''Papilio protenor'', the spangle, is a butterfly found in South East Asia belonging to the swallowtail family.

Morphology

Male

''Upperside'': velvety indigo-blue black, duller on the forewing than on the hindwing. Forewing with pale adnervular streaks broadened along the terminal margin and extended well into the cell. Hindwing: a broad pale yellowish-white sub costal streak; interspaces 4 to 6 irrorated (sprinkled) with bluish scales; tornal angle marked with red. ''Underside'': forewing dull black; adnervular streaks distinctly grey and much broader than on the upperside. Hindwing: ground colour as on the upperside, a large irregularly-shaped patch at the tornal angle that extends into interspace 2, and subterminal lunules in interspaces 2, 6, and 7 dull pinkish-red, cell irrorated more-or-less with a sprinkling of blue scales; the tornal patch with a black, outwardly blue-edged, round medial spot, and interspaces 4 and 5 with subterminal irrorations of blue scales. Antennae black; head, thorax and abdomen dark brownish-black.

Female

Similar to male. ''Upperside'': ground colour deep brownish-black; adnervular streaks on forewing yellowish; irroration of blue scales on outer portions of hindwing more dense; no white subcostal streak; red patch at tornal angle large with an oval medial black spot; another similar black spot subterminally in interspace 2 posteriorly bordered by a crescent-shaped red mark. ''Underside'': similar to that in the male, differs in the adnervular streaks on the forewing that are broader and paler; on the hindwing the tornal red patch is paler and larger, and is extended broadly anteriorly and outwards towards the

termen Termen is a Municipalities of Switzerland, municipality in the district of Brig (district), Brig in the Cantons of Switzerland, canton of Valais in Switzerland. History Termen is first mentioned in 1201 as ''Terman''. Geography Termen has an ar ...

into interspace 2; in the latter it coalesces with a broad subterminal black-centred red

ocellus A simple eye or ocellus (sometimes called a pigment pit) is a form of eye or an optical arrangement which has a single lens without the sort of elaborate retina that occurs in most vertebrates. These eyes are called "simple" to distinguish the ...

; the irroration of blue scales in interspace 5 with a small subterminal red lunule below it. Antennae, head, thorax and abdomen as in the male.

Sexual Dimorphism

Scientists have found that between the male and female, there are higher amounts of

linalool Linalool () refers to two enantiomers of a naturally occurring terpene alcohol found in many flowers and spice plants. Together with geraniol, nerol, and citronellol, linalool is one of the rose alcohols. Linalool has multiple commercial app ...

and

2,3-butanediol 2,3-Butanediol is the organic compound with the formula (CH3CHOH)2. It is classified as a ''vic''-diol (glycol). It exists as three stereoisomers, a chiral pair and the meso isomer. All are colorless liquids. Applications include precursors to ...

present in the male than in the female. The male also elicits stronger odour than female.

Range

Northern Pakistan Northern Pakistan ( ) is a tourism region in northern and north-western parts of Pakistan, comprising the administrative units of Gilgit-Baltistan (formerly known as '' Northern Areas''), Azad Kashmir, Khyber Pakhtunkhwa, Islamabad Capita ...

Jammu and Kashmir Jammu and Kashmir may refer to: * Jammu and Kashmir (union territory), a region administered by India as a union territory since 2019 * Jammu and Kashmir (state), a region administered by India as a state from 1952 to 2019 * Jammu and Kashmir (prin ...

Garhwal Himalaya The Garhwal Himalayas are mountain ranges located in the Indian state of Uttarakhand. Geology This range is also a part of the Himalayan Sivalik Hills, the outer most hills of the Himalaya located in Himachal Pradesh and Uttarakhand. Major p ...

( Govind Wildlife Sanctuary),

Sikkim Sikkim ( ; ) is a States and union territories of India, state in northeastern India. It borders the Tibet Autonomous Region of China in the north and northeast, Bhutan in the east, Koshi Province of Nepal in the west, and West Bengal in the ...

Assam Assam (, , ) is a state in Northeast India, northeastern India, south of the eastern Himalayas along the Brahmaputra Valley, Brahmaputra and Barak River valleys. Assam covers an area of . It is the second largest state in Northeast India, nor ...

Bangladesh Bangladesh, officially the People's Republic of Bangladesh, is a country in South Asia. It is the List of countries and dependencies by population, eighth-most populous country in the world and among the List of countries and dependencies by ...

Burma Myanmar, officially the Republic of the Union of Myanmar; and also referred to as Burma (the official English name until 1989), is a country in northwest Southeast Asia. It is the largest country by area in Mainland Southeast Asia and ha ...

, southern

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

(including

Hainan Hainan is an island provinces of China, province and the southernmost province of China. It consists of the eponymous Hainan Island and various smaller islands in the South China Sea under the province's administration. The name literally mean ...

Vietnam Vietnam, officially the Socialist Republic of Vietnam (SRV), is a country at the eastern edge of mainland Southeast Asia, with an area of about and a population of over 100 million, making it the world's List of countries and depende ...

, northern

Laos Laos, officially the Lao People's Democratic Republic (LPDR), is the only landlocked country in Southeast Asia. It is bordered by Myanmar and China to the northwest, Vietnam to the east, Cambodia to the southeast, and Thailand to the west and ...

Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...

North Korea North Korea, officially the Democratic People's Republic of Korea (DPRK), is a country in East Asia. It constitutes the northern half of the Korea, Korean Peninsula and borders China and Russia to the north at the Yalu River, Yalu (Amnok) an ...

South Korea South Korea, officially the Republic of Korea (ROK), is a country in East Asia. It constitutes the southern half of the Korea, Korean Peninsula and borders North Korea along the Korean Demilitarized Zone, with the Yellow Sea to the west and t ...

and

Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...

Nepal Nepal, officially the Federal Democratic Republic of Nepal, is a landlocked country in South Asia. It is mainly situated in the Himalayas, but also includes parts of the Indo-Gangetic Plain. It borders the Tibet Autonomous Region of China Ch ...

Habitat

Many species of butterfly occupy only a limited range of the host plant. They utilize the plant chemicals to assess potential host plants and decide if or not 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 ...

on them. Most species in 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'') ...

'' exclusively use rutaceous plants (

rutaceae The Rutaceae () is a family (biology), family, commonly known as the rueRUTACEAE
in Bo ...

) as host plant, and ''Papilio protenor'' is a specialist. Their major host plant is

citrus ''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes. ''Citrus'' is nativ ...

. Research indicates high affinity of the female for ''

Citrus unshiu ''Citrus unshiu'' is a semi-seedless and easy-peeling citrus species, also known as the satsuma mandarin or Japanese mandarin. During the Edo period of Japan, kishu mikans were more popular because there was a popular superstition that eating ' ...

''. There are various oviposition stimulants discovered, including sugar acid,

amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...

alkaloid Alkaloids are a broad class of natural product, naturally occurring organic compounds that contain at least one nitrogen atom. Some synthetic compounds of similar structure may also be termed alkaloids. Alkaloids are produced by a large varie ...

and

flavonoid Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...

. Flavanone glycosides such as

naringin Naringin is a flavanone-7-''O''-glycoside between the flavanone naringenin and the disaccharide neohesperidose. The flavonoid naringin occurs naturally in citrus fruits, especially in grapefruit, where naringin is responsible for the fruit's bitte ...

and

hesperidin Hesperidin is a flavanone glycoside found in citrus fruits. Its aglycone is hesperetin. Its name is derived from the word "hesperidium", for fruit produced by citrus trees. Hesperidin was first isolated in 1828 by French chemist M. Lebreton f ...

were investigated and both elicited positive responses from the female adults. Females do not oviposit on ''

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 wiktionary:黄, 黄wiktionary:檗, 檗), one of the 50 fundamental herbs used in traditional Ch ...

''. The avoidance is attributed to high concentration of

phellamurin Phellamurin, a flavonoid, is the 7-O-β-D-glucopyranoside, 8-C-prenyl derivative of the flavan-on-ol Aromadendrin, and may be seen as the 7-O-glucoside of noricaritin. Being a flavanonol, it has two stereocenters on the C-ring, so four stereoisom ...

. Researchers have found different adaptation to different host plants between adult females and larvae. For example, females are able to oviposit on ''

Trollius asiaticus ''Trollius asiaticus'', the Asian globeflower, is a species of ornamental plant in the family Ranunculaceae, which is native to Asia and Europe. This plant usually grows in wet places, especially in grasslands and forest A forest is an ...

'', but larvae fail to grow on it. Also, while the female adults are not able to oviposit on ''

'', larvae are found to be able to live on it. ''Papilio protenor'' males appear to be more site faithful than females.

Status

Common; not threatened.

Life history

Larva

"Green, with a yellow collar and brown lichen-like markings. Feeds on ''

Zanthoxylum ''Zanthoxylum'' is a genus of about 250 species of deciduous and evergreen trees, shrubs and climbers in the Family (biology), family Rutaceae that are native to warm temperate and Subtropics, subtropical areas worldwide. It is the type gen ...

alatum''." (Mackinnon, quoted in Bingham.) Experiments have been conducted on larvae of different

instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...

s, and it is revealed that among all the different instars, there are significant differences of chemical composition of

osmeteria The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlike ...

l secretion between the 5th instar and all the previous instars. 5th larvae secretion is mainly aliphatic acid and their esters, while the previous instars contain primarily mono- and sesquiterpenoids. Scientists suspect that these differences are associated with the colouration of the 4th larval

ecdysis Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remnant ...

Pupa

"Some pupae are coloured like rough bark, others are uniformly green." (Mackinnon, quoted in Bingham.) There are two colour-types in ''Papilio'' butterflies, green and brown. Pupae that grow on green-coloured branches are always green, while those that live on dead branches are brown in colour. Light condition does not seem to affect this relation. The green pigment is described as a "insectoverdin" type, which means it is a mixture of yellow and blue

chromoprotein A chromoprotein is a conjugated protein that contains a pigmented prosthetic group (or cofactor). A common example is haemoglobin, which contains a heme cofactor, which is the iron-containing molecule that makes Hemoglobin#Oxyhemoglobin, oxygenated ...

s. The chemical compounds separated from the green pupae (e.g., beta-carotene and

lutein Lutein (;"Lutein"

issued a

postage stamp A postage stamp is a small piece of paper issued by a post office, postal administration, or other authorized vendors to customers who pay postage (the cost involved in moving, insuring, or registering mail). Then the stamp is affixed to the f ...

depicting ''Papilio protenor''.

References

{{Taxonbar, from=Q556874 Fauna of Pakistan protenor Butterflies of Indochina Taxa named by Pieter Cramer Butterflies described in 1775 Butterflies of Japan Butterflies of Taiwan