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110-471: Pareronia ceylanica , the dark wanderer , is a medium-sized butterfly of the family Pieridae , that is, the yellows and whites. It is found in Sri Lanka and India . The dark wanderer resembles Pareronia valeria in colour and in the disposition of the markings, but differs as follows: Male upperside: ground colour a deeper blue. Forewing: the terminal black border much broader generally, entirely without
220-631: 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
330-438: A holoparasite such as Cuscuta derives all of its nutrients from another plant. Parasitic plants make up about one per cent of angiosperms and are in almost every biome in the world. All these plants have modified roots, haustoria , which penetrate the host plants, connecting them to the conductive system—either the xylem , the phloem , or both. This provides them with the ability to extract water and nutrients from
440-493: 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
550-570: 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
660-511: A cause of gastroenteritis , is spread by the fecal–oral route from animals, or by eating insufficiently cooked poultry , or by contaminated water. Haemophilus influenzae , an agent of bacterial meningitis and respiratory tract infections such as influenza and bronchitis , is transmitted by droplet contact. Treponema pallidum , the cause of syphilis , is spread by sexual activity . Viruses are obligate intracellular parasites, characterised by extremely limited biological function, to
770-417: 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
880-627: A faster rate than their hosts. Classic examples include interactions between vertebrate hosts and tapeworms , flukes , and those between the malaria-causing Plasmodium species, and fleas . Parasites reduce host fitness by general or specialised pathology , that ranges from parasitic castration to modification of host behaviour . Parasites increase their own fitness by exploiting hosts for resources necessary for their survival, in particular by feeding on them and by using intermediate (secondary) hosts to assist in their transmission from one definitive (primary) host to another. Although parasitism
990-571: 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
1100-444: 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
1210-606: 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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#17328731092201320-595: A hormone or by diverting nutrients. For example, the trematode Zoogonus lasius , whose sporocysts lack mouths, castrates the intertidal marine snail Tritia obsoleta chemically, developing in its gonad and killing its reproductive cells. Directly transmitted parasites, not requiring a vector to reach their hosts, include such parasites of terrestrial vertebrates as lice and mites; marine parasites such as copepods and cyamid amphipods; monogeneans ; and many species of nematodes, fungi, protozoans, bacteria, and viruses. Whether endoparasites or ectoparasites, each has
1430-503: 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
1540-416: 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
1650-547: 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
1760-448: 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
1870-836: A parasite employs to identify and approach a potential host are known as "host cues". Such cues can include, for example, vibration, exhaled carbon dioxide , skin odours, visual and heat signatures, and moisture. Parasitic plants can use, for example, light, host physiochemistry, and volatiles to recognize potential hosts. There are six major parasitic strategies , namely parasitic castration ; directly transmitted parasitism; trophically -transmitted parasitism; vector -transmitted parasitism; parasitoidism ; and micropredation. These apply to parasites whose hosts are plants as well as animals. These strategies represent adaptive peaks ; intermediate strategies are possible, but organisms in many different groups have consistently converged on these six, which are evolutionarily stable. A perspective on
1980-618: A parasitic alien species. First used in English in 1539, the word parasite comes from the Medieval French parasite , from the Latinised form parasitus , from Ancient Greek παράσιτος (parasitos) 'one who eats at the table of another' in turn from παρά (para) 'beside, by' and σῖτος (sitos) 'wheat, food'. The related term parasitism appears in English from 1611. Parasitism
2090-432: A phenomenon termed the biotrophy-necrotrophy switch . Pathogenic fungi are well-known causative agents of diseases on animals as well as humans. Fungal infections ( mycosis ) are estimated to kill 1.6 million people each year. One example of a potent fungal animal pathogen are Microsporidia - obligate intracellular parasitic fungi that largely affect insects, but may also affect vertebrates including humans, causing
2200-533: A predator, the European sparrowhawk , giving her time to lay her eggs in the host's nest unobserved. Host species often combat parasitic egg mimicry through egg polymorphism , having two or more egg phenotypes within a single population of a species. Multiple phenotypes in host eggs decrease the probability of a parasitic species accurately "matching" their eggs to host eggs. In kleptoparasitism (from Greek κλέπτης ( kleptēs ), "thief"), parasites steal food gathered by
2310-646: 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
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#17328731092202420-412: 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
2530-435: 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
2640-573: A single host-species. Within that species, most individuals are free or almost free of parasites, while a minority carry a large number of parasites; this is known as an aggregated distribution . Trophically -transmitted parasites are transmitted by being eaten by a host. They include trematodes (all except schistosomes ), cestodes , acanthocephalans , pentastomids , many roundworms , and many protozoa such as Toxoplasma . They have complex life cycles involving hosts of two or more species. In their juvenile stages they infect and often encyst in
2750-430: A suitable fungus soon after germinating. Parasitic fungi derive some or all of their nutritional requirements from plants, other fungi, or animals. Plant pathogenic fungi are classified into three categories depending on their mode of nutrition: biotrophs, hemibiotrophs and necrotrophs. Biotrophic fungi derive nutrients from living plant cells, and during the course of infection they colonise their plant host in such
2860-410: 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
2970-528: 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
3080-470: 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
3190-486: 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
3300-1047: A variety of methods to infect animal hosts, including physical contact, the fecal–oral route , free-living infectious stages, and vectors, suiting their differing hosts, life cycles, and ecological contexts. Examples to illustrate some of the many possible combinations are given in the table. social behaviour (grooming) Among the many variations on parasitic strategies are hyperparasitism, social parasitism, brood parasitism, kleptoparasitism, sexual parasitism, and adelphoparasitism. Hyperparasites feed on another parasite, as exemplified by protozoa living in helminth parasites, or facultative or obligate parasitoids whose hosts are either conventional parasites or parasitoids. Levels of parasitism beyond secondary also occur, especially among facultative parasitoids. In oak gall systems, there can be up to five levels of parasitism. Hyperparasites can control their hosts' populations, and are used for this purpose in agriculture and to some extent in medicine . The controlling effects can be seen in
3410-469: A variety of routes. To give a few examples, Bacillus anthracis , the cause of anthrax , is spread by contact with infected domestic animals ; its spores , which can survive for years outside the body, can enter a host through an abrasion or may be inhaled. Borrelia , the cause of Lyme disease and relapsing fever , is transmitted by vectors, ticks of the genus Ixodes , from the diseases' reservoirs in animals such as deer . Campylobacter jejuni ,
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3520-437: 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
3630-474: A way as to keep it alive for a maximally long time. One well-known example of a biotrophic pathogen is Ustilago maydis , causative agent of the corn smut disease. Necrotrophic pathogens on the other hand, kill host cells and feed saprophytically , an example being the root-colonising honey fungi in the genus Armillaria . Hemibiotrophic pathogens begin their colonising their hosts as biotrophs, and subsequently killing off host cells and feeding as necrotrophs,
3740-422: 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
3850-462: Is adapted structurally to this way of life. The entomologist E. O. Wilson characterised parasites as "predators that eat prey in units of less than one". Parasites include single-celled protozoans such as the agents of malaria , sleeping sickness , and amoebic dysentery ; animals such as hookworms , lice , mosquitoes , and vampire bats ; fungi such as honey fungus and the agents of ringworm ; and plants such as mistletoe , dodder , and
3960-563: Is a kind of symbiosis , a close and persistent long-term biological interaction between a parasite and its host. Unlike saprotrophs , parasites feed on living hosts, though some parasitic fungi, for instance, may continue to feed on hosts they have killed. Unlike commensalism and mutualism , the parasitic relationship harms the host, either feeding on it or, as in the case of intestinal parasites, consuming some of its food. Because parasites interact with other species, they can readily act as vectors of pathogens, causing disease . Predation
4070-450: 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
4180-675: Is aggregated. Coinfection by multiple parasites is common. Autoinfection , where (by exception) the whole of the parasite's life cycle takes place in a single primary host, can sometimes occur in helminths such as Strongyloides stercoralis . Vector-transmitted parasites rely on a third party, an intermediate host, where the parasite does not reproduce sexually, to carry them from one definitive host to another. These parasites are microorganisms, namely protozoa , bacteria , or viruses , often intracellular pathogens (disease-causers). Their vectors are mostly hematophagic arthropods such as fleas, lice, ticks, and mosquitoes. For example,
4290-443: Is by definition not a symbiosis, as the interaction is brief, but the entomologist E. O. Wilson has characterised parasites as "predators that eat prey in units of less than one". Within that scope are many possible strategies. Taxonomists classify parasites in a variety of overlapping schemes, based on their interactions with their hosts and on their life cycles , which can be complex. An obligate parasite depends completely on
4400-588: Is clothed with very minute hairs. Colour green, with a lateral row of conspicuous white spots from the 5th to the 12th segment and rows of smaller spots on the back. Food-plant, Capparis heyneana ." "Suspended by the tail and by a very long band ... thoracic portion bent back almost at right angles to the abdominal; head produced into a very long sharp snout; wing-cases forming a keel nearly half an inch in depth, and so thin as to be almost transparent. Colour uniform pale watery green." (Davidson & Aitken.) Butterfly Butterflies are winged insects from
4510-458: 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
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4620-403: 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
4730-410: 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,
4840-645: Is found in the ant Tetramorium inquilinum , an obligate parasite which lives exclusively on the backs of other Tetramorium ants. A mechanism for the evolution of social parasitism was first proposed by Carlo Emery in 1909. Now known as " Emery's rule ", it states that social parasites tend to be closely related to their hosts, often being in the same genus. Intraspecific social parasitism occurs in parasitic nursing, where some individual young take milk from unrelated females. In wedge-capped capuchins , higher ranking females sometimes take milk from low ranking females without any reciprocation. In brood parasitism ,
4950-451: 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
5060-563: 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
5170-585: Is often unambiguous, it is part of a spectrum of interactions between species , grading via parasitoidism into predation, through evolution into mutualism , and in some fungi, shading into being saprophytic . Human knowledge of parasites such as roundworms and tapeworms dates back to ancient Egypt , Greece , and Rome . In early modern times, Antonie van Leeuwenhoek observed Giardia lamblia with his microscope in 1681, while Francesco Redi described internal and external parasites including sheep liver fluke and ticks . Modern parasitology developed in
5280-465: 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
5390-419: Is seen in some species of anglerfish , such as Ceratias holboelli , where the males are reduced to tiny sexual parasites , wholly dependent on females of their own species for survival, permanently attached below the female's body, and unable to fend for themselves. The female nourishes the male and protects him from predators, while the male gives nothing back except the sperm that the female needs to produce
5500-692: 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
5610-405: 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
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#17328731092205720-469: Is then carried to a nest, sometimes alongside other prey if it is not large enough to support a parasitoid throughout its development. An egg is laid on top of the prey and the nest is then sealed. The parasitoid develops rapidly through its larval and pupal stages, feeding on the provisions left for it. Koinobiont parasitoids, which include flies as well as wasps, lay their eggs inside young hosts, usually larvae. These are allowed to go on growing, so
5830-455: 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 a chrysalis . When metamorphosis is complete, the pupal skin splits,
5940-473: 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
6050-469: 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
6160-452: 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
6270-437: The broomrapes . There are six major parasitic strategies of exploitation of animal hosts, namely parasitic castration , directly transmitted parasitism (by contact), trophically-transmitted parasitism (by being eaten), vector-transmitted parasitism, parasitoidism , and micropredation. One major axis of classification concerns invasiveness: an endoparasite lives inside the host's body; an ectoparasite lives outside, on
6380-530: 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 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
6490-402: 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
6600-600: 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
6710-505: The snubnosed eel is probably a facultative endoparasite (i.e., it is semiparasitic) that opportunistically burrows into and eats sick and dying fish. Plant-eating insects such as scale insects , aphids , and caterpillars closely resemble ectoparasites, attacking much larger plants; they serve as vectors of bacteria, fungi and viruses which cause plant diseases . As female scale insects cannot move, they are obligate parasites, permanently attached to their hosts. The sensory inputs that
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#17328731092206820-504: The 19th century. In human culture, parasitism has negative connotations. These were exploited to satirical effect in Jonathan Swift 's 1733 poem "On Poetry: A Rhapsody", comparing poets to hyperparasitical "vermin". In fiction, Bram Stoker 's 1897 Gothic horror novel Dracula and its many later adaptations featured a blood-drinking parasite. Ridley Scott 's 1979 film Alien was one of many works of science fiction to feature
6930-535: 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
7040-430: 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
7150-405: 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
7260-502: 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
7370-433: 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
7480-467: 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
7590-696: The air or soil given off by host shoots or roots , respectively. About 4,500 species of parasitic plant in approximately 20 families of flowering plants are known. Species within the Orobanchaceae (broomrapes) are among the most economically destructive of all plants. Species of Striga (witchweeds) are estimated to cost billions of dollars a year in crop yield loss, infesting over 50 million hectares of cultivated land within Sub-Saharan Africa alone. Striga infects both grasses and grains, including corn , rice , and sorghum , which are among
7700-434: 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
7810-658: 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
7920-864: The deer tick Ixodes scapularis acts as a vector for diseases including Lyme disease , babesiosis , and anaplasmosis . Protozoan endoparasites, such as the malarial parasites in the genus Plasmodium and sleeping-sickness parasites in the genus Trypanosoma , have infective stages in the host's blood which are transported to new hosts by biting insects. Parasitoids are insects which sooner or later kill their hosts, placing their relationship close to predation. Most parasitoids are parasitoid wasps or other hymenopterans ; others include dipterans such as phorid flies . They can be divided into two groups, idiobionts and koinobionts, differing in their treatment of their hosts. Idiobiont parasitoids sting their often-large prey on capture, either killing them outright or paralysing them immediately. The immobilised prey
8030-441: 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
8140-508: The dusky brownish black of the terminal margin as to be very indistinct and blurred. Antennae, head, thorax and abdomen in both sexes much as in P. valeria hippia . This species lives in southern India, the Western Ghats , the southern Andaman Islands and Sri Lanka . "Long, cylindrical or slightly depressed and tapering slightly from the head, which is large, to the tail which ends in two short strong spines clothed with bristles. The body
8250-409: The energy that would have gone into reproduction into host and parasite growth, sometimes causing gigantism in the host. The host's other systems remain intact, allowing it to survive and to sustain the parasite. Parasitic crustaceans such as those in the specialised barnacle genus Sacculina specifically cause damage to the gonads of their many species of host crabs . In the case of Sacculina ,
8360-419: 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
8470-491: The evolutionary options can be gained by considering four key questions: the effect on the fitness of a parasite's hosts; the number of hosts they have per life stage; whether the host is prevented from reproducing; and whether the effect depends on intensity (number of parasites per host). From this analysis, the major evolutionary strategies of parasitism emerge, alongside predation. Parasitic castrators partly or completely destroy their host's ability to reproduce, diverting
8580-444: The family Cuculidae , over 40% of cuckoo species are obligate brood parasites, while others are either facultative brood parasites or provide parental care. The eggs of some brood parasites mimic those of their hosts, while some cowbird eggs have tough shells, making them hard for the hosts to kill by piercing, both mechanisms implying selection by the hosts against parasitic eggs. The adult female European cuckoo further mimics
8690-710: 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
8800-449: The female of P. valeria , but on the upperside the outer black margins beyond the discal markings on both forewings and hindwings are proportionately much broader, the transverse subterminal series of spots that crosses the wing is further from the terminal edge. On the underside the terminal black borders are broader and darker, the subterminal series of spots on apex of the forewing and on the hindwing absent or so very thickly overlaid with
8910-409: 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
9020-544: 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
9130-424: The host and parasitoid develop together for an extended period, ending when the parasitoids emerge as adults, leaving the prey dead, eaten from inside. Some koinobionts regulate their host's development, for example preventing it from pupating or making it moult whenever the parasitoid is ready to moult. They may do this by producing hormones that mimic the host's moulting hormones ( ecdysteroids ), or by regulating
9240-442: The host to complete its life cycle, while a facultative parasite does not. Parasite life cycles involving only one host are called "direct"; those with a definitive host (where the parasite reproduces sexually) and at least one intermediate host are called "indirect". An endoparasite lives inside the host's body; an ectoparasite lives outside, on the host's surface. Mesoparasites—like some copepods , for example—enter an opening in
9350-422: The host's body and remain partly embedded there. Some parasites can be generalists, feeding on a wide range of hosts, but many parasites, and the majority of protozoans and helminths that parasitise animals, are specialists and extremely host-specific. An early basic, functional division of parasites distinguished microparasites and macroparasites. These each had a mathematical model assigned in order to analyse
9460-656: The host's endocrine system. A micropredator attacks more than one host, reducing each host's fitness by at least a small amount, and is only in contact with any one host intermittently. This behavior makes micropredators suitable as vectors, as they can pass smaller parasites from one host to another. Most micropredators are hematophagic , feeding on blood. They include annelids such as leeches , crustaceans such as branchiurans and gnathiid isopods, various dipterans such as mosquitoes and tsetse flies , other arthropods such as fleas and ticks, vertebrates such as lampreys , and mammals such as vampire bats . Parasites use
9570-402: The host's surface. Like predation, parasitism is a type of consumer–resource interaction , but unlike predators , parasites, with the exception of parasitoids, are much smaller than their hosts, do not kill them, and often live in or on their hosts for an extended period. Parasites of animals are highly specialised , each parasite species living on one given animal species, and reproduce at
9680-425: The host. A parasitic plant is classified depending on where it latches onto the host, either the stem or the root, and the amount of nutrients it requires. Since holoparasites have no chlorophyll and therefore cannot make food for themselves by photosynthesis , they are always obligate parasites, deriving all their food from their hosts. Some parasitic plants can locate their host plants by detecting chemicals in
9790-477: The host. The parasitism is often on close relatives, whether within the same species or between species in the same genus or family. For instance, the many lineages of cuckoo bees lay their eggs in the nest cells of other bees in the same family. Kleptoparasitism is uncommon generally but conspicuous in birds; some such as skuas are specialised in pirating food from other seabirds, relentlessly chasing them down until they disgorge their catch. A unique approach
9900-423: The hosts suffer increased parental investment and energy expenditure to feed parasitic young, which are commonly larger than host young. The growth rate of host nestlings is slowed, reducing the host's fitness. Brood parasites include birds in different families such as cowbirds , whydahs , cuckoos , and black-headed ducks . These do not build nests of their own, but leave their eggs in nests of other species . In
10010-472: The intermediate host. When the intermediate-host animal is eaten by a predator, the definitive host, the parasite survives the digestion process and matures into an adult; some live as intestinal parasites . Many trophically transmitted parasites modify the behaviour of their intermediate hosts, increasing their chances of being eaten by a predator. As with directly transmitted parasites, the distribution of trophically transmitted parasites among host individuals
10120-471: The intestinal infection microsporidiosis . Protozoa such as Plasmodium , Trypanosoma , and Entamoeba are endoparasitic. They cause serious diseases in vertebrates including humans—in these examples, malaria, sleeping sickness, and amoebic dysentery —and have complex life cycles. Many bacteria are parasitic, though they are more generally thought of as pathogens causing disease. Parasitic bacteria are extremely diverse, and infect their hosts by
10230-1030: 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
10340-441: The large blue butterfly, Phengaris arion , its larvae employing ant mimicry to parasitise certain ants, Bombus bohemicus , a bumblebee which invades the hives of other bees and takes over reproduction while their young are raised by host workers, and Melipona scutellaris , a eusocial bee whose virgin queens escape killer workers and invade another colony without a queen. An extreme example of interspecific social parasitism
10450-567: The largest group is the parasitoid wasps in the Hymenoptera. The phyla and classes with the largest numbers of parasitic species are listed in the table. Numbers are conservative minimum estimates. The columns for Endo- and Ecto-parasitism refer to the definitive host, as documented in the Vertebrate and Invertebrate columns. A hemiparasite or partial parasite such as mistletoe derives some of its nutrients from another living plant, whereas
10560-506: 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
10670-537: The marine worm Bonellia viridis has a similar reproductive strategy, although the larvae are planktonic. Examples of the major variant strategies are illustrated. Parasitism has an extremely wide taxonomic range, including animals, plants, fungi, protozoans, bacteria, and viruses. Parasitism is widespread in the animal kingdom, and has evolved independently from free-living forms hundreds of times. Many types of helminth including flukes and cestodes have complete life cycles involving two or more hosts. By far
10780-401: 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
10890-407: The next generation. Adelphoparasitism, (from Greek ἀδελφός ( adelphós ), brother ), also known as sibling-parasitism, occurs where the host species is closely related to the parasite, often in the same family or genus. In the citrus blackfly parasitoid, Encarsia perplexa , unmated females may lay haploid eggs in the fully developed larvae of their own species, producing male offspring, while
11000-407: 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. Parasite Parasitism is a close relationship between species , where one organism, the parasite, lives on or inside another organism, the host , causing it some harm, and
11110-417: The point where, while they are evidently able to infect all other organisms from bacteria and archaea to animals, plants and fungi, it is unclear whether they can themselves be described as living. They can be either RNA or DNA viruses consisting of a single or double strand of genetic material ( RNA or DNA , respectively), covered in a protein coat and sometimes a lipid envelope. They thus lack all
11220-535: The population movements of the host–parasite groupings. The microorganisms and viruses that can reproduce and complete their life cycle within the host are known as microparasites. Macroparasites are the multicellular organisms that reproduce and complete their life cycle outside of the host or on the host's body. Much of the thinking on types of parasitism has focused on terrestrial animal parasites of animals, such as helminths. Those in other environments and with other hosts often have analogous strategies. For example,
11330-529: 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
11440-431: 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
11550-466: The testes of over two-thirds of their crab hosts degenerate sufficiently for these male crabs to develop female secondary sex characteristics such as broader abdomens, smaller claws and egg-grasping appendages. Various species of helminth castrate their hosts (such as insects and snails). This may happen directly, whether mechanically by feeding on their gonads, or by secreting a chemical that destroys reproductive cells; or indirectly, whether by secreting
11660-422: The transverse sub-terminal series of bluish-white spots; in a few specimens there are one or two of these spots present, but nothing like the series so conspicuous in P. valeria hippia . Hindwing: the terminal black border very broad, narrowing slightly but distinctly towards the tornal angle. Proportionately this border is even broader than in the forewing. Underside: as in P. valeria . The female closely resembles
11770-403: The tropics, however effectively cheat by taking carbon from a fungus rather than exchanging it for minerals. They have much reduced roots, as they do not need to absorb water from the soil; their stems are slender with few vascular bundles , and their leaves are reduced to small scales, as they do not photosynthesize. Their seeds are small and numerous, so they appear to rely on being infected by
11880-506: The usual machinery of the cell such as enzymes , relying entirely on the host cell's ability to replicate DNA and synthesise proteins. Most viruses are bacteriophages , infecting bacteria. Parasitism is a major aspect of evolutionary ecology; for example, almost all free-living animals are host to at least one species of parasite. Vertebrates, the best-studied group, are hosts to between 75,000 and 300,000 species of helminths and an uncounted number of parasitic microorganisms. On average,
11990-585: The way that the CHV1 virus helps to control the damage that chestnut blight , Cryphonectria parasitica , does to American chestnut trees, and in the way that bacteriophages can limit bacterial infections. It is likely, though little researched, that most pathogenic microparasites have hyperparasites which may prove widely useful in both agriculture and medicine. Social parasites take advantage of interspecific interactions between members of eusocial animals such as ants , termites , and bumblebees . Examples include
12100-471: The world's most important food crops. Orobanche also threatens a wide range of other important crops, including peas , chickpeas , tomatoes , carrots , and varieties of cabbage . Yield loss from Orobanche can be total; despite extensive research, no method of control has been entirely successful. Many plants and fungi exchange carbon and nutrients in mutualistic mycorrhizal relationships. Some 400 species of myco-heterotrophic plants, mostly in
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