brain is already dead. The digestive system begins with a muscular throat ( pharynx ) that pumps food from the mouth into the gullet ( esophagus ). It is designed as a tube and leads into the food store. It is connected to the midgut, where nutrients are absorbed into the blood. This is followed by the rectum. There, as well as the two tubular kidneys ( Malpighian vessels ), metabolic products are absorbed from the organs and excreted through the anus . As already mentioned, there are not only butterflies that do not eat and whose digestive system is useless, but also those that have no digestive system at all, as is the case with some peacock spiders .

The internal sexual organs in males consist of two testes lying above the intestine , which in many species have grown together. They are connected by narrow tubes ( vasa deferentia ) to the ejaculatory duct , which leads to the aedeagus . It acts as a penis and transfers the sperm into the female. Most of the time, the sperm are stored in a bladder with a harder shell ( spermatophore ) and transferred. The females have two ovaries , each connected by four tubes, in which the eggs are formed.

Caterpillars

Poplar Hawk caterpillar

Abdominal legs of the swallowtail caterpillar

Head detail of the agate owl caterpillar

→ Main article : caterpillar

The caterpillar is the actual feeding stage of the butterfly. With some (e.g. peacock moth (Saturniidae), clucking (Lasiocampidae)) it is even the only one in which food is taken in at all. The moths of these species then only live to reproduce and die soon after hatching. Since the body volume of the caterpillars increases significantly, they have to moult several times until they have reached their final size. As a rule, they molt four to five times, each time about doubling in volume. For hormonally controlled moulting, the caterpillar swells until the old skin bursts and can be pushed backwards by muscle movements.

Segments and limbs

Similar to other insects with complete metamorphosis , the trunk of the caterpillar consists of evenly strung together segments. Butterfly caterpillars have 14 segments, consisting of the head, three breast segments and ten abdomen segments, the last three of which are mostly fused to form an anal segment. Like the moths, the caterpillars can be divided into the three areas of head, chest and abdomen.

The head is usually hardened by chitin deposits. Outside on the underside they have two to eight (mostly three) pairs of point eyes ( stemmata ). The most important characteristic, which also dominates the appearance of the head, is the mouthparts. In contrast to the stubby shaped antennae, they are very pronounced.

The three breast segments together form the breast. A pair of legs is placed on each of them (sternum), which are designed like the butterflies, but shorter and with which food is held. On the back of the first breast segment there is usually a neck shield, which consists of a plate hardened with chitin. On each side there is a pore-like opening ( stigma ) with which the tracheal system is supplied with oxygen . Such openings are only rarely found on the other breast segments.

The 10 abdominal segments that follow form the abdomen, which, however, is not clearly separated from the front part of the body. Each of these abdominal segments carries a stigma on breathing. The first two segments are legless, in contrast to the very similar larvae of sawflies , which only have a leg free segment. The next segments, mostly from the third to the sixth, carry limbs. These so-called belly legs , however, are not actually legs, but only unstructured skin protuberances that end with wreaths of hooks for better clinging. They are much more domed in shape than the real legs and usually widened like a suction cup at the end. On the tenth segment of the abdomen there is another pair of real legs that are much stronger. These are called pushers .

A spinneret is located between the breastbones and the bellybones, with which the caterpillars can create a spider thread.

variants

Several families deviate from this classic caterpillar shape: the primeval moths (Micropterigidae) have provided the first two abdominal segments with belly legs, the tensioners (Geometridae) lack the first three pairs of belly legs that have receded, and in some owl butterflies (Noctuidae) the first two are missing. The sternum of the snail spiders (Limacodidae) has receded into tiny stumps. All other pairs of legs have also receded, so that these caterpillars crawl like nudibranchs .

The smooth caterpillars of the knightly butterflies (Papilionidae) have a neck fork which they can quickly turn out when threatened and which gives off an unpleasant odor for enemies. The caterpillars of the spider moths and bud moths (Yponomeutidae), Noctuoidea and Geometroidea , for example, have similar protruding glands on their backs .

Many caterpillars have thorns or hairs to protect them from birds or parasitoid wasps and flies. Thorns are common in many butterfly caterpillars, and many bear moth caterpillars have thick hairs . Many bearer moths (Lymantriidae) have an unusual appearance due to tufts of hair of various colors and lengths. In humans, hair can sometimes cause skin irritation due to toxins, and it often loosens when it is brushed against the grain. If the hair does not contain toxins, it can cause itching and redness just by penetrating the skin, which looks like many small pinpricks. The caterpillar of the oak processionary moth ( Thaumetopoea processionea ) has over 600,000 poisonous hairs that can trigger allergies if people are only under infested trees.

internal structure

A small brain is located in the lower part of the head. A tubular bowel runs from the mouth to the end of the abdomen. In the caterpillar stage, the testicles of the later moths are already laid, they are located under the upper side of the abdomen. The inner organs are supplied with oxygen and nutrients by the green or yellow colored blood, also called the hemolymph . This is pumped through the body from a back vessel.

Mimicry, camouflage and warning

butterfly

Peacock butterfly

Hornet Glasswing

Due to the many different predators of butterflies, in the course of evolution for camouflage , deception and warning, drawings have often developed on their wings that either look like animal eyes, imitate dangerous and poisonous animals ( mimicry ) or through conspicuous coloration ( aposematism ) before they are inedible or warn about toxicity. Animal eyes can be found on the wings of the peacock butterfly , in the genus of the night peacock butterfly and the neotropical genus Caligo . The wrong eyes confuse robbers and tempt them to snap shut in the wrong place.

The hornet glass-winged bird ( Sesia apiformis ) looks very similar to hornets .

Some butterflies have colorful drawings on the upper side of the wings, but the underside is usually simply drawn and often appears like withered leaves. This means that they are well camouflaged with closed wings and adapted to their surroundings. Some moths imitate leaf veins on the underside of their wings. Especially moths, which usually sit on tree bark during the day, have a bark-like wing color. Bird droppings are also a widely used camouflage method. Caterpillars, butterflies and pupae not only look like bird droppings on the upper side of the leaf, they can also reinforce this impression by adopting an appropriate posture. An example of this is the elm harlequin ( Calospilos sylvata ).

There are also butterflies with a frightening color such as the Red Ribbon ( Catocala nupta ). When these moths rest inconspicuously on tree trunks, you can only see their brown-gray fore wings. But if they are startled and fly away, their bright red hind wings become visible.

Moths, whose bodies contain poisons and are therefore inedible for most of their potential enemies, warn them with a conspicuous color. Many moths of the Danaidae warn predators with conspicuous colors, as do the diurnal rams , which belong to the moths and contain hydrogen cyanide or other poisons.

In many species of white butterflies, noble butterflies and swallowtails, the females imitate poisonous butterflies from other families. Since the males are chosen by the females for mating, they must be easily identifiable by the females. Therefore, these show the original appearance of the species and have not developed any mimicry.

Caterpillars

Middle hawk caterpillar

The caterpillars also have many predators and have adapted just like the moths. Caterpillars that live on conifers , for example , usually have a longitudinal pattern that makes them appear to disappear between the needles. Thera firmata (Geometridae) imitates the pine needles of its forage plant with longitudinal stripes, its head also resembles a bud in shape and color.

Some caterpillars of the knightly butterfly family (Papilionidae), the caterpillars of Acronicta alni (Noctuidae) and some caterpillars of the genus Trilocha ( real spinner ) have developed a bird droppings mimesis .

Many caterpillars (Geometridae) imitate the shape of branches for camouflage. If they sit upside down on a branch, they become blurred, as if they were not as thick as a caterpillar but as flat as a leaf. A counter-shadow color, which is supposed to hide the cylindrical body of the caterpillar, also serves as camouflage. Other tensioners remain motionless with their backs clinging to a branch and thus look like a small branch with their perfectly matched color. They even form bud-like thickenings. The caterpillars of Nemoria arizonaria have developed a seasonal dimorphism , which means that they look different depending on the season. In spring they imitate the catkins of the forage plant, an oak. In summer they eat their leaves and imitate the branches they sit on.

An effective deterrent to predators through replicas of animal eyes has developed in parallel with some swallowtails and swarmers. They usually have two eye patches and can imitate small snakes through their posture . An example of this can be found in the swarmers of the genus Hemeroplanes.

Caterpillars, which are poisonous, warn predators with conspicuous colors, such as many species of the subfamily of the Danaidae , which also includes the monarch butterfly. These caterpillars do not hide and show themselves undisturbed on their feeding plants. Other caterpillars that are not poisonous but that have a similar coloration (mimicry) benefit from them.

Way of life and behavior

Nutrition of the adults

Painted lady (Cynthia cardui) sucks nectar

Small cabbage whites ( Pieris rapae ) gathered on a salt / mineral lick

Large and small blue butterflies suckle on a frog carcass

The adult animals ( Imagines ) take with her proboscis usually only liquid diet to mostly flower nectar . They fly to a large number of different flowers and are therefore also important for their pollination . Some plants with deep calyxes can only be pollinated by butterflies. But there are also other sweet liquids such as B. Vegetable juices, honeydew sucked by lice and the juice of rotting fruit. The skull hawk ( Acherontia atropos ) has specialized in vacuuming honeycombs .

On hot days, butterflies also like to suck water from small puddles. But they also do this to absorb mineral salts .

Few butterfly species feed wholly or partially on animal excrement , urine , sweat and tear fluid. Among the noble butterflies there are some species that prefer to suckle on animal excrement and sweat, such as the great blue butterfly and the kingfisher . The moth Lobocraspis griseifusa , Arcyophora sp. and Filodes fulvidorsalis of the families common borer (Pyralidae), owl butterfly (Noctuidae) and spanner (Geometridae) from Africa , Brazil and Southeast Asia drink tear fluid. By irritating the eyeball, the victim's tear production is stimulated. Usually larger animals such as large mammals and crocodiles are approached. However , there are no such animals in Madagascar , and since smaller mammals such as lemurs can drive away moths, the lachryphagous butterfly species such as Hemiceratoides hieroglyphica that live there have to switch to birds. This can only be done during the night. The moths have specially shaped proboscis that they can slide under the lids of the sleeping birds. Some tear fluid-drinking butterfly species also like to suck blood from open wounds. In some other species such as the Calyptra Thalictri ( Calyptra thalictri ) and subtropical species Calyptra eustrigata , Calyptra minuticornis , Calyptra orthograpta and Calyptra labilis from the family of cutworms (Noctuidae) is proboscis to a proboscis transformed and can deep to seven millimeters penetrate the skin of the host animal . These butterfly species sometimes feed on the blood of certain mammals and humans. They can therefore also transmit pathogens such as viruses .

The proboscis of some butterflies (for example peacock moth (Saturniidae), cluck (Lasiocampidae)) is more or less regressed. These butterflies can then no longer take in food. They live on their reserves for a few days to mate. Another exception are the primeval moths (Micropterigidae), which still have mandibles and eat pollen. The Heliconius species use pollen that they digest on their proboscis to get additional nitrogen, which enables them to live up to 8 months.

Diet and way of life of the caterpillars

The caterpillars , which are designed completely differently than the moths, also feed very differently. Usually the eggshell is eaten first after hatching. Afterwards, the caterpillars of most butterfly species eat leaves, needles, flowers, seeds or fruits of various plants, with many species specializing and dependent on certain plants ( monophagy ). Especially among these there are miners such as the leaf miners (Gracillariidae). These eat leaves or needles between their upper and lower skin ( cuticula ). Typically shaped feeding tunnels (mines) are created. Other caterpillars feed on organic waste, algae , lichens, or even predatory. Cannibalism also occurs in caterpillars when there is a lack of food. Some species even feed on a zoophag , including the caterpillar of the Hawaiian Hyposmocoma molluscivora , which feeds mainly on very small snails , which they stick to their seat with their silk threads .

The caterpillars of some butterfly families live in symbiosis or as social parasites with ants. In Central Europe, some bluebell caterpillars live together with scale ants and knot ants . The caterpillar secretes a sugary liquid with glands on its back. This attracts ants, which do not kill the caterpillar, as is usually the case with small insects, but only collect the sweet liquid and protect the caterpillar. The ants drum their legs on the caterpillar's back to stimulate the production of the sweet liquid. In the last stage of the caterpillar, they drag the caterpillar into their burrow. Here she takes on the smell of ants. It no longer lives symbiotically with the ants, but appears here as a social parasite and feeds on the brood and can also be fed by the ants, since it begs just like the brood of the ants. Although it still secretes a sugary fluid, that is out of proportion to the damage the ants suffer. It pupates during construction and hibernates depending on the season. This makes these butterflies one of the few that can survive in areas heavily populated by ants. In Mexico, the cuboid butterfly (Riodinidae) Anatole rossi lives symbiotically with ants. The ants lock the caterpillar in a burrow every evening to protect it from other predatory ants. During the day they guard the caterpillar and repel parasitoid wasps, for example. Even when it is dry, the ants bring the caterpillar into a cave where it can even survive forest fires. As a reward, the ants also receive a sweet liquid.

Flight behavior

Aurora butterfly in flight

Pigeon tails sucking nectar

Depending on the type of butterfly and its wing shape, several flight forms can be distinguished. From the fast soaring flight to the fast to the slow fluttering and gliding, they all show the gradations caused by the speed of the wing beat and the type of locomotion. The wings are propelled by several muscles of the thorax. The wings of the moth do not just flap up and down in flight, but rotate at the base so that they describe an "8". The swarmers (Sphingidae), whose wings flap like a hummingbird , are among the fastest butterflies . They can move at up to 50  km / h and in flight, while sucking the nectar, also stand still and even fly backwards. In addition to their normal flight, moths also have a repertoire of various courtship flights to offer. Longhorn moths (Adelidae) prance on the spot and rootworms (Hepialidae) swing back and forth. The very small butterflies can be carried by the wind and float in the air like plankton in water. In general, the wind is an important means of transport, not just to accelerate the spread of species. Some species of butterflies, especially real moths (Tineidae), are airborne.

As cold-blooded animals, they first have to warm up in order to be able to fly. Butterflies use the sun for this . Due to the large wing area, you can do this even when the sky is overcast. Moths need to heat themselves by vibrating the wings and the heat resulting from the movement of the muscles. When the body temperature gets too high on sunny, very hot days, the moths sit in the shade and cool by flapping their wings.

wintering

Butterflies that live in climates with cold seasons have to hibernate. As an image do this z. B. the lemon butterfly ( Gonepteryx rhamni ) or the peacock butterfly ( Inachis io ). They hide in hollow trees or in animal structures and stay there motionless. Most butterflies overwinter as a caterpillar, pupa or unhatched in an egg. Some caterpillars even wake up on very warm winter days and eat before they fall into winter rigor again. Likewise, some moths interrupt their winter rigor on warm days and fly around, without necessarily ingesting food. Depending on the region, the caterpillars or pupae need more than one winter to complete their development. This is the case, for example, in high mountains , for example with Parnassius species.

hikes

Some species of butterflies make long migrations; they are called migrant butterflies . This behavior is e.g. B. known from over 200 tropical species. The monarch butterfly ( Danaus plexippus ), known for its mass migrations in North America over thousands of kilometers, and the painted lady ( Vanessa cardui ), which migrates in many parts of the world, including from North Africa to Europe, have been particularly well researched . Painted ladies can often be spotted a few seconds apart as they fly in the same direction over the Alps or the open landscape.

In Europe, many species north of the Alps are not indigenous, which means that they cannot survive permanently and immigrate again every year. Examples of this are the painted lady, the pigeon tail ( Macroglossum stellatarum ) and the admiral . They fly north from their habitats in southern Europe and north Africa in spring ; partially they cross the Alps. Over the summer they live in Central Europe and parts of Northern Europe . They even form new generations here. When winter approaches, most of them fly back south. Some specimens try to hibernate and survive in mild winters or in specially protected hiding places.

The reason for the migration has not been sufficiently clarified, as the vast majority of animals cannot survive the winter. A strategy of accidental area expansion is obvious, at least the cause is also a withering of nectar plants in the Mediterranean area in summer. It is also possible that the migratory butterflies still follow a behavioral atavism, i.e. are influenced by other climatic conditions.

Reproduction and development

Primitive insects do not change their shape during their lifetime, they just get bigger and therefore have to shed their skin. In butterflies, metamorphosis fundamentally changes their appearance. It is called complete metamorphosis here, because besides the larval stage there is another stage, namely that of the pupa . Thus the butterflies have four stages of development: egg, caterpillar, pupa and butterfly.

Courtship behavior

Courtship is a very strictly observed ritual. It usually begins with a special flight and continues on the ground by walking around the female. During the flight the wings of the couple often touch each other or the female touches the wings of the male with her antennae. The partner's willingness to mate is reinforced by fragrances . The males only work at a short distance, but the female moths in particular attract the males over long distances. The territorial behavior of the males goes hand in hand with courtship . Depending on the species, certain areas such as treetops and hilltops ("Gipfelbalz"), sections of the path or small, bare spots are defended.

Great Restoration Bluebirds when mating

pairing

After the female has received the male's spermatophore , it enters a bladder ( bursa copulatrix ), from which the sperm finally enter the seminal vesicle, where they are often stored for long periods of time. The eggs are fertilized only during egg-laying, during which they slide past the opening of the seminal vesicle. In some butterfly species, the female can fertilize her eggs without the male ( parthenogenesis ).

Egg and egg laying

Females of the hard hay spanner ( Siona lineata ) laying eggs

Clutches of the tree whitefly ( Aporia crataegi )

The eggs of the butterflies are among the most complex of the insects. There is an enormous variety of shapes that vary between narrow spindle-shaped, oval, spherical, hemispherical, lens-shaped and flat cylindrical. In addition, there are various surface structures that often make the hard eggshell ( chorion ) look bizarre. The eggs are rarely smooth. There are ribbed, dented, star-shaped, decorated, hairy and jagged eggs. The pattern is basically regular. There are two main types of eggs: flat and upright eggs. In the former, the umbilical bulge ( micropyle ) through which the sperm penetrates the egg cell during fertilization is on the upper side, in the case of upright eggs the micropyle is on the back ( dorsal ). Most species of butterflies have eggs of the second type. The oxygen supply is carried out by pores ( aeropyles ), are in the egg itself but even chambers filled with air. The size of the eggs varies between 0.5 and 2 millimeters. The coloring of the eggs couldn't be more varied. Above all, the color varies during development, which is mostly due to the fact that you can see the color of the growing caterpillar through the slightly translucent eggshells. Most eggs are initially light and darken to a black or dark blue by the time the caterpillar hatches.

The laying of eggs is very different depending on the species. It is also adapted to the shape and color of the eggs. Most butterflies lay their eggs individually, in pairs, or in small and large groups. Some put their entire egg supply in a single position. Depending on the type, 20 to over 1,000 eggs are laid. They are usually attached to the base with a sticky substance. These are arranged either randomly in single or multi-layer clusters or regularly in ice mirrors next to one another or in a ring around plant parts or stems. They can also be covered by hairs of the anal bush, which the female strips off and sticks to the eggs with a secretion. This means that they are better camouflaged from predators. However, some species just drop their eggs randomly on the ground. It is usually placed on the appropriate forage plant so that the caterpillars find food as soon as they hatch. But there are also species that randomly distribute their eggs on unsuitable plants. The caterpillars usually hatch after two to three weeks, but this is also possible after less than a week. When the eggs overwinter, which happens in many species, the caterpillars sometimes only hatch after six months. Their first development is usually completed before winter, only hatching is delayed. After hatching, many species eat the eggshell first. Presumably, this serves to absorb nutrients and vital microorganisms that have been passed from the mother to the egg.

Caterpillar

Caterpillar of a tensioner

Sloe-brush caterpillar

The caterpillars mostly lead a hidden life and are also well adapted to their surroundings. They are usually green or brown in color. The caterpillars of the swarmers are among the largest in Europe. They can reach a length of 15 centimeters. Some caterpillars spin the leaves of the food plants together, such as in the case of the admiral, or they roll up a leaf and eat this tube from the inside, as do many curlers (Tortricidae), which is how this family got its name.

In some species one can observe a social behavior. The caterpillars of the processionary moth (Thaumetopoeidae), for example, live together in large webs and move together in long “processions” to their food sources.

Doll

Pupation of a peacock butterfly in 60-second steps

Doll of the Elm Harlequin

When the caterpillar is an adult, it begins to pupate by shedding its skin for the last time. Then the metamorphosis to the butterfly takes place. The caterpillar organs are broken down or reshaped and transformed into butterfly organs and the entire external shape of the animals changes. The pupae of the butterflies are basically mummy pupae. This means that all body attachments (antennae, leg systems and wing sheaths) are glued to the body with a putty. Only the primeval moths (Micropterigidae) have free pupae in which the limbs are not glued together. The three body sections of the mummy dolls are difficult, but the appendages and the head are easy to recognize. The doll is almost immobile. She can only swing her abdomen sideways and make rolling movements. Some (e.g. glass winged birds (Sesiidae) and wood drills (Cossidae)) have thorns on their abdominal segments, with which they can work their way forward in their feeding ducts in the wood to open them more easily after hatching. In the more primitive families, however, the appendages of the body can also be moved easily.

In the butterfly families summarized under the butterflies, two types of pupae are distinguished according to the type of attachment to the base. Some are called fall dolls, the other belt dolls. Falling dolls hang freely dangling with hooks from a web anchor that is attached to the base. Belt dolls are connected to a twig or the like by a thread that is spun around the middle of the body like a belt. In addition, like the first, these dolls are anchored to the underside with a web cushion, but these dolls usually do not hang down, but are attached to the top by the web. This type is also used with other types of butterflies. The pupae of the remaining butterfly families pupate either freely on the ground or in a more or less tightly spun silk web. This is called a cocoon . The silk is made from special spinning glands located on the lower lip. Precautions are necessary so that the finished butterfly can leave its sometimes very solid pupa. Either a round lid is provided, which is then pushed open from the inside, or a trap allows the butterfly to crawl out without uninvited guests being able to penetrate through it. This is e.g. B. the case with the lesser night peacock butterfly ( Saturnia pavonia ). Other species secrete a liquid from the mouth opening that softens the cocoon.

Because in some species (especially butterflies) the pupae are formed without a protective web and are therefore very sensitive, they must be well camouflaged. Unlike the webbed species, they cannot rely on enemies to abandon their attack because of the tough and difficult-to-penetrate outer skin. Therefore, the defenseless pupae are often perfectly camouflaged like fresh or dried leaves. Some pupae can even make chirping noises to confuse predators.

The doll phase (so-called doll rest) usually lasts two to four weeks. However, some species overwinter as a pupa. Here the moths develop before winter, but only hatch in spring. Sometimes the moths rest in the pupae for longer than one winter. The spring wool juicers ( Eriogaster lanestris ) can stay in their pupae for up to seven years before they hatch.

The silk moth ( Bombyx mori ) is probably the best-known example of a cocoon, because it is used to make silk . The cocoon consists of a single thread, over 500 meters long, which is automatically unwound and rewound onto spools.

Slip

Play media file

hatching peacock butterfly. The speed of the film is true to the original.

When the pupa reaches the final stage of its development, it is visibly darker in color and the drawing of the wings can often be seen through the pupal shell. The pupa bursts open at predetermined seams and the butterfly hatches. In the case of cocoons, the butterfly either leaves the web through a provided lid or it forces itself outside. Then he begins to pump air into the body to allow the doll's shell to burst open further. Then he pulls the body out of the shell and clings to the outside with his legs. The wings are still hanging limply from the body, they are inflated by the moths pumping blood into the veins that are still empty. When the wings are unfolded to full size, they have flattened out at the same time. After unfolding, the wing veins lose their function in that they also dry out. While the wings are drying, the moth excretes metabolic products from pupa in the form of a red drop ( meconium ). Then the butterfly finally takes off on its first flight. It can mate, with the new oviposition the life cycle is completed.

The lifespan of the moths varies greatly. It is only a single day for real sack-carriers (Psychidae), but can also last up to ten months including periods of rest, as for example with the lemon butterfly ( Gonepteryx rhamni ). Ingesting moths understandably live longer than those with stunted mouthparts. The average age of butterflies is two to three weeks.

Natural enemies

The moths and especially the caterpillars are exposed to countless predators . The main enemies of adults and caterpillars are probably insectivorous birds worldwide. How strong the predation pressure is from birds is shown by the countless adaptations of butterflies to optically preying predators, such as mimicry, camouflage colors, eye spots etc. (see above). In Europe, for example, tits live mainly on caterpillars during the breeding season and raise their young with them. Some species of birds, such as cuckoos , specialize in long-haired caterpillars, which other birds avoid.

Only a few species of birds hunt moths, e.g. B. the goat milkers . The main enemy of nocturnal butterflies are bats. Here, too, numerous evolutionary adaptations of moths to their main enemies have arisen as part of an "evolutionary race". B. the ability to hear ultrasound sounds or utter them yourself.

In addition, adults such as caterpillars are preyed on by numerous insectivorous vertebrate and invertebrate species.

Specialized predators and parasitoids

Hawk-caterpillar with numerous parasitoid cocoons

There are also numerous predators and parasites or parasitoids that specialize in butterflies or certain butterfly species . Above all, there are some common wasps , parasitic wasps , wasps , brackish wasps and flies . The butterfly-hunting silver-mouthed wasp ( Lestica subterranea ), for example, paralyzes small butterflies with its poison and pulls them into their burrow, where several butterflies are made available to each larva in several chambers. Most of the parasitoids feed as larvae in the eggs, caterpillars and butterflies. There are those who live individually and grow up to the size of the host, others are very small and eat the host by the dozen together. They usually feed on the non-vital areas of the caterpillars first and only kill them at the end. Among the beetles, pupa predators ( Calosoma spec. ) In particular live mainly on caterpillars.

Distribution and habitats

One of the most important properties of butterflies is that they have the ability to adapt to a wide range of environmental conditions. They have occupied almost all ecological niches and can be found almost everywhere. Some groups show a fine niche like the high mountain butterflies (including Boloria , Parnassius , Erebia ), which often have very narrow habitat requirements. Butterflies can thus be used as bio-indicators in nature conservation. You not only need the appropriate larval habitats with the forage plants, but also require the presence of certain habitat types of the alpine level such as alpine mats and rubble heaps in which cushion plants ( e.g. Silene acaulis ) and espalier bushes ( Dryas octopetala or Salix retusa ) occur. These are needed as fodder plants or hiding places during bad weather.

As herbivores ( phytophagous ) butterflies are closely tied to the flora and thus the vegetation periods . This describes the period between the start of plant growth in spring and the onset of the next winter. The further north you go in Europe or the higher you climb in the mountains, the shorter the growing seasons. This is accompanied by a change in the flora, including the butterfly fauna. Butterflies can be found at great heights. In the Alps, for example, the Matterhorn bear ( Holoarctica cervini ) comes up to 3,200 meters above sea level. NN and the glacier butterfly ( Oenis glacialis ) lives at 2000 to 3000 meters above sea level. NN. In the Rocky Mountains, Colias meadii reaches up to 3,600 meters above sea level. NN. In the Himalayas, a subspecies of the swallowtail , Papilio machaon ladakensis, which also occurs in Europe , often flies at 3500 to 4200 meters above sea level. NN. Some species of the genus Parnassius go even higher , such as P. acco, P. delphius and P. simo, which was found at 4900 meters, and P. delphius workmani, which was found at 5700 meters. Butterflies are particularly rich in species in the tropics , as the temperature conditions are ideal for their vital activity. Solitary biodiversity can be found in the Iguaçu National Park, for example, with around 800 species . Species diversity decreases towards the temperate and cold zones. This climatic dependence can also be determined in mountainous areas; the number of species here decreases with increasing altitude. Their spread is limited by the species-specific conditions of temperature and humidity, but also by the occurrence of the caterpillar forage plants.

Habitat requirements

Each species of butterfly makes diverse, species-specific demands on the properties of its environment. Only when these are fulfilled can the animals survive. One of the most important conditions for the distribution and occurrence of the predominantly herbivorous ( phytophagous ) butterflies is the sufficient availability of food plants, both for butterflies and caterpillars. While some species adopt many food plants and are widely distributed, many species rely on a few or only one nutrient plant. They are therefore also limited in their distribution. In many cases, special landscape or vegetation structures are imperative. Further essential parameters for habitat suitability are the microclimate , the intensity of the daily and seasonal temperature fluctuations and the duration of the vegetation period . For the various butterfly species, a distinction is made between so-called biotope residents who are loyal to their location and biotope complex residents who leave the “caterpillar site” in the larval stage, and different biotope inhabitants who can inhabit several different biotopes and live in them like biotope inhabitants. Butterflies colonize almost all types of biotope from forests, dry grasslands , meadows , wetlands and ruderal meadows to parks and gardens.

Butterfly plants

Numerous flowers of North American columbines specialize in the pollination of swarms. Their long nectar extensions are only attractive to this group of pollinators

Many butterflies and plants are closely coordinated in their development. Even Charles Darwin had to adapt the flowers of Angraecum sesquipedale the existence of a then unknown Motte postulated. In the genus Columbine, there have been adaptations to pollinators ("pollination syndrome"), hawks (pschophilia), bumblebees (cantharophilia) or hummingbirds (ornithophilia). They serve as selective evolutionary features; here, for example, the flowers of Aquilegia pubescens and Aquilegia caerulea are adapted to enthusiasts. They always have white flowers, while those species that are visited by hummingbirds are yellow or red ( Aquilegia formosa ) and have developed shorter nectar processes. In Central Europe, a few plant species are the food source for a large number of butterfly caterpillars, such as the nettle , whose numerous guests are also known as nettle butterflies . These include peacock butterfly , small fox , C-butterfly , painted lady , admiral and the nettle borer .

The record as food plants for caterpillars is held in Central Europe by oak and willow , on which caterpillars of over 100 species live. The willows include the large and small Schiller butterfly , mourning cloak , large fox , evening peacock butterfly , night peacock butterfly , red ribbon , large and small forked tail .

Almost a hundred species settle on poplars and birches , including many that also live on willows. The caterpillars of 65 species live on hawthorn , 54 each on sloe , blackberry and raspberry ,  44 on hazel and  33 on roses .

Hazard and protection

Large fire butterfly (endangered)

Numerous species of butterflies are endangered. In Europe, the main cause of this is habitat loss. Wetlands have been drained, open land has been afforested or cultivated, and agriculture has been intensified. This not only lost the habitat of butterflies and caterpillars. Often important forage plants are deprived of their growth base and the butterfly species adapted to them can no longer develop without them. The application of nitrogen fertilizers not only increases the nitrogen content in the forage plants but also the mortality rate in the butterflies. According to a study, the survival rate of the larvae of all six examined species has decreased by at least a third, and this with the amounts of fertilizer used today in agriculture. Accordingly, the fertilizer law (in the case of Germany) should urgently be tightened. The cultivation of GM maize can also have a negative impact on protected species. As well as the overgrowth of areas that have just been created by humans, such as heaths and semi- arid grasslands , which without continued use, e.g. B. as sheep pasture , shrub, reduces the habitats that sensitive butterfly species need to survive.

A number of species are dependent on very special landscapes. The species that are true to their location disappear when their biotope is destroyed or changes. Numerous rams (Zygaenidae) are z. B. rely on dry grass and the plants growing there. If these areas are not cared for and covered in bush, the rams will also disappear. Other important habitats are bogs and other wetlands that some species of bluish rely on. The blue gentian ant ( Maculinea alcon ) and the dark blue button ( Maculinea nausithous ) are extremely faithful to their location and only stay in the vicinity of their forage plants, which grow on wet meadows and moors. This close connection to a certain type of biotope represents a high risk potential for these species. As soon as there is serious interference in the habitat of these species, a process of displacement or an islanding of the distribution areas takes place, which can quickly lead to the complete extinction of a population . The causes of displacement are sometimes complex and have not yet been adequately researched.

The increasing light pollution is a great danger, especially for moths . They are attracted to street lights and other lighting and stay near the light sources all night. The next day they are either eaten by birds or they die of malnutrition or exhaustion. Lots of pesticides such as cyhalothrin and tebufenozide are sprayed to combat moths in the commercial forests .

Numerous specially protected butterfly species are listed in Appendix 1 to the Federal Species Protection Ordinance. It prohibits the gathering and trapping of these species, but it cannot address the actual threat posed by habitat loss. You can also get an overview of the endangered species in the red list of large butterflies. Only 50% of all butterfly species in Germany are not endangered, 2% are already extinct or lost.

One form of commercial exploitation of nature that is particularly prevalent in many Asian countries is the use of butterfly wings for mosaics and other questionable decorations. Here the risk of inventory damage cannot be dismissed out of hand.

Collections

history

Butterfly collection in the natural history cabinet Waldenburg in Saxony

The collecting of butterflies began in the 17th and 18th centuries, where the collector was met with a lot of ridicule. It was operated by a few, since after the emergence of entomology as a separate science in the 18th century, insect collections generally provided an important basis for scientific work as a documentation and comparison tool. This importance continues to this day; the creation and maintenance of type collections is particularly important for the foundation of the nomenclature .

In the middle of the 19th century collecting began to establish itself as a hobby . It was a popular hobby of numerous nature-loving people of all ages and countries that left its mark on literature, music and film. Most butterfly collections were limited to butterflies , as these had the most magnificent colors.

rating

The effect of collecting on the population of butterflies is often controversial. There is also the cliché that the former popularity of butterfly collecting contributed to the decline of many species. A distinction must be made here according to the collector's intention.

There is no evidence whatsoever that collecting activities within the framework of a scientific objective using the usual fishing methods has ever permanently damaged the population of a species. On the contrary, it is precisely the butterfly experts who, through their research, can assess population developments and provide the basis for butterfly protection. In this context, collecting always serves to secure evidence and the possibility of carrying out reliable determinations and also being able to control them retrospectively.

Hobby collectors without scientific ambitions, on the other hand, are generally viewed more critically, as they are primarily interested in an attractive collection. Nevertheless, in the opinion of experts, their activity is in most cases not very important, especially since this group of nature lovers has now mostly chosen photography as an alternative. But also in earlier times other reasons for the decline in stocks were decisive. On the other hand, hobby collectors are often customers of insect fairs and thus create demand for exotic, sometimes particularly rare species that are smuggled on a large scale.

Famous collectors and collections

A famous butterfly collector was Walter Rothschild, 2nd Baron Rothschild from London , who bequeathed his collection to the Natural History Museum after his death . Another collector was the Russian-American writer and butterfly researcher Vladimir Nabokov .

With more than 7 million butterfly specimens, the butterfly collection of the Bavarian Zoological State Collection is one of the largest natural history collections in the world and probably the largest butterfly collection ever. The collection of the Berlin Natural History Museum with almost four million copies is also important .

The butterfly collection of the Natural History Museum in Vienna houses one of the largest collections of day and night moths in the world. Approximately 3.5 million prepared and several hundred thousand unprepared specimens are in around 11,000 shops.

There are butterfly zoos that specialize in displaying butterflies, and most of them are also bred there. Examples of this are the Butterfly House in Vienna and the Butterfly Garden in the Museum of Science in Boston .

Ecological importance

Marsh horn clover ram

Rose lichen bear

Bioindicators

Butterflies are suitable as so-called pointer animals or bioindicators in nature and landscape planning due to their diverse and specific habitat ties and their often very close ties to only a few plant species . Important bio-indicators include various representatives from the ram family (Zygaenidae). Studies in the Düben Heath have shown that some species from the bear moth family (Arctiidae) can become important as bio-indicators. These are the rose lichen bear ( Miltochrista miniata ), the red-collar lichen bear ( Atolmis rubricollis ) and the yolk-yellow lichen bear ( Eilema sororcula ). These species occur mainly in areas in which more than 40% of an area is overgrown with lichen ( lichen dominance more than 40%). The lack of these species in areas with less than 10% lichen dominance is most likely due to pollution from pollutants. The general rule is that unaffected areas have a high number of species, areas with high pollution levels have a low number of species. The above-mentioned species can already be missing if there is still a sufficient stock of food plants.

Pollinators

Numerous species of butterflies specialize in certain plants and their flowers. As a result, the plants are also dependent on the moths, because only they can pollinate the specially shaped flowers. For example, Xanthopan morgani can penetrate the particularly narrow flower necks of orchids and pollinate them with its 20 centimeter long trunk . The yucca z. B. is dependent on the yuccamot ( Tegeticula yuccasella ) as a pollinator, which can penetrate the pollen with greatly extended lip probes.

Development history

Since some caddis flies also have similarly scaled wings (e.g. the African species Pseudoleptocerus chirindensis ) and both secrete or process silk via their labial glands in the larval stage , they are traced back to a common ancestor " Amphiesmenoptera ". Genetic studies also point in this direction: the Serpin -2 gene is well comparable in butterflies and caddis flies, which suggests that the gene is highly conserved and the two orders are more closely related .

The history of the development of butterflies begins around 135 million years ago with the beginning of the Cretaceous ( Mesozoic ). The emergence and upswing of butterflies is closely related to the appearance of flowering plants and is inconceivable without them. Butterflies turned out to be an extremely successful group of animals. They managed to conquer all areas of the mainland.

Researchers found the oldest known fossils in Siberian sedimentary rocks from the Jura , which undoubtedly belong to the butterflies, they were assigned to the species Eolepidopterix jurassica Rasnitsyn, 1983. A paraphyletic collection of older fossils, called “Necrotauliidae”, is now part of the common parent group of butterflies and caddis flies. Fossil butterflies, such as Undopterix sukatshevae, also originate from the Cretaceous Period of Siberia . Parasabatinca aftimacrai made of Cretaceous Lebanese amber is the oldest undoubted fossil representative of the primeval moths , the most original living butterflies. Some animals of this species got caught in the resin of conifers and were preserved in amber .

One of the youngest groups of butterflies, and one of those with the most incomplete fossil record, are the butterfly families. With Doritites bosniaskii a European representative comes from the Miocene of Italy. Prodryas persephone was described from the older Eocene sediments of Florissant, Colorado, USA, about 34 million years old . The Middle Eocene ( Lutetium ) Praepapilio gracilis and Praepapilio colorado ( Papilionidae ) are probably the oldest described butterflies. Richard Vane-Wright dates the origin of the butterflies to "not older than 70 million years".

Systematics

External system

Within the subclass of flying insects (Pterygota), the butterflies belong to the superordinate order of the new winged insects (Neoptera). The caddis flies are the closest relatives of the butterflies within the parent group. The butterflies probably split off from the caddis flies in the Mesozoic Era.

The following cladogram illustrates the relationships:

New winged wing (Neoptera)	Eumetabola    Paraneoptera   Holometabola     NN    Reticulated winged (Neuropterida)      Coleopteroid   NN    Hymenoptera (Hymenoptera)   Mecopteroida     Amphiesmenoptera    Caddisflies (Trichoptera)      Butterflies (Lepidoptera)   Antliophora    Beak fly (Mecoptera)   NN    Fly (Diptera)      Fleas (Siphonaptera)      Paurometabola

Internal system

Little fox (butterfly)

Reporting corridor owl (moth)

→ Main article : Systematics of butterflies

The classic system of butterflies is presented very inconsistently. With over 180,000 species in around 130 families and 44 superfamilies, the butterflies represent the second largest insect order next to the beetles . They are divided into four sub- orders : Zeugloptera , Aglossata , Heterobathmiina and Glossata .

The suborder Zeugloptera includes the most pristine butterflies, which, like the representatives of the Aglossata and Heterobathmiina, still have biting and chewing mouthparts. The caterpillars of the Zeugloptera are the only ones to have developed ventral feet. The suborder Glossata covers the majority of the butterfly families. They have specialized mouthparts ( proboscis ) and their caterpillars do not have belly feet, just skin protuberances.

Other doctrines argue that butterflies can only be grouped into two sub-categories. This is on the one hand the subordination of the Zeugloptera, which have chewing mouthparts (and feed on pollen), on the other hand the order of the glossata, which have small to very large proboscis (and feed on plant saps and nectar).

The division into small butterflies (Microlepidoptera) and large butterflies (Macrolepidoptera) or into butterflies and moths has no scientific basis. It is used to simplify practical work with butterflies. Spanners (Geometridae), which are classically counted among the moths, are, for example, more closely related to the moths than to the other moths. Nevertheless, the terms are still used today for practical reasons. Butterflies were divided into these four categories according to external characteristics and the supposed day and night activity or according to an arbitrarily chosen size.

Butterflies and humans

GDR postage stamp ( Alpenapollo )

Silk production in Khota, China

Dried caterpillars in the Orodara market, Burkina Faso

Avenue eaten by the horse chestnut leaf miner

Economic use

The spider threads of the caterpillars of the East Asian family of real spinners (Bombycidae), especially the silk moth ( Bombyx mori ), are the raw material for silk . In order to obtain the yarn of this textile fiber , the pupae are killed with boiling water or hot steam around the tenth day after the cocoon has been completed. The spinning thread is carefully unwound and carefully cleaned before further processing in the silk weaving mill (see silk construction ). The caterpillars are bred for silk in China , Japan , India and southern Europe. By crossing the silk threads you get different colors like golden yellow and other nuances. The caterpillars feed exclusively on the leaves of the mulberry trees , which were cultivated for their breeding and were also imported into Europe .

The caterpillars and pupae of some species are used as protein-rich food. In East Asia, boiled silkworm pupae are eaten as a snack. The caterpillars of Gonimbrasia belina , the so-called “mopane worms”, are eaten in southern Africa . Dried caterpillars are also offered in West Africa.

Parasitic species can be used to control plants that are classified as pests . The cactus moth was used in Australia to contain neophytes .

pest

Clothes moth larva in felt

Agricultural monocultures offer optimal conditions for certain butterfly species, as well as for other insects, under which large numbers of individuals can develop. This is especially the case in the tropics and subtropics , because higher temperatures encourage rapid development of the animals.

The culture damage is caused by caterpillar damage on leaves. The caterpillars of the large ( Pieris brassicae ) and the small cabbage white butterfly ( Pieris rapae ) can destroy entire cabbage fields . Other species such as oak moth ( Tortrix viridana ) are considered forest pests. In Europe there are repeated mass reproductions that lead to large parts of the herbaceous plants being eaten up. The trees are severely weakened and can eventually die off due to additional factors such as drought. Renewing leaves are often also attacked by oak powdery mildew ( Microsphaera quercina ). Healthy oaks can even survive an infestation of several years. Some areas are affected differently from year to year, others only occasionally. Other caterpillars develop in fruit , potatoes , flower bulbs , cotton or seeds .

For those who think economically, mass developments ( calamities ) are a problem. In addition to the use of insecticides , nesting boxes for birds have been found to be very effective as part of biological pest control . In addition, attractant traps and the confusion method are also used to catch males or to confuse them so that they cannot find the females.

The clothes moth ( Tineola bisselliella ) is particularly worth mentioning . Your caterpillars feed on a multitude of different animal substances, such as wool , felt , feathers , silk and fur , and thus become a problem in the household.

Vectors of disease

As already explained under nutrition , some subtropical species that feed on tear fluid and / or mammalian blood can transmit various infectious diseases by mechanical means (see infection routes , blood-sucking insects or haematophagy ) .

Butterflies in Art and the Non-Biological Sciences

Mythological meaning

By pupating and hatching from the apparently lifeless cocoon after months of external rest, the butterfly was the symbol of rebirth and immortality in antiquity and is still the symbol of resurrection in Christian art today . Moths and / or dolls can therefore be found on numerous tombs. In many Asian regions they are seen as bad luck charms and heralds of death, but often also as symbols of a new beginning.

In ancient Greece ψυχή psyche was the name for the butterfly, because the adults were seen as the souls of the dead. The doll was called νεκύδαλλο, which means "cover of the dead". In Greek and Roman mythology, the soul often appears with butterfly wings. Released from death, the soul can move away from its shell and rise freely upwards.

The peoples of Central America also associated the butterfly and its metamorphosis with myths. Different species have been equated with different goddesses or fire or the messenger of death. Black butterflies were and are still considered messengers of death.

Many customs and rites associated with butterflies are also known from the Indians of North America. This mystical relationship has partially survived to the present day and there is rich literature on it.

Visual arts

Carl Spitzweg : The butterfly hunter

There are numerous paintings, such as The Butterfly Hunter by Carl Spitzweg , which show pictorial representations of butterflies.

literature

Poems

Butterflies are also a popular motif in poetry. Examples of poems about butterflies:

• Johann Gottfried Herder : Dear, light, airy thing, butterfly
• Johann Wolfgang von Goethe : The butterfly
• Friedrich von Schlegel : The butterfly
• Heinrich Heine : The butterfly is in love with the rose
• Eduard Mörike : To Clara (In the vineyard) , Citronenfalter in April
• Christian Morgenstern : A butterfly flies over me
• Rainer Maria Rilke : In the churchyard of Ragaz written down, I
  (from: Exposed on the mountains of the heart )
• Hermann Hesse : Blue butterfly (from: The blossom branch )
• Inger Christensen : The butterfly valley

Of Wilhelm Busch 's poem comes She was a little flower :

It was a pretty and delicate flower,
bloomed brightly in the sunshine.
He was a young butterfly
Who hung blissfully on the flower.
Often a
bee would come with a hum and nibble and whisper around there.
A bug often crawled
up and down the pretty little flower.
Oh God, how it
went so painfully through the soul of the butterfly .
But what horrifies him most,
the worst of all came at last.
An old donkey ate the whole
plant that he loved so dearly.

Even Friedrich Hebbel dealt with butterflies:

On a flower, red and burning, sat
a butterfly sucking its honey,
And so forgetting itself in its lust
that it did not even fly further in front of me.
I wanted to see how sweet the flower was,
And broke it off: it stayed where it was;
I braided it in my lover's hair:
he sucked away, as if dissolved in delight!

Novels

The novel Papillon (French butterfly) is one of the most famous prison novels. In it, the author Henri Charrière describes his experiences in French Guiana, where he spent over ten years before he finally managed to escape. He then lived a contented life until he reads what he believed to be a very poorly written fictional novel. Sure that his life was more exciting, he wrote his experiences in exercise books, the publication of which became a world best seller. The novel was later filmed twice and so also went successfully around the world. 1973 with Dustin Hoffman and Steve McQueen in the lead roles; The remake followed in 2017 based on the same script.

The novel The Butterfly Catcher by Sabine M. Gruber describes in "Lolita manner" the obsessive love of the young pianist Herbie for his underage piano student, the nanny Aurelia. Herbie's second great obsession is catching and skewering butterflies. The background story is musical: a recital at New York's Carnegie Hall with the Schöne Müllerin by Franz Schubert .

The wonderful description of a (Bavarian) lepidopterist can be found in Lord Jim by Joseph Conrad .

Natural sciences

In the context of chaos theory there is a pictorial illustration of non-linear dynamic, deterministic systems: the “ butterfly effect ”. A well-known example from meteorology is the question of whether the delicate flapping of a butterfly's wings in one region of the world can trigger a storm in another region far away with all possible dynamic consequences and chain reactions.

See also

Portal: Butterflies  - Overview of Wikipedia content on butterflies

literature

• MJ Scoble: The Lepidoptera: Form, Function and Diversity. The Oxford University Press, Oxford UK 1995.
• NP Kristensen (Ed.): Lepidoptera, Moths and Butterflies. In: Handbook of Zoology. 2 volumes. Walter de Gruyter, Berlin / New York 1999, 2003.
• David J. Carter, Brian Hargreaves, Alexander Pelzer: Caterpillars and butterflies of Europe and their fodder plants (OT: A Field Guide to Caterpillars of Butterflies and Moths in Britain and Europe. ) Paul Parey, Hamburg / Berlin 1987, ISBN 3-490-13918 -6 .
• Günter Ebert (Ed.): The butterflies of Baden-Württemberg , Ulmer Verlag, Stuttgart
  • Volume 1 (1993): Butterfly I = Knight Butterfly (Papilionidae), Whiteflies (Pieridae), Edelfalter (Nymphalidae). ISBN 3-8001-3451-9 .
  • Volume 2 (1993): Tagfalter II = Augenfalter (Satyridae), Bluebirds (Lycaenidae), Dickkopffalter (Hesperidae). ISBN 3-8001-3459-4 .
  • Volume 3 (1993): Moth I = root borer (Hepialidae), wood borer (Cossidae), ram (Zygaenidae), snail moth (Limacodidae), sack bearer (Psychidae), window spot (Thyrididae). ISBN 3-8001-3472-1 .
  • Volume 4 (1994): Nachtfalter II = Bombycidae, Endromidae, Lasiocampidae, Lemoniidae, Saturniidae, Sphingidae, Drepanidae, Notodontidae, Dilobidae, Lymantriidae, Ctenuchidae, Nolidae. ISBN 3-8001-3474-8 .
  • Volume 5 (1997): Moths III = Sesiidae, Arctiidae, Noctuidae. ISBN 3-8001-3481-0 .
  • Volume 6 (1997): Moth IV = Owls (Noctuidae), Part 2. ISBN 3-8001-3482-9 .
  • Volume 7 (1998): Moths V = Owls (Noctuidae), 3rd part. ISBN 3-8001-3500-0 .
  • Volume 8 (2001): Moth VI = Spanner (Geometridae), 1st part. ISBN 3-8001-3497-7 .
  • Volume 9 (2003): Moth VII = Spanner (Geometridae), Part 2. ISBN 3-8001-3279-6 .
• Manfred Koch , Wolfgang Heinicke: We identify butterflies. 3. Edition. Neumann, Radebeul 1991, ISBN 3-7402-0092-8 .
• Lepidopterists working group: Butterflies and their habitats, Volume 1. Swiss Confederation for Nature Conservation, Basel 1987 ISBN 3-85587-402-2 .
• Heiko Bellmann : The new Kosmos butterfly guide. Butterflies, caterpillars and forage plants. Franckh-Kosmos, Stuttgart 2003, ISBN 3-440-09330-1 .
• Bernard Skinner: Color Identification Guide to Moths of the British Isles. Penguin UK, 1999, ISBN 0-670-87978-9 .
• Paul Smart et al .: Cosmos Encyclopedia of Butterflies. The butterflies of the earth - Over 2000 life-size species. (OT: The Illustrated Encyclopedia of the Butterfly World in Color. ) 2nd edition. Franckh-Kosmos, Stuttgart 1987, ISBN 3-440-04466-1 .
• Tom Tolman, Richard Lewington: The butterflies of Europe and Northwest Africa. Franckh-Kosmos, Stuttgart 1998, ISBN 3-440-07573-7 .
• Hans-Josef Weidemann: Butterflies: observe, determine. Naturbuch-Verlag, Augsburg 1995, ISBN 3-89440-115-X .
• Hans-Josef Weidemann, Jochen Köhler: Moths. Weirdos and hawkers. Naturbuch-Verlag, Augsburg 1996, ISBN 3-89440-128-1 .
• F. Nemos: Europe's most famous butterflies. Description of the most important species and instructions for knowing and collecting butterflies and caterpillars. Oestergaard, Berlin approx. 1895, with 18 color plates hdl.handle.net (PDF; 77 MB).

On the motif “butterfly” in mythology and art

• Reinhard Breymayer : Book and Butterfly. A portrait of Friedrich Hölderlin's dean of Nürtingen, Jakob Friedrich Klemm (1733–1793). With reference to the parallel motifs in Mörike's poem "In the Vineyard" . In: Stuttgart works on German studies, No. 307. Verlag Hans-Dieter Heinz, Akademischer Verlag Stuttgart, Stuttgart 1994 [1995], ISBN 3-88099-311-4 , pp. 83-113.
• Roswitha Kirsch-Stracke, Petra Widmer: Butterfly and Opium Poppy. On the symbolic content of animal and plant representations on tombs. In: Stadt und Grün (The Garden Department). Vol. 48, no. 8, pp. 520-526, Patzer Verlag, 1999.
• Peter Godzik : What does the caterpillar know about the butterfly? Departure to change, freedom and immortal life , EB-Verlag, Hamburg-Schenefeld 2007, ISBN 3-936912-72-6

Web links

Commons : Butterflies  - Collection of images, videos and audio files

Wiktionary: Butterfly  - explanations of meanings, word origins, synonyms, translations

Wikiquote: Butterfly  Quotes

• Lepidoptera near Fauna Europaea
• deutschlandfunk.de , Environment and Consumers, May 9, 2016, Peter Kaiser: Why the butterflies are disappearing
• funet.fi: Lepidoptera and some other life forms (engl.)
• lepiforum.de: Identification of butterflies (Lepidoptera) and their pre-imaginal stages
• leps.it: Moths and Butterflies of Europe and North Africa (engl.)
• nhm.ac.uk: host plants of butterflies (database)
• Distribution of butterflies in Germany
• schmetterling-raupe.de ( Association for the Environment and Nature Conservation Germany )
  • Information on the topic of butterfly death from the BUND
• How are our butterflies? ( Federal Office for the Environment )
• www.naturraum-stux.de/ Website with many photos suitable for identifying butterflies and caterpillars

Individual evidence

1. ↑ Erik J. van Nieukerken, Lauri Kaila, Ian J. Kitching, Niels P. Kristensen, David C. Lees, Joël Minet, Charles Mitter, Marko Mutanen, Jerome C. Regier, Thomas J. Simonsen, Niklas Wahlberg, Shen-Horn Yen, Reza Zahiri, David Adamski, Joaquin Baixeras, Daniel Bartsch, Bengt Å. Bengtsson, John W. Brown, Sibyl Rae Bucheli, Donald R. Davis, Jurate De Prins, Willy De Prins, Marc E. Epstein, Patricia Gentili-Poole, Cees Gielis, Peter Hättenschwiler, Axel Hausmann, Jeremy D. Holloway, Axel Kallies , Ole Karsholt, Akito Y. Kawahara, Sjaak (JC) Koster, Mikhail V. Kozlov, J. Donald Lafontaine, Gerardo Lamas, Jean-François Landry, Sangmi Lee, Matthias Nuss, Kyu-Tek Park, Carla Penz, Jadranka Rota, Alexander Schintlmeister, B. Christian Schmidt, Jae-Cheon Sohn, M. Alma Solis, Gerhard M. Tarmann, Andrew D. Warren, Susan Weller, Roman V. Yakovlev, Vadim V. Zolotuhin, Andreas Zwick (2011): Order Lepidoptera Linnaeus , 1758. In: Zhang, Z.-Q. (Editor) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 212-221.
2. ↑ Christopher O'Toole: Firefly Encyclopedia of Insects and Spiders. 2002, ISBN 1-55297-612-2 .
3. ^ Ole Karsholt: The lepidoptera of Europe: a distributional checklist. 380 pp., Apollo Books, Stenstrup 1996, ISBN 87-88757-01-3 .
4. ↑ See Lemma Fifalter in the Swiss Idiotikon .
5. ↑ Cf. Lemma Butterfly In: Kluge. Etymological dictionary of the German language . Arranged by Elmar Seebold. 25th, revised and expanded edition. Berlin, Boston (2011): De Gruyter.
6. ↑ Audio sample: Video about Lepidoptera at 0:07.
7. ↑ See Online Etymology Dictionary: Lepidoptera und lepido- (English)
8. ↑ Kenneth F. Kitchell Jr .: Animals in the Ancient World from A to Z . Routledge, 2014, ISBN 978-1-317-57743-0 . P. 19
9. ^ Hermann & Anna Levinson (2009): Butterflies (Lepidoptera) in Egyptian and Greek antiquity. News of the German Society for General and Applied Entomology 23 (3): 121-132.
10. ^ Ian C. Beavis: Butterflies, Moths and Wood-Boring Larvae . In Ian C. Beavis (editor): Insects and other Invertebrates in Classical Antiquity . Liverpool University Press, 1988, ISBN 978-0-85989-284-1 -
11. ↑ ^{a b c d e f g h i j k l} Heiko Bellmann: The new Kosmos butterfly guide. Butterflies, caterpillars and forage plants. Franckh-Kosmos, Stuttgart 2003, ISBN 3-440-09330-1 .
12. ↑ ^{a b c d e f g h i j k l m} Roger Philips, David Carter: Kosmos Atlas butterfly guide, European day and night butterflies. Franckh-Kosmos, Stuttgart 1991, ISBN 3-440-06306-2 .
13. ^ Lionel G. Higgins, Norman D. Rilley: The butterflies of Europe and Northwest Africa. (A Field Guide to the Butterflies of Britain and Europe) Verlag Paul Parey, 1971, ISBN 3-490-02418-4 .
14. ↑ ^{a b} Thomas C. Emmel: Wonderful and mysterious world of butterflies . Bertelsmann Lexikon-Verlag, Gütersloh and Berlin 1976, ISBN 3-570-00893-2 , p. 22nd f .
15. ↑ ^{a b c} Malcolm J. Scoble: The Lepidoptera: Form, Function and Diversity . Oxford University Press, Oxford 1995, ISBN 0-19-854952-0 , pp. 108 ff . (English).
16. ↑ ^{a b} Thomas C. Emmel: Wonderful and mysterious world of butterflies . Bertelsmann Lexikon-Verlag, Gütersloh and Berlin 1976, ISBN 3-570-00893-2 , p. 60 .
17. ↑ Thomas C. Emmel: Wonderful and mysterious world of butterflies . Bertelsmann Lexikon-Verlag, Gütersloh and Berlin 1976, ISBN 3-570-00893-2 .
18. ↑ Malcolm J. Scoble: The Lepidoptera: Form, Function and Diversity . Oxford University Press, Oxford 1995, ISBN 0-19-854952-0 , pp. 121 ff . (English).
19. ↑ Lawrence E. Gilbert: Pollen Feeding and Reproductive Biology of Heliconius Butterflies. Proc. Nat. Acad. Sci. USA, 69 (6): 1403-1407, 1972 PDF
20. ^ The University of Arizona College of Agriculture and Life Sciences and The University of Arizona Library: Heliconius. In: Tree of Life Web Project. Retrieved November 12, 2007 . 
21. ↑ Thomas C. Emmel: Wonderful and mysterious world of butterflies . Bertelsmann Lexikon-Verlag, Gütersloh and Berlin 1976, ISBN 3-570-00893-2 , p. 55 f .
22. ↑ ^{a b} Otakar Kudrna: Butterflies of Europe, Vol. 2, Introduction in Lepidopterology. Aula-Verlag, Wiesbaden 1990, ISBN 3-89104-033-4 , p. 78.
23. ↑ Zoltan Varga 2014: Biogeography of the High-Mountain Lepidoptera of the Balkan Peninsula. Ecologia Montenegrina, 1 (3): 140-168
24. ↑ ^{a b c d} Hans-Josef Weidemann: Butterfly: observe, determine. Naturbuch-Verlag, Augsburg 1995, ISBN 3-89440-115-X .
25. ^ Justen B. Whittall & Scott A. Hodges 2007: Pollinator shifts drive increasingly long nectar spursin columbine flowers. Nature, Vol 447/7 June 2007: 706-711
26. ↑ Helmut Hintermeier: The privet and his guests , in Allgemeine Deutsche Beekeeper newspaper , November 2008, pages 30-31.
27. ↑ Stefan Abrahamczyk, Thomas Wohlgemuth, Michael Nobis, Reto Nyffeler, Michael Kessler: Shifts in food plant abundance for flower ‐ visiting insects between 1900 and 2017 in the canton of Zurich, Switzerland. In: Ecological Applications. 2020, doi : 10.1002 / EAP.2138 .
28. ↑ Dramatic loss of forage crops for insects. In: uni-bonn.de . April 24, 2020, accessed April 28, 2020 . 
29. ↑ Susanne Kurz, Thilo Heinken, Thomas Fartmann: Nitrogen enrichment in host plants increases the mortality of common Lepidoptera species . In: Oecologia . tape 188 , 2018, p. 1227-1237 , doi : 10.1007 / s00442-018-4266-4 .
30. ↑ GM maize can harm protected butterflies. In: LID.CH Agricultural Information Service. August 7, 2020, accessed on August 17, 2020 . 
31. ↑ Jens Blankennagel: Controversial insecticide use in Brandenburg: From Monday, "Liquid Karate" will fall from the sky. In: berliner-kurier.de . May 3, 2019, accessed May 4, 2019 . 
32. ↑ Franconian forests sprayed with insecticide: This pest is the reason. In: nordbayern.de . May 2, 2019, accessed May 4, 2019 . 
33. ↑ lepidoptera collection. In: nhm-wien.ac.at. Natural History Museum Vienna, accessed on September 22, 2018 . 
34. ↑ Günter Ebert (Ed.): The Butterflies of Baden-Württemberg, Volume 5, Moths III (Sesiidae, Arctiidae, Noctuidae). Ulmer Verlag, Stuttgart 1997, ISBN 3-8001-3481-0 .
35. ↑ John Huxley, Peter C. Barnard: Wing ‐ scales of Pseudoleptocerus chirindensis Kimmins (Trichoptera: Leptoceridae) . In: Zoological journal of the Linnean Society . 92, No. 3, March 15, 1988, pp. 285-312. doi : 10.1111 / j.1096-3642.1988.tb01514.x .
36. ^ Paul Whalley: A review of the current fossil evidence of Lepidoptera in the Mesozoic . In: Biological Journal of the Linnean Society . 28, No. 3, June 28, 2008, pp. 253-271. doi : 10.1111 / j.1095-8312.1986.tb01756.x .
37. ↑ D. Grimaldi & MS Engel: Evolution of the Insects . Cambridge University Press, 2005, ISBN 0-521-82149-5 .
38. ↑ Naoyuki Yonemura, et al. : Conservation of a pair of serpin 2 genes and their expression in Amphiesmenoptera . In: Insect Biochemistry and Molecular Biology . 42, No. 5, 2012, pp. 371-380. doi : 10.1016 / j.ibmb.2012.01.008 .
39. ↑ Otakar Kudrna: Butterfly - Life, Endangerment, Protection. Mayer-Verlag, Ravensburg, 1991, ISBN 3-473-46082-6 .
40. ↑ ^{a b} Jae-Cheon Sohn, Conrad Labandeira, Donald Davis, Charles Mitter (2002): An annotated catalog of fossil and subfossil Lepidoptera (Insecta: Holometabola) of the world. Zootaxa 3286: 1-132.
41. ^ Richard Vane-Wright: Butterflies at that awkward age. In: Nature. Vol. 498, 477-479. nature.com (PDF)
42. ↑ Endopterygota. Tree of Life Web Project. 1995, accessed August 31, 2006 . 
43. ↑ Roswitha Kirsch-Stracke, Petra Widmer: Butterfly and Opium Poppy. On the symbolic content of animal and plant representations on tombs. In: Stadt und Grün (The Garden Department). Vol. 48, no. 8, pp. 520-526, Patzer Verlag, 1999.
44. ↑ Strongly abbreviated from "petalouda (butterflies) " - πεταλούδα in the Greek language Wikipedia, there also numerous references.
45. ↑ see The LiederNet Archive
46. ^ Rainer Maria Rilke (1875–1926), What was written down in the churchyard in Ragaz. insects.ch, accessed on August 30, 2013 . 
47. ↑ Blue butterfly. Karolina Hoffmann, accessed on August 31, 2006 . 
48. ^ Wilhelm Busch: Critique of the Heart . Friedrich Bassermann, Heidelberg 1874 ( She was a flower [accessed on September 20, 2018]).
49. ^ Friedrich Hebbel: Poems. In: Spiegel Online. April 26, 2006, archived from the original on August 28, 2014 ; accessed on September 22, 2018 .