87-407: There are 31 genera and about 600 species: Swallowtail butterflies are large, colorful butterflies in the family Papilionidae , and include over 550 species . Though the majority are tropical , members of the family inhabit every continent except Antarctica . The family includes the largest butterflies in the world, the birdwing butterflies of the genus Ornithoptera . Swallowtails have
174-487: A behavior in which the butterflies' appearance closely resemble that of distasteful species that prevents predation. Swallowtails differ from many animals that practice mimicry . The tiger swallowtail butterfly ( Papilio glaucus ), exhibits a female-limited polymorphism for Batesian mimicry and others, such as the Canadian tiger swallowtail ( Papilio canadensis ) do not display any form of mimicry. Predators include
261-505: 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 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
348-637: 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
435-476: A black-and-white pattern for Batesian mimicry , a black-and-yellow pattern that resembles the males of the species, and a pattern with orange patches that resembles the elderly males of the species. Given that the males of the species, which do not have Batesian mimicry, are preyed upon much more frequently by predators than the females, an ongoing question is why females would exhibit the non-mimetic wing pattern, which would seemingly lower their fitness compared to
522-493: 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
609-570: 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
696-417: 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
783-576: 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
870-444: 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
957-606: 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
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#17327720515341044-508: 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
1131-416: 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
1218-547: 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
1305-452: 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
1392-399: A number of distinctive features; for example, the papilionid caterpillar bears a repugnatorial organ called the osmeterium on its prothorax. The osmeterium normally remains hidden, but when threatened, the larva turns it outward through a transverse dorsal groove by inflating it with fluid. The forked appearance in some of the swallowtails' hindwings, which can be seen when the butterfly
1479-658: 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
1566-412: 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
1653-435: 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
1740-610: A single genus Praepapilio , includes two species of extinct butterflies, each member being described from single fossils found in a middle Eocene deposit in Colorado , United States (Durden and Rose, 1978). A phylogeny of the Papilionidae based on Nazari (2007) is given: (†) Praepapilioninae Baroniinae Parnassiini Zerynthiini Luehdorfiini Leptocircini Teinopalpini Papilionini Troidini It
1827-555: A subfamily of essentially Holarctic butterflies. The vast majority of species, mostly Parnassius , can be found in mountain habitats. Parnassiinines can also be found in other habitats such as "arid deserts ( Hypermnestra ), humid forests ( Luehdorfia ) and even lowland meadows ( Zerynthia )". The tribes recognized in the Parnassiinae are Parnassiini , Zerynthiini , and Luehdorfiini . Tribe Parnassiini contains two genera, Hypermnestra , largely confined to central Asia and
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#17327720515341914-410: 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
2001-541: 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
2088-470: 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
2175-486: 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
2262-437: 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
2349-422: 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
2436-450: 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
2523-458: 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
2610-403: 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
2697-410: 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,
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2784-451: 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
2871-554: Is mimetic, and whether the mimicry is monomorphic or polymorphic. A phenomenon which has received particular attention is female-limited polymorphism, in which only the females of a species are mimetic and polymorphic, often mimicking different, distantly-related aposematic butterflies. This polymorphism is seen in Papilio dardanus , the African swallowtail butterfly, whose females have three different morphs for wing color pattern:
2958-581: 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
3045-611: Is now accepted that the subfamily Papilioninae is monophyletic . The swallowtail butterflies in the nominate tribe Papilionini number about 225 species and studies have been made on their host plant coevolution and phylogeny . Old morphological classifications were also found to be valid in that they formed clusters. Species belonging to the groups that use Rutaceae as host plants formed two groups corresponding to Old World and American taxa. Those that fed on Lauraceae and Magnoliaceae were found to form another cluster which includes both Asian and American taxa. The Parnassinae, like
3132-402: Is resting with its wings spread, gave rise to the common name swallowtail . As for its formal name, Linnaeus chose Papilio for the type genus , as papilio is Latin for "butterfly". For the specific epithets of the genus, Linnaeus applied the names of Greek figures to the swallowtails. The type species : Papilio machaon honored Machaon , one of the sons of Asclepius , mentioned in
3219-465: 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
3306-698: 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
3393-405: 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
3480-680: Is the Siberian Apollo ( Parnassius arcticus ), found in the Arctic Circle in northeastern Yakutia , at altitudes of 1500 meters above sea level. In the Himalayas , various Apollo species such as Parnassius epaphus , have been found at altitudes of 6,000 meters above sea level. The caterpillars of various swallowtail butterfly species feed on a wide range of different plants, most depending on only one of five families: Aristolochiaceae , Annonaceae , Lauraceae , Umbelliferae ( Apiaceae ) and Rutaceae . By eating some of these toxic plants,
3567-574: The Iliad . Further, the species Papilio homerus is named after the Greek poet, Homer . The Mon of the Taira clan of Japan is an Agehachō (swallowtail butterfly). The genera of extant swallowtails are usually classified into three subfamilies , Baroniinae , Parnassiinae , and Papilioninae , the latter two being further divided into tribes . In swallowtails, besides morphological characteristics,
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3654-473: The B. philenor by predators, the Papilio glaucus's background wing color matches that of the B. philenor residing in the same regional area. Studies support this theory; in the southeastern United States, the relative abundance of melanic females has been found to geographically correlate with B. philenor . Only certain subsets of swallowtails practice mimicry . Species differ in whether one or both sexes
3741-479: 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
3828-474: 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
3915-455: 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
4002-426: The birdwing butterflies are particularly sought after and are cultured in butterfly farms to supply collectors. 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 the superfamilies Hedyloidea (moth-butterflies in
4089-402: 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
4176-600: 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
4263-412: The red-winged blackbird , Pennsylvania firefly , five-lined skink , green darner , goldenrod spider , Chinese mantis , fiery searcher , and striped skunk . Not all individuals in some species are identical in appearance. For example, Papilio glaucus (eastern tiger swallowtail), Y-linkage determines whether the females are either wild-type (yellow and black) or melanic (dark melanin replaces
4350-680: 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 the food plant on which their larvae , known as caterpillars , will feed. The caterpillars grow, sometimes very rapidly, and when fully developed, pupate in
4437-540: 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
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#17327720515344524-465: The Batesian females suffered the fewest predators but their fitness was reduced the most by sexual harassment, while the other two faced lower sexual harassment but also lost fitness from predators' attacks. After mating , the male Parnassines produce a glue-like substance that is used to seal the female genital opening and prevent other males from mating. They lay individual eggs on the underside of
4611-464: The Papilionidae, as quoted in Bingham (1905) are as follows: Egg . "Dome-shaped, smooth or obscurely facetted, not as high as wide, somewhat leathery, opaque." ( Doherty .) Larva . Stout, smooth or with a series of fleshy tubercles on the dorsum: sometimes with a raised fleshy protuberance (the so-called hood or crest) on the fourth segment. The second segment has a transverse opening, out of which
4698-721: The Papilioninae, were also believed to be monophyletic based on morphological studies but recent studies based on both morphological and molecular characteristics suggest that this is not the case. Of the Parnassiinae , the genera Parnassius and Hypermnestra were found to be extremely close based on molecular studies and are now considered to be part of the tribe Parnassiini. The two taxa, Archon and Luehdorfia , have been found to be closely related through analysis of nuclear and mitochondrial DNA, and, though they share no morphological similarities, have now been united in
4785-430: 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
4872-405: 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
4959-423: The aberrant genera Parnassius and Hypermnestra ) with all 12 veins present and in addition a short internal vein, vein 1 a, that invariably terminates on the dorsal margin. Stages of development of a papilionid, the giant swallowtail ( Papilio cresphontes ) The key characteristics that differentiate the Papilionidae from the other butterfly families are: Swallowtail butterflies practice Batesian mimicry ,
5046-502: 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
5133-467: 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
5220-434: 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
5307-405: The archetypical papilionid food plant, the lowland vine Aristolochia . Zerynthiini comprises four genera – Sericinus , Bhutanitis , Zerynthia and Allancastria . Subfamily: Parnassiinae . The tribes recognized in the Papilioninae are Leptocircini , Teinopalpini, Troidini , and Papilionini . Subfamily: Papilioninae . An additional subfamily, Praepapilioninae , consisting of
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#17327720515345394-658: 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
5481-644: The case of most species of Parnassius . The majority of swallowtail species and the greatest diversity are found in the tropics and subtropical regions between 20°N and 20°S, particularly Southeast Asia, and between 20°N and 40°N in East Asia. Only 12 species are found in Europe and only one species, Papilio machaon is found in the British Isles . North America has 40 species, including several tropical species and Parnassius . The northernmost swallowtail
5568-486: The caterpillars sequester aristolochic acid which renders both the caterpillars and the butterflies of some of these as toxic, thus protecting them from predators. Similarly, the Parnassius smintheus sequesters sarmentosin from its host plant Sedum lanceolatum for protection from predators. Swallowtail tribes Zerynthiini (Parnassiinae), Luehdorfiini (Parnassiinae) and Troidini (Papilioninae), almost exclusively use
5655-524: The choice of food plants and ecological lifestyle reflect phylogeny and classification. The Baroniinae are a monotypic subfamily, restricted to a very small region in Mexico and are considered to be the most basal of the subfamilies. Baronia brevicornis is considered to be a relict species, and shares features with a fossil taxon Praepapilio . Baronia is unique among papilionidae' as having an Acacia species (family Leguminosae ) as its food plant. Subfamily: Baroniene . The Parnassiinae are
5742-441: 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
5829-419: 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
5916-406: The family Aristolochiaceae as their host plants. For example, the eastern black swallowtail's ( Papilio polyxenes ) main host plant in the wild is Queen Anne's lace , but they also eat garden plants in the carrot family, including carrots, parsley, dill, and fennel. Adult swallowtails sip nectar, but also mud and sometimes manure. The detailed descriptions of morphological characteristics of
6003-741: 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
6090-409: 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
6177-492: The genus Parnassius (the Apollos), a distinctive group of many species, all of which are alpine and capable of living at high altitudes. Most Parnassius have two small reddish spots on their hindwings . The tribe Luehdorfiini contains the genera Archon of Asia minor and the genus Luehdorfia of China and Japan. These two tribes have evolved to change their food plants, while the third tribe, Zerynthiini, has retained
6264-555: 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
6351-1030: 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
6438-408: The larva protrudes at will and an erect, forked, glandular fleshy organ that emits a strong, penetrating, and somewhat unpleasant odor. Pupa . Variable in form but most often curved backwards. It is angulate, with the head truncate or rounded and the back of abdomen is smooth or tuberculate. It is attached by the tail, normally in a perpendicular position, and further secured by a silken girth round
6525-517: The leaves of their food plants. There is no parental investment once the eggs have been laid. The pupae are typically attached to the substrate by the cremaster but with head up held by a silk girdle. The Apollos, however, pupate in debris on the ground and also build a loose cocoon . In the temperate regions, the winters are passed in a pupal diapause stage. Since swallowtail butterflies are large, colorful, and attractive, they have been targeted by butterfly collectors . The largest of these,
6612-506: 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
6699-499: The male avoidance hypothesis, female butterflies disguise themselves in an attempt to evade male harassment, as courtship can be harmful, time-consuming, and attract predators. One study recorded male responses to females of each morphs and found that the males consistently favored the Batesian mimics, then the black and yellow, and then the morph with orange patches. The scientists concluded that frequency-dependent selection did lead to equal success for all three alternative strategies:
6786-426: The middle. In Parnassius , the pupa is placed in a loose silken web between leaves. Imago . Wings extraordinarily variable in shape. Hindwing very frequently has a tail, which may be slender, or broad and spatulate, but is always an extension of the termen at vein 4. In one genus, Armandia , the termen of the hindwing is prolonged into tails at the apices of veins 2 and 3 as well as at vein 4. Forewing (except in
6873-445: The mimicry form. The pipevine swallowtail exhibits Batesian mimicry as well. Several hypotheses for this phenomenon were made, the two noteworthy being the pseudosexual selection hypothesis and the male avoidance hypothesis. In the pseudosexual hypothesis, male butterflies aggressively approached the male -looking females and then mellowed their behavior into sexual behavior when they were close enough to identify them as females. In
6960-401: 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
7047-474: 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. Zerynthia See text See text Zerynthia is a genus of swallowtail butterflies placed in the subfamily Parnassiinae . The genus has a complex history; a multiplicity of names have been applied to its species. Zerynthia consists of
7134-498: The pipevine swallowtail. There are also Papilio glaucus that are not wholly black; several possess an intermediate "sooty" color and are sensitive to temperature . The different polymorphisms (wild-type, melanic, and the 'sooty' intermediate) depend upon the geographical distribution and abundance of its mimic, the Battus philenor , whose wing color varies depending on its geographical location. In order to be successfully confused for
7221-529: 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
7308-431: 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
7395-561: The swallowtails. Recent research suggests that this may not be the case, the Baroniinae being closely related to only the Parnassiinae, and Praepapilio to only the Papilionini and neither taxa being sister to the rest of the swallowtails. As of 2005, 552 extant species have been identified which are distributed across the tropical and temperate regions. Various species inhabit altitudes ranging from sea level to high mountains, as in
7482-462: The tribe Luehdorfiini. The subfamily Baroniinae is represented by the sole representative species Baronia brevicornis . They are unique in the family to use the Fabaceae (Leguminosae) as their larval host plants. The Baronninae and the extinct subfamily Praepapilioninae share many external similarities and are traditionally considered to be the most primitive subfamilies and sister to the rest of
7569-438: The yellow background). This genetic difference stems from the fact that melanism is controlled by a single gene , which controls the level of dopamine in the organism. The enzyme BAS, which assists dopamine in producing the yellow pigmentation , normally found on the wings' background, is suppressed. Without the pigmentation, the butterfly appears mostly black (the melanic form) and is a Batesian mimic of Battus philenor ,
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