94-435: Polygonia comma , the eastern comma , is a North American butterfly in the family Nymphalidae , subfamily Nymphalinae . This butterfly is seasonally variable. The upperside of the summer form's hindwings are all black, whereas the winter forms hindwings are reddish orange. The underside of both forms is striped with dark and light brown. There is a silvery comma mark in the middle of the hindwing in both forms. Its wingspan
188-631: A diapause (resting) stage, and the hatching may take place only in spring. Some temperate region butterflies, such as the Camberwell beauty , lay their eggs in the spring and have them hatch in the summer. Butterfly larvae, or caterpillars, consume plant leaves and spend practically all of their time searching for and eating food. Although most caterpillars are herbivorous, a few species are predators : Spalgis epius eats scale insects , while lycaenids such as Liphyra brassolis are myrmecophilous , eating ant larvae. Some larvae, especially those of
282-493: A butterfly cannot fly until the wings are unfolded. A newly emerged butterfly needs to spend some time inflating its wings with hemolymph and letting them dry, during which time it is extremely vulnerable to predators. The colourful patterns on many butterfly wings tell potential predators that they are toxic. Hence, the genetic basis of wing pattern formation can illuminate both the evolution of butterflies as well as their developmental biology . The colour of butterfly wings
376-570: A butterfly through metamorphosis has held great appeal to mankind. To transform from the miniature wings visible on the outside of the pupa into large structures usable for flight, the pupal wings undergo rapid mitosis and absorb a great deal of nutrients. If one wing is surgically removed early on, the other three will grow to a larger size. In the pupa, the wing forms a structure that becomes compressed from top to bottom and pleated from proximal to distal ends as it grows, so that it can rapidly be unfolded to its full adult size. Several boundaries seen in
470-417: A common family. In some species, such as the great spangled fritillary , the eggs are deposited close to but not on the food plant. This most likely happens when the egg overwinters before hatching and where the host plant loses its leaves in winter, as do violets in this example. The egg stage lasts a few weeks in most butterflies, but eggs laid close to winter, especially in temperate regions, go through
564-571: A few butterflies (e.g., harvesters ) eat harmful insects, and a few are predators of ants , while others live as mutualists in association with ants. Culturally, butterflies are a popular motif in the visual and literary arts. The Smithsonian Institution says "butterflies are certainly one of the most appealing creatures in nature". The Oxford English Dictionary derives the word straightforwardly from Old English butorflēoge , butter-fly; similar names in Old Dutch and Old High German show that
658-444: A few species. Some butterflies have organs of hearing and some species make stridulatory and clicking sounds. Many species of butterfly maintain territories and actively chase other species or individuals that may stray into them. Some species will bask or perch on chosen perches. The flight styles of butterflies are often characteristic and some species have courtship flight displays. Butterflies can only fly when their temperature
752-498: A function of water depth. For example, the Spanish Dancer nudibranch (genus Hexabranchus ), among the largest of tropical marine slugs, potently chemically defended, and brilliantly red and white, is nocturnal and has no known mimics. Mimicry is to be expected as Batesian mimics with weak defences can gain a measure of protection from their resemblance to aposematic species. Other studies have concluded that nudibranchs such as
846-606: A hard-ridged outer layer of shell, called the chorion . This is lined with a thin coating of wax which prevents the egg from drying out before the larva has had time to fully develop. Each egg contains a number of tiny funnel-shaped openings at one end, called micropyles ; the purpose of these holes is to allow sperm to enter and fertilize the egg. Butterfly eggs vary greatly in size and shape between species, but are usually upright and finely sculptured. Some species lay eggs singly, others in batches. Many females produce between one hundred and two hundred eggs. Butterfly eggs are fixed to
940-503: A leaf before eggs are laid on it. Many butterflies use chemical signals, pheromones ; some have specialized scent scales ( androconia ) or other structures ( coremata or "hair pencils" in the Danaidae). Vision is well developed in butterflies and most species are sensitive to the ultraviolet spectrum. Many species show sexual dimorphism in the patterns of UV reflective patches. Colour vision may be widespread but has been demonstrated in only
1034-416: A leaf with a special glue which hardens rapidly. As it hardens it contracts, deforming the shape of the egg. This glue is easily seen surrounding the base of every egg forming a meniscus. The nature of the glue has been little researched but in the case of Pieris brassicae , it begins as a pale yellow granular secretion containing acidophilic proteins. This is viscous and darkens when exposed to air, becoming
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#17328557536321128-610: A letter to Alfred Russel Wallace dated 23 February 1867, Charles Darwin wrote, "On Monday evening I called on Bates & put a difficulty before him, which he could not answer, & as on some former similar occasion, his first suggestion was, 'you had better ask Wallace'. My difficulty is, why are caterpillars sometimes so beautifully & artistically coloured?" Darwin was puzzled because his theory of sexual selection (where females choose their mates based on how attractive they are) could not apply to caterpillars since they are immature and hence not sexually active. Wallace replied
1222-547: A longer lifespan of several months as adults. The thorax of the butterfly is devoted to locomotion. Each of the three thoracic segments has two legs (among nymphalids , the first pair is reduced and the insects walk on four legs). The second and third segments of the thorax bear the wings. The leading edges of the forewings have thick veins to strengthen them, and the hindwings are smaller and more rounded and have fewer stiffening veins. The forewings and hindwings are not hooked together ( as they are in moths ) but are coordinated by
1316-448: A necessity for their successful establishment. Many butterflies, such as the painted lady , monarch, and several danaine migrate for long distances. These migrations take place over a number of generations and no single individual completes the whole trip. The eastern North American population of monarchs can travel thousands of miles south-west to overwintering sites in Mexico . There is
1410-442: A predator from a distance, but are warning-like from a close proximity, allowing for an advantageous balance between camouflage and aposematism. Warning coloration evolves in response to background, light conditions, and predator vision. Visible signals may be accompanied by odors, sounds or behavior to provide a multi-modal signal which is more effectively detected by predators. Unpalatability, broadly understood, can be created in
1504-646: A reverse migration in the spring. It has recently been shown that the British painted lady undertakes a 9,000-mile round trip in a series of steps by up to six successive generations, from tropical Africa to the Arctic Circle — almost double the length of the famous migrations undertaken by monarch. Spectacular large-scale migrations associated with the monsoon are seen in peninsular India. Migrations have been studied in more recent times using wing tags and also using stable hydrogen isotopes . Butterflies navigate using
1598-412: A segmented palp. Adjoining these is the labium-hypopharynx which houses a tubular spinneret which is able to extrude silk. Caterpillars such as those in the genus Calpodes (family Hesperiidae) have a specialized tracheal system on the 8th segment that function as a primitive lung. Butterfly caterpillars have three pairs of true legs on the thoracic segments and up to six pairs of prolegs arising from
1692-435: A silken girdle may be spun to keep the pupa in a head-up position. Most of the tissues and cells of the larva are broken down inside the pupa, as the constituent material is rebuilt into the imago. The structure of the transforming insect is visible from the exterior, with the wings folded flat on the ventral surface and the two halves of the proboscis, with the antennae and the legs between them. The pupal transformation into
1786-665: A single generation, and a few in cold locations may take several years to pass through their entire life cycle. Butterflies are often polymorphic , and many species make use of camouflage , mimicry , and aposematism to evade their predators. Some, like the monarch and the painted lady , migrate over long distances. Many butterflies are attacked by parasites or parasitoids , including wasps , protozoans , flies , and other invertebrates, or are preyed upon by other organisms. Some species are pests because in their larval stages they can damage domestic crops or trees; other species are agents of pollination of some plants. Larvae of
1880-399: A single leaf together with silk. The chrysalis is brown and covered with spines. Winter-form adults overwinter; some will also migrate south for the winter. The dark form of comma is frequently confused with the dark form of the question mark ( P. interrogationis ), but the two can readily be distinguished by the shape of the comma mark on the underside. The pale form is easily confused with
1974-410: A time-compensated sun compass. They can see polarized light and therefore orient even in cloudy conditions. The polarized light near the ultraviolet spectrum appears to be particularly important. Many migratory butterflies live in semi-arid areas where breeding seasons are short. The life histories of their host plants also influence butterfly behaviour. Butterflies in their adult stage can live from
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#17328557536322068-528: A total of about 20,000 species. Traditionally, butterflies have been divided into the superfamilies Papilionoidea and the moth-like Hedyloidea . Recent work has discovered that Hedylidae, the only family within Hedyloidea, is nested within the Papilionoidea, meaning that Papilionoidea would be synonymous with Rhopalocera. The relationships between the rest of the 6 families are extremely well resolved, which
2162-470: A trend towards multivoltinism . Courtship is often aerial and often involves pheromones . Butterflies then land on the ground or on a perch to mate. Copulation takes place tail-to-tail and may last from minutes to hours. Simple photoreceptor cells located at the genitals are important for this and other adult behaviours. The male passes a spermatophore to the female; to reduce sperm competition, he may cover her with his scent, or in some species such as
2256-486: A tubular proboscis which is curled up at rest and expanded when needed to feed. The first and second maxillae bear palps which function as sensory organs. Some species have a reduced proboscis or maxillary palps and do not feed as adults. Many Heliconius butterflies also use their proboscis to feed on pollen; in these species only 20% of the amino acids used in reproduction come from larval feeding, which allow them to develop more quickly as caterpillars, and gives them
2350-725: A variety of ways. Some insects such as the ladybird or tiger moth contain bitter-tasting chemicals, while the skunk produces a noxious odor, and the poison glands of the poison dart frog , the sting of a velvet ant or neurotoxin in a black widow spider make them dangerous or painful to attack. Tiger moths advertise their unpalatability by either producing ultrasonic noises which warn bats to avoid them, or by warning postures which expose brightly coloured body parts (see Unkenreflex ), or exposing eyespots . Velvet ants (actually parasitic wasps) such as Dasymutilla occidentalis both have bright colors and produce audible noises when grabbed (via stridulation ), which serve to reinforce
2444-437: A water-insoluble, rubbery material which soon sets solid. Butterflies in the genus Agathymus do not fix their eggs to a leaf; instead, the newly laid eggs fall to the base of the plant. Eggs are almost invariably laid on plants. Each species of butterfly has its own host plant range and while some species of butterfly are restricted to just one species of plant, others use a range of plant species, often including members of
2538-422: A week to nearly a year depending on the species. Many species have long larval life stages while others can remain dormant in their pupal or egg stages and thereby survive winters. The Melissa Arctic ( Oeneis melissa ) overwinters twice as a caterpillar. Butterflies may have one or more broods per year. The number of generations per year varies from temperate to tropical regions with tropical regions showing
2632-902: Is 4.5–6.4 cm (1.8–2.5 in). The eastern comma may be spotted in woods near rivers, ponds, marshes, swamps, and other water sources. This butterfly seldom visits flowers, but rather feeds on sap, rotting fruit, salts and minerals from puddling , and dung. Caterpillars feed on paper birch ( Betula papyrifera ), false nettle ( Boehmeria cylindrica ), hackberry ( Celtis occidentalis ), hops ( Humulus ), wood nettle ( Laportea canadensis ), currants and gooseberries ( Ribes spp.), basswood ( Tilia americana ), American elm ( Ulmus americana ), nettle ( Urtica ), and clearweed ( Pilea pumila ). The green eggs are laid singly or in stacks under host plant leaves and stems. The spiny larva varies in color from pale green to yellow to white and to even black. The solitary larva feeds on leaves at night. Older larvae construct daytime leaf shelters by pulling
2726-432: Is a sufficiently successful strategy to have had significant effects on the evolution of both aposematic and non-aposematic species. Non-aposematic species have often evolved to mimic the conspicuous markings of their aposematic counterparts. For example, the hornet moth is a deceptive mimic of the yellowjacket wasp; it resembles the wasp, but has no sting. A predator which avoids the wasp will to some degree also avoid
2820-450: Is above 27 °C (81 °F); when it is cool, they can position themselves to expose the underside of the wings to the sunlight to heat themselves up. If their body temperature reaches 40 °C (104 °F), they can orientate themselves with the folded wings edgewise to the sun. Basking is an activity which is more common in the cooler hours of the morning. Some species have evolved dark wingbases to help in gathering more heat and this
2914-520: Is argued to be sufficient for such species to be considered aposematic. It has been proposed that aposematism and mimicry is less evident in marine invertebrates than terrestrial insects because predation is a more intense selective force for many insects, which disperse as adults rather than as larvae and have much shorter generation times. Further, there is evidence that fish predators such as blueheads may adapt to visual cues more rapidly than do birds, making aposematism less effective. However, there
Polygonia comma - Misplaced Pages Continue
3008-458: Is derived from tiny structures called scales, each of which have their own pigments . In Heliconius butterflies, there are three types of scales: yellow/white, black, and red/orange/brown scales. Some mechanism of wing pattern formation are now being solved using genetic techniques. For instance, a gene called cortex determines the colour of scales: deleting cortex turned black and red scales yellow. Mutations, e.g. transposon insertions of
3102-403: Is especially evident in alpine forms. As in many other insects, the lift generated by butterflies is more than can be accounted for by steady-state, non-transitory aerodynamics . Studies using Vanessa atalanta in a wind tunnel show that they use a wide variety of aerodynamic mechanisms to generate force. These include wake capture , vortices at the wing edge, rotational mechanisms and
3196-469: Is experimental evidence that pink warty sea cucumbers are aposematic, and that the chromatic and achromatic signals that they provide to predators both independently reduce the rate of attack. Blue-ringed octopuses are venomous. They spend much of their time hiding in crevices whilst displaying effective camouflage patterns with their dermal chromatophore cells. However, if they are provoked, they quickly change colour, becoming bright yellow with each of
3290-523: Is exploited in Müllerian mimicry , where species with strong defences evolve to resemble one another. By mimicking similarly coloured species, the warning signal to predators is shared, causing them to learn more quickly at less of a cost. A genuine aposematic signal that a species actually possesses chemical or physical defences is not the only way to deter predators. In Batesian mimicry , a mimicking species resembles an aposematic model closely enough to share
3384-410: Is extruded and inserted into the female's vagina. A spermatophore is deposited in the female, following which the sperm make their way to a seminal receptacle where they are stored for later use. In both sexes, the genitalia are adorned with various spines, teeth, scales and bristles, which act to prevent the butterfly from mating with an insect of another species. After it emerges from its pupal stage,
3478-451: Is good, especially in some species in the blue/violet range. The antennae are composed of many segments and have clubbed tips (unlike moths that have tapering or feathery antennae). The sensory receptors are concentrated in the tips and can detect odours. Taste receptors are located on the palps and on the feet. The mouthparts are adapted to sucking and the mandibles are usually reduced in size or absent. The first maxillae are elongated into
3572-563: Is native to the Americas, but in the nineteenth century or before, spread across the world, and is now found in Australia, New Zealand, other parts of Oceania, and the Iberian Peninsula . It is not clear how it dispersed; adults may have been blown by the wind or larvae or pupae may have been accidentally transported by humans, but the presence of suitable host plants in their new environment was
3666-399: Is no statistical relationship between the two factors. Nudibranch molluscs are the most commonly cited examples of aposematism in marine ecosystems, but the evidence for this has been contested, mostly because (1) there are few examples of mimicry among species, (2) many species are nocturnal or cryptic, and (3) bright colours at the red end of the colour spectrum are rapidly attenuated as
3760-465: Is restricted to the males, and studies have suggested that the nutrients collected may be provided as a nuptial gift , along with the spermatophore, during mating. In hilltopping , males of some species seek hilltops and ridge tops, which they patrol in search for females. Since it usually occurs in species with low population density, it is assumed these landscape points are used as meeting places to find mates. Butterflies use their antennae to sense
3854-430: Is selected by predators. Concurrent reciprocal selection (CRS) may entail learning by predators or it may give rise to unlearned avoidances by them. Aposematism arising by CRS operates without special conditions of the gregariousness or the relatedness of prey, and it is not contingent upon predator sampling of prey to learn that aposematic cues are associated with unpalatability or other unprofitable features. Aposematism
Polygonia comma - Misplaced Pages Continue
3948-781: Is significant. The sound-producing rattle of rattlesnakes is an acoustic form of aposematism. Sound production by the caterpillar of the Polyphemus moth, Antheraea polyphemus , may similarly be acoustic aposematism, connected to and preceded by chemical defences. Similar acoustic defences exist in a range of Bombycoidea caterpillars. The existence of aposematism in marine ecosystems has been debated. Many marine organisms, particularly those on coral reefs, are brightly coloured or patterned, including sponges, corals, molluscs, and fish, with little or no connection to chemical or physical defenses. Caribbean reef sponges are brightly coloured, and many species are full of toxic chemicals, but there
4042-441: Is strong enough to allow seemingly maladaptive traits to persist despite other factors working against the trait. Once aposematic individuals reach a certain threshold population, for whatever reason, the predator learning process would be spread out over a larger number of individuals and therefore is less likely to wipe out the trait for warning coloration completely. If the population of aposematic individuals all originated from
4136-692: Is summarized in the below cladogram. Papilionidae [REDACTED] Hedylidae [REDACTED] Hesperiidae [REDACTED] Pieridae [REDACTED] Nymphalidae [REDACTED] Lycaenidae [REDACTED] Riodinidae [REDACTED] Butterfly adults are characterized by their four scale-covered wings, which give the Lepidoptera their name ( Ancient Greek λεπίς lepís, scale + πτερόν pterón, wing). These scales give butterfly wings their colour: they are pigmented with melanins that give them blacks and browns, as well as uric acid derivatives and flavones that give them yellows, but many of
4230-405: Is taken up by the gut, but there may also be large silk glands, and special glands which secrete distasteful or toxic substances. The developing wings are present in later stage instars and the gonads start development in the egg stage. When the larva is fully grown, hormones such as prothoracicotropic hormone (PTTH) are produced. At this point the larva stops feeding, and begins "wandering" in
4324-461: Is to prevent attack, by warning potential predators that the prey animal has defenses such as being unpalatable or poisonous. The easily detected warning is a primary defense mechanism, and the non-visible defenses are secondary. Aposematic signals are primarily visual, using bright colors and high-contrast patterns such as stripes. Warning signals are honest indications of noxious prey, because conspicuousness evolves in tandem with noxiousness. Thus,
4418-473: The Lycaenidae , form mutual associations with ants. They communicate with the ants using vibrations that are transmitted through the substrate as well as using chemical signals. The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions . Large blue ( Phengaris arion ) caterpillars trick Myrmica ants into taking them back to the ant colony where they feed on
4512-469: The Weis-Fogh ' clap-and-fling ' mechanism. Butterflies are able to change from one mode to another rapidly. Butterflies are threatened in their early stages by parasitoids and in all stages by predators, diseases and environmental factors. Braconid and other parasitic wasps lay their eggs in lepidopteran eggs or larvae and the wasps' parasitoid larvae devour their hosts, usually pupating inside or outside
4606-557: The X chromosome . If so, predators would learn to associate the colour with unpalatability from males with the trait, while heterozygous females carry the trait until it becomes common and predators understand the signal. Well-fed predators might also ignore aposematic morphs, preferring other prey species. A further explanation is that females might prefer brighter males, so sexual selection could result in aposematic males having higher reproductive success than non-aposematic males if they can survive long enough to mate. Sexual selection
4700-452: The ZW sex-determination system where females are the heterogametic sex (ZW) and males homogametic (ZZ). Butterflies are distributed worldwide except Antarctica, totalling some 18,500 species. Of these, 775 are Nearctic ; 7,700 Neotropical ; 1,575 Palearctic ; 3,650 Afrotropical ; and 4,800 are distributed across the combined Oriental and Australian / Oceania regions. The monarch butterfly
4794-600: The non-coding DNA around the cortex gene can turn a black-winged butterfly into a butterfly with a yellow wing band. When the butterfly Bicyclus anynana is subjected to repeated inbreeding in the laboratory, there is a dramatic decrease in egg hatching. This severe inbreeding depression is considered to be likely due to a relatively high mutation rate to recessive alleles with substantial damaging effects and infrequent episodes of inbreeding in nature that might otherwise purge such mutations. Although B. anynana experiences inbreeding depression when forcibly inbred in
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#17328557536324888-400: The satyr comma ( P. satyrus ), which usually occurs north and west of the eastern comma's range. They can be distinguished by the upperside color, which is orange brown in the comma and tawny yellowish brown in P. satyrus ; by the underside pattern, which tends to be mottled in the comma but appears to be more longitudinally streaked in P. satyrus ; and by the row of pale submarginal spots on
4982-557: The superfamilies Hedyloidea (moth-butterflies in the Americas) and Papilionoidea (all others). The oldest butterfly fossils have been dated to the Paleocene , about 56 million years ago, though molecular likely originated in the Cretaceous . Butterflies have a four-stage life cycle , and like other holometabolous insects they undergo complete metamorphosis . Winged adults lay eggs on
5076-449: The 50-60 rings flashing bright iridescent blue within a third of a second. It is often stated this is an aposematic warning display, but the hypothesis has rarely if ever been tested. The mechanism of defence relies on the memory of the would-be predator; a bird that has once experienced a foul-tasting grasshopper will endeavor to avoid a repetition of the experience. As a consequence, aposematic species are often gregarious. Before
5170-535: The Apollos ( Parnassius ) plugs her genital opening to prevent her from mating again. The vast majority of butterflies have a four-stage life cycle: egg , larva (caterpillar), pupa (chrysalis) and imago (adult). In the genera Colias , Erebia , Euchloe , and Parnassius , a small number of species are known that reproduce semi-parthenogenetically ; when the female dies, a partially developed larva emerges from her abdomen. Butterfly eggs are protected by
5264-430: The abdomen, generally with short prolegs on segments 3–6 and 10; the three pairs of true legs on the thorax have five segments each. Many are well camouflaged; others are aposematic with bright colours and bristly projections containing toxic chemicals obtained from their food plants. The pupa or chrysalis, unlike that of moths, is not wrapped in a cocoon. Many butterflies are sexually dimorphic . Most butterflies have
5358-405: The abdominal segments. These prolegs have rings of tiny hooks called crochets that are engaged hydrostatically and help the caterpillar grip the substrate. The epidermis bears tufts of setae , the position and number of which help in identifying the species. There is also decoration in the form of hairs, wart-like protuberances, horn-like protuberances and spines. Internally, most of the body cavity
5452-502: The adult colour pattern are marked by changes in the expression of particular transcription factors in the early pupa. The reproductive stage of the insect is the winged adult or imago . The surface of both butterflies and moths is covered by scales, each of which is an outgrowth from a single epidermal cell. The head is small and dominated by the two large compound eyes . These are capable of distinguishing flower shapes or motion but cannot view distant objects clearly. Colour perception
5546-467: The air for wind and scents. The antennae come in various shapes and colours; the hesperiids have a pointed angle or hook to the antennae, while most other families show knobbed antennae. The antennae are richly covered with sensory organs known as sensillae . A butterfly's sense of taste is coordinated by chemoreceptors on the tarsi , or feet, which work only on contact, and are used to determine whether an egg-laying insect's offspring will be able to feed on
5640-434: The ant eggs and larvae in a parasitic relationship. Caterpillars mature through a series of developmental stages known as instars . Near the end of each stage, the larva undergoes a process called apolysis , mediated by the release of a series of neurohormones . During this phase, the cuticle , a tough outer layer made of a mixture of chitin and specialized proteins , is released from the softer epidermis beneath, and
5734-658: The blues, greens, reds and iridescent colours are created by structural coloration produced by the micro-structures of the scales and hairs. As in all insects, the body is divided into three sections: the head, thorax , and abdomen . The thorax is composed of three segments, each with a pair of legs. In most families of butterfly the antennae are clubbed, unlike those of moths which may be threadlike or feathery. The long proboscis can be coiled when not in use for sipping nectar from flowers. Nearly all butterflies are diurnal , have relatively bright colours, and hold their wings vertically above their bodies when at rest, unlike
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#17328557536325828-658: The brighter and more conspicuous the organism, the more toxic it usually is. This is in contrast to deimatic displays , which attempt to startle a predator with a threatening appearance but which are bluffing, unsupported by any strong defences. The most common and effective colors are red, yellow, black, and white. These colors provide strong contrast with green foliage, resist changes in shadow and lighting, are highly chromatic, and provide distance dependent camouflage . Some forms of warning coloration provide this distance dependent camouflage by having an effective pattern and color combination that do not allow for easy detection by
5922-524: The brown & green eatable catterpillars, would enable birds to recognise them easily as at a kind not fit for food, & thus they would escape seizure which is as bad as being eaten ." Since Darwin was enthusiastic about the idea, Wallace asked the Entomological Society of London to test the hypothesis. In response, the entomologist John Jenner Weir conducted experiments with caterpillars and birds in his aviary, and in 1869 he provided
6016-440: The concept aposematism in his book The Colours of Animals . He described the derivation of the term as follows: The second head (Sematic Colours) includes Warning Colours and Recognition Markings: the former warn an enemy off, and are therefore called Aposematic [Greek, apo , from, and sema , sign] Aposematism is paradoxical in evolutionary terms, as it makes individuals conspicuous to predators, so they may be killed and
6110-441: The desiccated husk. Most wasps are very specific about their host species and some have been used as biological controls of pest butterflies like the large white butterfly . When the small cabbage white was accidentally introduced to New Zealand, it had no natural enemies. In order to control it, some pupae that had been parasitised by a chalcid wasp were imported, and natural control was thus regained. Some flies lay their eggs on
6204-419: The epidermis begins to form a new cuticle. At the end of each instar, the larva moults , the old cuticle splits and the new cuticle expands, rapidly hardening and developing pigment. Development of butterfly wing patterns begins by the last larval instar. Caterpillars have short antennae and several simple eyes . The mouthparts are adapted for chewing with powerful mandibles and a pair of maxillae, each with
6298-710: The family Hesperiidae (skippers). Molecular clock estimates suggest that butterflies originated sometime in the Late Cretaceous , but only significantly diversified during the Cenozoic, with one study suggesting a North American origin for the group. The oldest American butterfly is the Late Eocene Prodryas persephone from the Florissant Fossil Beds , approximately 34 million years old. Butterflies are divided into seven families that contain
6392-417: The first experimental evidence for warning coloration in animals. The evolution of aposematism surprised 19th-century naturalists because the probability of its establishment in a population was presumed to be low, since a conspicuous signal suggested a higher chance of predation. Wallace coined the term "warning colours" in an article about animal coloration in 1877. In 1890 Edward Bagnall Poulton renamed
6486-403: The food plant on which their larvae , known as caterpillars , will feed. The caterpillars grow, sometimes very rapidly, and when fully developed, pupate in a chrysalis . When metamorphosis is complete, the pupal skin splits, the adult insect climbs out, expands its wings to dry, and flies off. Some butterflies, especially in the tropics, have several generations in a year, while others have
6580-409: The friction of their overlapping parts. The front two segments have a pair of spiracles which are used in respiration. The abdomen consists of ten segments and contains the gut and genital organs. The front eight segments have spiracles and the terminal segment is modified for reproduction. The male has a pair of clasping organs attached to a ring structure, and during copulation, a tubular structure
6674-544: The grass was growing. The earliest Lepidoptera fossils date to the Triassic - Jurassic boundary, around 200 million years ago. Butterflies evolved from moths, so while the butterflies are monophyletic (forming a single clade ), the moths are not. The oldest known butterfly is Protocoeliades kristenseni from the Palaeocene aged Fur Formation of Denmark, approximately 55 million years old, which belongs to
6768-422: The hindwing upperside, which tend to be separate and surrounded by dark shading in comma, but are larger and tend to run together into a pale band in P. satyrus . Butterfly Butterflies are winged insects from the lepidopteran suborder Rhopalocera , characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The group comprises
6862-1030: The laboratory it recovers within a few generation when allowed to breed freely. During mate selection, adult females do not innately avoid or learn to avoid siblings, implying that such detection may not be critical to reproductive fitness. Inbreeding may persist in B anynana because the probability of encountering close relatives is rare in nature; that is, movement ecology may mask the deleterious effect of inbreeding resulting in relaxation of selection for active inbreeding avoidance behaviors. Butterflies feed primarily on nectar from flowers. Some also derive nourishment from pollen , tree sap, rotting fruit, dung, decaying flesh, and dissolved minerals in wet sand or dirt. Butterflies are important as pollinators for some species of plants. In general, they do not carry as much pollen load as bees , but they are capable of moving pollen over greater distances. Flower constancy has been observed for at least one species of butterfly. Adult butterflies consume only liquids, ingested through
6956-506: The majority of moths which fly by night, are often cryptically coloured (well camouflaged), and either hold their wings flat (touching the surface on which the moth is standing) or fold them closely over their bodies. Some day-flying moths, such as the hummingbird hawk-moth , are exceptions to these rules. Butterfly larvae , caterpillars , have a hard ( sclerotised ) head with strong mandibles used for cutting their food, most often leaves. They have cylindrical bodies, with ten segments to
7050-708: The memory of a bad experience attenuates, the predator may have the experience reinforced through repetition. Aposematic organisms are often slow-moving, as they have little need for speed and agility. Instead, their morphology is frequently tough and resistant to injury, thereby allowing them to escape once the predator is warned off. Aposematic species do not need to hide or stay still as cryptic organisms do, so aposematic individuals benefit from more freedom in exposed areas and can spend more time foraging, allowing them to find more and better quality food. They may make use of conspicuous mating displays, including vocal signals, which may then develop through sexual selection . In
7144-906: The most toxic alkaloids among all living species. Within the same family, there are also cryptic frogs (such as Colostethus and Mannophryne ) that lack these toxic alkaloids. Although these frogs display an extensive array of coloration and toxicity, there is very little genetic difference between the species. Evolution of their conspicuous coloration is correlated to traits such as chemical defense, dietary specialization, acoustic diversification, and increased body mass. Some plants are thought to employ aposematism to warn herbivores of unpalatable chemicals or physical defences such as prickled leaves or thorns. Many insects, such as cinnabar moth caterpillars, acquire toxic chemicals from their host plants. Among mammals, skunks and zorillas advertise their foul-smelling chemical defences with sharply contrasting black-and-white patterns on their fur, while
7238-402: The moth. This is known as Batesian mimicry , after Henry Walter Bates , a British naturalist who studied Amazonian butterflies in the second half of the 19th century. Batesian mimicry is frequency dependent: it is most effective when the ratio of mimic to model is low; otherwise, predators will encounter the mimic too often. A second form of mimicry occurs when two aposematic organisms share
7332-401: The name is ancient, but modern Dutch and German use different words ( vlinder and Schmetterling ) and the common name often varies substantially between otherwise closely related languages. A possible source of the name is the bright yellow male of the brimstone ( Gonepteryx rhamni ); another is that butterflies were on the wing in meadows during the spring and summer butter season while
7426-438: The next day with the suggestion that since some caterpillars "...are protected by a disagreeable taste or odour, it would be a positive advantage to them never to be mistaken for any of the palatable catterpillars [ sic ], because a slight wound such as would be caused by a peck of a bird's bill almost always I believe kills a growing catterpillar. Any gaudy & conspicuous colour therefore, that would plainly distinguish them from
7520-448: The outside of caterpillars and the newly hatched fly larvae bore their way through the skin and feed in a similar way to the parasitoid wasp larvae. Predators of butterflies include ants, spiders, wasps, and birds. Aposematism Aposematism is the advertising by an animal, whether terrestrial or marine, to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make
7614-582: The poisonous beetles they ingest, could be included. It has been proposed that aposematism played a role in human evolution, body odour carrying a warning to predators of large hominins able to defend themselves with weapons. Perhaps the most numerous aposematic vertebrates are the poison dart frogs (family: Dendrobatidae ). These neotropical anuran amphibians exhibit a wide spectrum of coloration and toxicity. Some species in this poison frog family (particularly Dendrobates , Epipedobates , and Phyllobates ) are conspicuously coloured and sequester one of
7708-484: The prey difficult to kill and eat, such as toxicity, venom , foul taste or smell, sharp spines, or aggressive nature. These advertising signals may take the form of conspicuous coloration , sounds , odours , or other perceivable characteristics. Aposematic signals are beneficial for both predator and prey, since both avoid potential harm. The term was coined in 1877 by Edward Bagnall Poulton for Alfred Russel Wallace 's concept of warning coloration . Aposematism
7802-529: The proboscis. They sip water from damp patches for hydration and feed on nectar from flowers, from which they obtain sugars for energy, and sodium and other minerals vital for reproduction. Several species of butterflies need more sodium than that provided by nectar and are attracted by sodium in salt; they sometimes land on people, attracted by the salt in human sweat. Some butterflies also visit dung and scavenge rotting fruit or carcasses to obtain minerals and nutrients. In many species, this mud-puddling behaviour
7896-563: The protection, while many species have bluffing deimatic displays which may startle a predator long enough to enable an otherwise undefended prey to escape. The term aposematism was coined by the English zoologist Edward Bagnall Poulton in his 1890 book The Colours of Animals . He based the term on the Ancient Greek words ἀπό apo 'away' and σῆμα sēma 'sign', referring to signs that warn other animals away. The function of aposematism
7990-431: The quest for a suitable pupation site, often the underside of a leaf or other concealed location. There it spins a button of silk which it uses to fasten its body to the surface and moults for a final time. While some caterpillars spin a cocoon to protect the pupa, most species do not. The naked pupa, often known as a chrysalis, usually hangs head down from the cremaster, a spiny pad at the posterior end, but in some species
8084-550: The same anti-predator adaptation and non-deceptively mimic each other, to the benefit of both species, since fewer individuals of either species need to be attacked for predators to learn to avoid both of them. This form of mimicry is known as Müllerian mimicry , after Fritz Müller , a German naturalist who studied the phenomenon in the Amazon in the late 19th century. Many species of bee and wasp that occur together are Müllerian mimics. Their similar coloration teaches predators that
8178-506: The same few individuals, the predator learning process would result in a stronger warning signal for surviving kin, resulting in higher inclusive fitness for the dead or injured individuals through kin selection . A theory for the evolution of aposematism posits that it arises by reciprocal selection between predators and prey, where distinctive features in prey, which could be visual or chemical, are selected by non-discriminating predators, and where, concurrently, avoidance of distinctive prey
8272-457: The signal's meaning. All of these results contradict the idea that novel, brightly coloured individuals would be more likely to be eaten or attacked by predators. Other explanations are possible. Predators might innately fear unfamiliar forms ( neophobia ) long enough for them to become established, but this is likely to be only temporary. Alternatively, prey animals might be sufficiently gregarious to form clusters tight enough to enhance
8366-407: The similarly-patterned badger and honey badger advertise their sharp claws, powerful jaws, and aggressive natures. Some brightly coloured birds such as passerines with contrasting patterns may also be aposematic, at least in females; but since male birds are often brightly coloured through sexual selection , and their coloration is not correlated with edibility, it is unclear whether aposematism
8460-510: The slugs of the family Phyllidiidae from Indo-Pacific coral reefs are aposematically coloured. Müllerian mimicry has been implicated in the coloration of some Mediterranean nudibranchs, all of which derive defensive chemicals from their sponge diet. The crown-of-thorns starfish , like other starfish such as Metrodira subulata , has conspicuous coloration and conspicuous long, sharp spines, as well as cytolytic saponins , chemicals which could function as an effective defence; this evidence
8554-768: The trait eliminated before predators learn to avoid it. If warning coloration puts the first few individuals at such a strong disadvantage, it would never last in the species long enough to become beneficial. There is evidence for explanations involving dietary conservatism , in which predators avoid new prey because it is an unknown quantity; this is a long-lasting effect. Dietary conservatism has been demonstrated experimentally in some species of birds and fish. Further, birds recall and avoid objects that are both conspicuous and foul-tasting longer than objects that are equally foul-tasting but cryptically coloured. This suggests that Wallace's original view, that warning coloration helped to teach predators to avoid prey thus coloured,
8648-438: The warning signal. If the species was already unpalatable, predators might learn to avoid the cluster, protecting gregarious individuals with the new aposematic trait. Gregariousness would assist predators to learn to avoid unpalatable, gregarious prey. Aposematism could also be favoured in dense populations even if these are not gregarious. Another possibility is that a gene for aposematism might be recessive and located on
8742-445: The warning. Among mammals, predators can be dissuaded when a smaller animal is aggressive and able to defend itself, as for example in honey badgers . Aposematism is widespread in insects, but less so in vertebrates , being mostly confined to a smaller number of reptile , amphibian , and fish species, and some foul-smelling or aggressive mammals . Pitohuis , red and black birds whose toxic feathers and skin apparently comes from
8836-436: Was correct. However, some birds (inexperienced starlings and domestic chicks) also innately avoid conspicuously coloured objects, as demonstrated using mealworms painted yellow and black to resemble wasps, with dull green controls. This implies that warning coloration works at least in part by stimulating the evolution of predators to encode the meaning of the warning signal, rather than by requiring each new generation to learn
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