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''Papilio xuthus'', the Asian swallowtail, Chinese yellow swallowtail or Xuthus swallowtail, is a yellow-colored, medium to large sized
swallowtail butterfly 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 large ...
found in northeast
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
, northern
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
, southern
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
,
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia, at the junction of the East and South China Seas in the northwestern Pacific Ocean, with the People's Republic of China (PRC) to the northwest, Japan to the nort ...
, the
Korean Peninsula Korea ( ko, 한국, or , ) is a peninsular region in East Asia. Since 1945, it has been divided at or near the 38th parallel, with North Korea (Democratic People's Republic of Korea) comprising its northern half and South Korea (Republic o ...
,
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
(from
Hokkaidō is Japan's second largest island and comprises the largest and northernmost prefecture, making up its own region. The Tsugaru Strait separates Hokkaidō from Honshu; the two islands are connected by the undersea railway Seikan Tunnel. The la ...
to the
Yaeyama Islands The Yaeyama Islands (八重山列島 ''Yaeyama-rettō'', also 八重山諸島 ''Yaeyama-shotō'', Yaeyama: ''Yaima'', Yonaguni: ''Daama'', Okinawan: ''Yeema'', Northern Ryukyuan: ''Yapema'') are an archipelago in the southwest of Okinawa ...
),
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...
and the
Hawaiian Islands The Hawaiian Islands ( haw, Nā Mokupuni o Hawai‘i) are an archipelago of eight major islands, several atolls, and numerous smaller islets in the North Pacific Ocean, extending some from the island of Hawaii in the south to northernmost Kur ...
. It was also recorded in the state of
Arunachal Pradesh Arunachal Pradesh (, ) is a state in Northeastern India. It was formed from the erstwhile North-East Frontier Agency (NEFA) region, and became a state on 20 February 1987. It borders the states of Assam and Nagaland to the south. It shares int ...
,
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
, in 2014. It mates multiple times in its life, leading to an increased genetic diversity in its young. It is preyed upon by a host of organisms, including the tree cricket '' Oecanthus longicauda'', ant ''
Lasius niger ) , official_languages = , languages_type = National languagesPolistes Wasps of the cosmopolitan genus ''Polistes'' (the only genus in the tribe Polistini) are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. Walter Ebeling coined the vernacular name "umbrella ...
'' and ''Trogus mactator''). ''P. xuthus'' utilizes color vision and color constancy while foraging for plants of the family ''
Rutaceae The Rutaceae is a family, commonly known as the rueRUTACEAE
in BoDD – Botanical Derm ...
''. It is common and not threatened. ''Papilio xuthus'' is common in urban, suburban, woods and orange orchards. The flight period is from May to August. Males use both physical and visual cues to attract mates during the
breeding season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
. Females of the species regularly mate with multiple partners. After mating, females use habitat and food quality to determine where they will oviposit their eggs.


Taxonomy

''Papilio xuthus'' is a member of the genus ''
Papilio ''Papilio'' is a genus in the swallowtail butterfly family, Papilionidae, as well as the only representative of the tribe Papilionini. The word ''papilio'' is Latin for butterfly. It includes the common yellow swallowtail (''Papilio machaon''), ...
''. It is of the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Papilionidae 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 larges ...
and order
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
. It was first introduced in
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
in 1971 from
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
or
Guam Guam (; ch, Guåhan ) is an organized, unincorporated territory of the United States in the Micronesia subregion of the western Pacific Ocean. It is the westernmost point and territory of the United States (reckoned from the geographic cent ...
. ''P. xuthus'' makes significant seasonal migrations over 200 km. There are two subspecies of ''P. xuthus'': ''Papilio xuthus koxingus'' and ''Papilio xuthus neoxuthus''.


Description

The Asian swallowtail is a mid-sized, yellow, prominently-tailed butterfly. It has a wingspan of 90 to 110 mm. Its normal color pattern consists of a black pattern on a yellow background. Wing coloration is sexually dimorphic, with females showing broader proximal marginal bands in the hindwing. Blue-iridescent and orange scales separate the black bands on the hindwing. The black bands also run in stripes of varying thickness along the forewing. The young caterpillar
mimics Materialise Mimics is an image processing software for 3D design and modeling, developed by Materialise NV, a Belgian company specialized in additive manufacturing software and technology for medical, dental and additive manufacturing industries ...
bird feces and has a white and brown spot on its head. As they mature, caterpillars develop a light green body color with brown spots.


Wing Coloration

''P. xuthus'' have highly sensitive photoreceptors, or
ommatidia The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ou ...
, that can detect a wide range of wavelengths. This adaptation helps them identify members of their own species. Their distinct coloring and patterning are what sets them apart from other butterflies. Wing color is derived from "scales", and ''P. xuthus'' has scales of yellow (cream), orange, black, and blue from the light each scale absorbs and reflects. Spectrophotometer studies saw that the yellow scales absorbed in the UV to violet range, orange absorbed in the lower half-wavelengths of the visible spectrum, black scales absorbed in most of the visible light range, and blue scales absorbed very little light. These absorbance ranges are due to the pigment molecules contained in each wing scale. Yellow and orange contain papiliochrome II and black contains
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
. The blue scales are unique in that they don't have a pigment molecule, and are instead colored by an optical phenomenon. The interaction of light with the upper and lower lamina of the scale gives them the distinct blue color that we see. Experimental manipulation of wing coloration has been achieved through administration of the acid carboxypeptidase, Molsin. Injection of Molsin into 0-2 day pupae, butterflies had high amounts of black wing scales. These modified subjects had produced high amounts of melanin and papiliochrome II. Injection of Molsin into 3-4 day pupae shows no change in wing coloration. This indicates that ''P. xuthus'' likely has an acid carboxypeptidase of its own that naturally induces production of these pigments, and its action lasts during the first two days of pupation. Administration of a higher amount of acid carboxypeptidase to the pupae at 0–2 days induces an overproduction of pigmentation molecules, which leads to a high number of black scales. ''P. xuthus'' uses wing coloration and patterning to identify insects of the same species. Sexual behaviors are elicited from males by visual identification alone. Adult ''P. xuthus'' males were able to find odorless conspecifics, and while they do not discriminate by sex, they can identify a female once physical contact is made.


Courtship and mating

During the mating season, Asian swallowtail males fly through the
foliage A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
of trees in search of a mate. If a male encounters a female resting on a plant with open and horizontal wings, it approaches her and examines her by contacting his forelegs with the tips of her wings. In addition, males use visual cues to conduct the mating ritual and are particularly attracted to closely spaced yellow patches on female wings. Most females of the Asian swallowtail butterfly mate more than once in their life. This behavior is typical of many other swallowtail butterflies including '' P. glaucus'', '' P. helenus'' and '' P. protenor''.Watanabe, M., and Nozato, K. (1986). Fecundity of the yellow swallowtail butterflies,Papilio xuthus and P. machaon hippocrates, in a wild environment.Zool. Sci. 3: 509–515. By mating multiply, females are able to increase the
genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...
of their offspring. In addition, multiple matings allow females to produce a greater number of eggs, of which there can be up to thirty.


Oviposition

Female butterflies disperse when they are about to
oviposit The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
. They decide on the number of eggs they deposit in different subhabitats by certain criteria such as the quality of the subhabitat, the existing adult population in the area, the quantity of young leaves in the vicinity, the degree of sun exposure (females prefer sunnier areas), and host plant height.


Mortality factors


Predation

Some insects, such as the grasshopper ''Oeanthus longicauda'', are capable of removing Asian swallowtail larvae out of their eggs before they hatch. During the first and second larval
instars An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
, their main predators are
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s (particularly ''
Lasius niger ) , official_languages = , languages_type = National languagesspider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s, and various other bugs. Asian swallowtail caterpillars may also be parasitized by wasps, such as ''Trogus mactator'' and ''Pteromarus puparum'', during development. These wasps ultimately kill the larvae after pupation. In fact, wasps have been observed emerging from the host pupa four or five days after the predicted date of emergence of the host.


Environment

Aside from predation, larval
mortality rate Mortality rate, or death rate, is a measure of the number of deaths (in general, or due to a specific cause) in a particular population, scaled to the size of that population, per unit of time. Mortality rate is typically expressed in units of de ...
is also affected by diseases that appear to correlate with the duration of the rainy season. Long days of rain in autumn lead to the rotting of infected larvae.


Foraging

Asian swallowtail butterflies exhibit
color vision Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different wavelengths (i.e., different spectral power distributions) independently of light intensity. Color perception is a part of ...
which gives them the ability to differentiate objects of contrasting colors irrespective of brightness. They also exhibit color constancy, which allows them to recognize the color of an organism regardless of the background light. Asian swallowtail butterflies show this color constancy even when the background light is yellow and red. This ability is important for
foraging Foraging is searching for wild food resources. It affects an animal's Fitness (biology), fitness because it plays an important role in an animal's ability to survive and reproduce. Optimal foraging theory, Foraging theory is a branch of behaviora ...
, as it allows butterflies to recognize certain flowers in an area irrespective of whether the flower is in the shade or in the sun.


Food plants

The
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e of the species feed on plants of family
Rutaceae The Rutaceae is a family, commonly known as the rueRUTACEAE
in BoDD – Botanical Derm ...
. Recorded species of food plants include: * ''
Phellodendron amurense ''Phellodendron amurense'' is a species of tree in the family Rutaceae, commonly called the Amur cork tree. It is a major source of '' huáng bò'' ( or 黄 檗), one of the 50 fundamental herbs used in traditional Chinese medicine. The Ainu pe ...
'' * ''
Poncirus trifoliata The trifoliate orange, ''Citrus trifoliata'' or ''Poncirus trifoliata'', is a member of the family Rutaceae. Whether the trifoliate oranges should be considered to belong to their own genus, ''Poncirus'', or be included in the genus ''Citrus'' is ...
'' * ''
Zanthoxylum ''Zanthoxylum'' is a genus of about 250 species of deciduous and evergreen trees, shrubs and climbers in the family Rutaceae that are native to warm temperate and subtropical areas worldwide. It is the type genus of the tribe Zanthoxyleae in ...
'' species, such as the Japanese pepper tree ('' Z. piperitum''), '' Z. nitidum'' and the Japanese prickly-ash '' Z. ailanthoides'' * Cultivated ''
Citrus ''Citrus'' is a genus of flowering plant, flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as Orange (fruit), oranges, Lemon, lemons, grapefruits, pomelos, and lim ...
'' species such as
mandarin orange The mandarin orange (''Citrus reticulata''), also known as the mandarin or mandarine, is a small citrus tree fruit. Treated as a distinct species of orange, it is usually eaten plain or in fruit salads. Tangerines are a group of orange-coloured ...
and
yuzu Yuzu (''Citrus junos'', from Japanese or ) is a citrus fruit and plant in the family Rutaceae of East Asian origin. Yuzu has been cultivated mainly in East Asia, though recently also in New Zealand, Australia, Spain, Italy, and France. I ...
* '' Euodia ruaecarpa'' * ''
Ruta graveolens ''Ruta graveolens'', commonly known as rue, common rue or herb-of-grace, is a species of '' Ruta'' grown as an ornamental plant and herb. It is native to the Balkan Peninsula. It is grown throughout the world in gardens, especially for its blui ...
'' (rue) * "
Bauhinia ''Bauhinia'' () is a large genus of flowering plants in the subfamily Cercidoideae and tribe Bauhinieae, in the large flowering plant family Fabaceae, with a pantropical distribution. The genus was named after the Bauhin brothers Gaspard and Jo ...
" species, such as Hong Kong Orchid Tree


Development and population dynamics

Despite its wide distribution, populations of the Asian swallowtail remain at a stable, relatively low, level. There are typically four to five generations per year with the adults of the first generation emerging from mid-April to early May and adults of the second generation emerging in mid-June.Inaba, S. (1937) The ecological observation on the swallowtail butterflyPapilio xuthus L. (II).Acta Zooecologica 1, (2): 17–29. ''P. xuthus'' will lay single eggs on leaves of host plants, allowing hatched
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e to feed on ''
Poncirus trifoliata The trifoliate orange, ''Citrus trifoliata'' or ''Poncirus trifoliata'', is a member of the family Rutaceae. Whether the trifoliate oranges should be considered to belong to their own genus, ''Poncirus'', or be included in the genus ''Citrus'' is ...
'', ''
Zanthoxylum ailanthoides ''Zanthoxylum ailanthoides'', also called ailanthus-like prickly ash, (, lit. " Ailanthus-leaved pepper",; the name is obviously adaptation of latin ailanthoides "ailanthus-like" , lit. " Yue pepper",, p.503 食茱萸 ''shi zhu yu'',, 「澳名� ...
'' and various citrus species. The larval stage lasts for approximately three to five weeks while the
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
l stage is around two weeks. According to a study, the population fluctuation of the Asian swallowtail butterfly has been surprisingly stable compared to that of various other insects. The stability of the population appeared to be regulated by the population dynamics of egg the parasitoids, ''
Trichogramma ''Trichogramma'' is a genus of minute polyphagous wasps that are endoparasitoids of insect eggs. ''Trichogramma'' is one of around 80 genera from the family Trichogrammatidae, with over 200 species worldwide.Consoli FL, Parra JRP, Zucchi RA (2010 ...
'', and the pupal parasitoids, ''Pteromalus puparum'', which maintained shorter life spans and generation times than the eggs of the Asian swallowtail. When the butterfly population reaches a high density, because of the shorter generational turn around of the parasites, intergeneration responses of the parasites are faster than those of the butterflies, resulting in a responding increased growth of the parasites. This creates an evolutionary stable system to keep the population of both the parasites and the host in check. It was also found that among the subpopulations studied, exchanges between each subpopulation helped to maintain the stability of the entire population as a whole.


Gallery

File:Egg of Papilio xuthus.JPG, Egg File:Papilio xuthus Larva 2011-10-15.jpg, Caterpillar File:Papilio xuthus Pupa 2011-07-16.jpg, Pupa File:Papilio xuthus and HIGANBANA Lycoris radiata.jpg, On ''
Lycoris radiata ''Lycoris radiata'', known as the red spider lily, red magic lily, corpse flower, or equinox flower, is a plant in the amaryllis family, Amaryllidaceae, subfamily Amaryllidoideae. Originally from China, Korea, and Nepal, it was introduced into J ...
''


See also

*
Papilionidae 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 larges ...


References

*Harry, J. L. 1994. ''Papilio xuthus'' in Hawaii: A Photo Life History (Lepidoptera: Papilionidae). ''Tropical Lepidoptera'' 5(1): 6–7
pdf
{{Taxonbar, from=Q132092
xuthus In Greek mythology, Xuthus (; grc, Ξοῦθος ''Xouthos'') was a Phthian prince who later became a king of Peloponnesus. He was the founder (through his sons) of the Achaean and Ionian nations. Etymology According to the author, Robert ...
Butterflies of Asia Butterflies of Oceania Butterflies of Indochina Butterflies of Japan Insects of Hawaii Taxa named by Carl Linnaeus Articles containing video clips Butterflies described in 1767