N A T U R K U N D L I C H E S   I N F O R M A T I O N S S Y S T E M

Papilio machaon Linné, 1758

(zoological nomenclature: valid name, available)

General information:

Papilio machaon Linnaeus, 1758: 462
Type locality: Sweden
Type: Lectotype : "27. Machaon"; "Machaon 750."; in coll. The Linnean Collections at The Linnean Society of London (pictures)

Synonyms, misspellings, wrong determinations, etc.:
The following compilation is incomplete and provisional only. It is based on several sources like Savela 2011 (see there for further sources), anonymous 2011b, Bisby et al. 2011 and different other webpages and does not intend to introduce any taxonomical changes.

Europe:
Papilio machaon machaon Linnaeus, 1758: 462. Subspecies. Type locality: Sweden.
Papilio sphyrus Hübner, 1823: pl. 155, f. 775-776. Junior subjective synonym of Papilio machaon machaon Linnaeus, 1758. Type locality: None given (Europe).
? Papilio machaon var. aurantiaca de Selys-Longchamps, 1857: 4. Junior subjective synonym of Papilio machaon machaon Linnaeus, 1758.
Papilio machaon var. marginalis Robbe, 1891: 395. Junior subjective synonym of Papilio machaon machaon Linnaeus, 1758.
Papilio machaon ab. nigrofasciata Rothke, 1895: 303. Unavailable infrasubspecific form.
Papilio machaon ab. niger Heyne, [1895]: 694. Unavailable infrasubspecific form.
Papilio machaon britannicus Seitz, 1907. Junior subjective synonym of Papilio machaon machaon Linnaeus, 1758.
Papilio machaon lapponica Verity, 1911. Junior subjective synonym of Papilio machaon machaon Linnaeus, 1758.
Papilio machaon gorganus Fruhstorfer, 1922. Junior subjective synonym of Papilio machaon machaon Linnaeus, 1758.
Papilio machaon hispanicus Eller, 1936. Junior subjective synonym of Papilio machaon machaon Linnaeus, 1758.
Papilio machaon melitensis Eller, 1936. Junior subjective synonym of Papilio machaon machaon Linnaeus, 1758.

North Africa:
Papilio machaon mauretanica Verity, 1905: 12, pl. 2, f. 5. Subspecies. Type locality: Algeria.
Papilio machaon var. asiatica ab. caerulescens Holl, 1910: 139, f. 5. Unavailable infrasubspecific form. Type locality: Algeria, Harrach.
Papilio machaon var. asiatica ab. djezïrensis Holl, 1910: 140. Unavailable infrasubspecific form. Type locality: Algeria
Papilio machaon maxima Verity, 1911: 296, pl. 52, f. 2. Junior subjective synonym of Papilio machaon mauretanica Verity, 1905. Type locality: Morocco, Tangier.
Papilio machaon maxima gen.aest. angulata Verity, 1911: 296, pl. 60, f. 14. Unavailable seasonal form. Type locality: Morocco, Tangier.
Papilio machaon f. chrysostoma Chnéour, 1934: 41, f. 1. Unavailable infrasubspecific form. Type locality: Tunisia.

Middle East:
Papilio machaon cypriae Verity, 1905. Subspecies. Type locality: Cyprus.
Papilio machaon syriacus Verity, 1908: 12. Subspecies or junior subjective synonym of Papilio machaon cypriae Verity, 1905 ?. Type locality: Caucasus Minor, Armenia, Talysh Mts.
Papilio machaon ruettimanni Seyer, 1977. Junior subjective synonym of Papilio machaon syriacus Verity, 1908.
Papilio machaon muetingi Seyer, 1976: 74. Subspecies. Type locality: Southern Arabia.

Central Asia, Himalayas, Tibet:
Papilio machaon var. asiatica Ménétries, 1855: 70. Subspecies. Type locality: Nepal.
Papilio machaon everesti Riley, 1927. Junior subjective synonym of Papilio machaon asiatica Ménétries, 1855.
Papilio machaon rinpoche Wyatt, 1959. Junior subjective synonym of Papilio machaon asiatica Ménétries, 1855. Type locality: Nepal.
Papilio machaon chitralensis Bang-Haas, 1934. Subspecies ?. Type locality: Pakistan.
Papilio ladakensis Moore, 1886: 46. Subspecies or distinct species?. Type locality: India, Eastern Pamir, Jammu and Kashmir, Ladakh.
Papilio machaon var. centralis Staudinger, 1886: 193. Subspecies.
Papilio machaon pendjabensis Eimer, 1895. Subspecies or junior subjective synonym of Papilio machaon centralis ?. Type locality: India, Pundjab.
Papilio machaon oreinus Sheljuzhko, 1919: Subspecies or junior subjective synonym of Papilio machaon centralis ?. Type locality: Tian-Shan.
Papilio machaon sikkimensis Moore, 1884: 47. Praeoccupavit by Papilio sikkimensis Wood-Mason, 1882 ?. Type locality: India, Sikkim.
Papilio machaon kiyonobu Morita, 1997. Subspecies or junior subjective synonym of Papilio machaon sikkimensis Moore, 1884. Type locality: Tibet.
Papilio machaon nolico Morita, 1997. Junior subjective synonym of Papilio machaon sikkimensis Moore, 1884 ?. Type locality: Tibet.
Papilio machaon weidenhofferi Seyer, 1976. Subspecies. Type locality: Turkmenistan, Kopet-Dagh.

Siberia and Far East:
Papilio hippocrates C. & R. Felder, 1864: 314, 362. Subspecies. Type locality: Japan.
Papilio machaon kaolimachaon Bryk 1946. Junior subjective synonym of Papilio hippocrates C. & R. Felder, 1864.
Papilio machaon koreae Eller, 1939. Nomen nudum.
Papilio machaon kamtschadalus Alphéraky, 1897. Subspecies. Type locality: Kamchatka.
Papilio machaon mandschuriensis Zeller, 1852. Subspecies ?. Type locality: China: Manchuria.
Papilio machaon orientis Verity, 1911. Subspecies or junior subjective synonym of Papilio machaon machon ?. Type locality: Russia: Siberia.
Papilio machaon sachalinensis Matsumura, 1911. Subspecies ?. Type locality: Russia, Sakhalin.
Papilio machaon schapiroi Seyer, 1976. Subspecies ?. Type locality: Russia, Southern Ussuri region.
Papilio machaon septentrionalis Verity, 1911. Subspecies ?. Type locality: Russia, Kurile islands.
Papilio machaon urupensis: Bryk, 1942. Junior subjective synonym of Papilio machaon septentrionalis
Papilio machaon ussuriensis Sheljuzhko, 1910: Infrasubspecific ? (described as seasonal form and therefore unavailable ?). Type locality: Russia, northern and central Ussuri region.

China, Taiwan, Indochina:
Papilio machaon annae Gistel, 1857. Subspecies or distinct species ?. Type locality: China, Qinghai.
Papilio machaon archias Fruhstorfer, 1907. Subspecies or junior subjective synonym of Papilio machaon annae Gistel, 1857. Type locality: China, Sichuan, Fou-Lin.
Papilio machaon chinensis gen. vern. neochinensis Sheljuzhko, 1913. Unavailable infrasubspecific name. Type locality: China: Sichuan.
Papilio machaon chinensis Verity, 1906. Primary homonym of Papilio paris chinensis Rothschild, 1895. Type locality: Sichuan, Wenchuan.
Papilio machaon venchuanus Moonen, 1984. Subspecies or junior subjective synonym of Papilio machaon annae Gistel, 1857.
Papilio machaon montanus Alphéraky, 1897. Subspecies or junior subjective synonym of Papilio machaon annae Gistel, 1857. Type locality: China, Ta-tsien-lu.
Papilio machaon alpherakyi Bang-Haas, 1933. Junior subjective synonym of Papilio machaon montanus Alphéraky, 1897. Type locality: China, Gansu.
Papilio machaon minschani Bang-Haas: Nomen nudum.
Papilio machaon chinensomandschuriensis Eller, 1939. Nomen nudum.
Papilio machaon kunkalaschani Eller, 1939. Junior subjective synonym of Papilio machaon montanus Alphéraky, 1897 ?. Type locality: China, Western Sichuan.
Papilio machaon hieromax Hemming, 1934. Unnecessary replacement name for Papilio machaon montanus Alphéraky, 1897.
Papilio machaon schantungensis Eller, 1936. Subspecies. Type locality: China, Shandong, Tsingtau.
Papilio machaon baijangensis Huang & Murayama, 1992. Subspecies. Type locality: China, Xinjiang.
Papilio machaon verityi Fruhstorfer, 1907: 301 Subspecies or distinct species ?. Type locality: Yunnan.
Papilio machaon birmanicus Rothschild, 1908: 168. Junior subjective synonym of Papilio machaon verityi Fruhstorfer, 1907. Type locality: Myanmar, Shan States.
Papilio machaon suroia Tytler, 1939. Subspecies or junior subjective synonym of Papilio machaon verityi Fruhstorfer, 1907 ?. Type locality: India, Assam, Manipur.
Papilio machaon taliensis Eller, 1939. Subspecies ?, nomen nudum ?. Type locality: China, Northern Yunnan, Dali.
Papilio machaon sylvinus: Hemming, 1933. Subspecies. Type locality: Taiwan.

North America:
Papilio machaon aliaska Scudder, 1869: 407. Subspecies. Type locality: USA, Alaska
Papilio machaon joannisi Verity, [1907]: pl. 10, f. 16. Junior subjective synonym of Papilio machaon aliaska Scudder, 1869.
Papilio machaon petersii Clark, 1932: pl. 6, f. 3. Junior subjective synonym of Papilio machaon aliaska Scudder, 1869.
Papilio machaon bairdii Edwards, 1866: 200. Subspecies or distinct species ?. Type locality: USA, Arizona, Fort Whipple, near Prescott.
Papilio asterius var. utahensis Strecker, [1878]. Junior subjective synonym of Papilio machaon bairdii Edwards, 1866. Type locality: USA, Utah.
Papilio hollandii Edwards, 1892: 50. Junior subjective synonym of Papilio machaon bairdii Edwards, 1866. Type locality: USA, Colorado.
Papilio machaon brucei Edwards, 1893: 253. Subspecies. Type locality: USA, Colorado.
Papilio machaon hudsonianus Clark, 1932: 6, pl. 3, f. 1-2. Subspecies or junior subjective synonym of Papilio machaon brucei ?. Type locality: Canada, Quebec.
Papilio machaon dodi Mc Dunnough, 1939. Subspecies or junior subjective synonym of Papilio machaon brucei Edwards, 1893.
Papilio machaon pikei Sperling, 1987. Subspecies or junior subjective synonym of Papilio machaon brucei Edwards, 1893. Type locality: Canada, Alberta, Dunvegan.
Papilio oregonia Edwards, 1876: 208. Subspecies or distinct species ?. Type locality: USA, Columbia River, near The Dalles.

Old World Swallowtail
Le Machaon, Le Grand Porte-queue
Schwalbenschwanz
Macaone
Makaon

Habit:

   
P. machaon machaon : Salzburg, Flachgau, Thalgau, Thalgauegg, 1980.08.21, coll. Michael Kurz P. machaon machaon , underside: Salzburg, Flachgau, Thalgau, Thalgauegg, 1980.08.21, coll. Michael Kurz    
Picture from: Kurz Michael
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Picture from: Kurz Michael
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Description:
Examined: 1 , 2 of ssp. machaon from central Europe, pictures of ssp. syriacus from the Middle East, ssp. centralis from Tadjikistan, ssp. neochinensis from Sichuan, ssp. chinensis ? from Shaanxi, ssp. hippocrates from Korea and Japan and several ssp. from North America. Wing expanse: 62 mm, 71 - 79 mm. Head and thorax black with a yellow line each, extending to the base of the wing; eyes brown, antennae black; upperside of wings light yellow to sulphurous, forewing with a broad black postdiscal band; the submarginal area black too, interrupted by a line of yellow dots between the veins; basal third of wing black, with scattered yellow scales; veins and three big spots on the anterior margin of the wing also black; hindwings distinctly tailed, the black postdiscal band scattered with blue; in the anal angle of the wing a red and blue, eyelike dot, delicately bordered with black, at least along the margin; veins black too, the curved inner margin black with yellow hairs; underside of wings distinctly lighter than the upper side, the black postdiscal band in the forewing strongly suffused with yellow in the anterior part, the submarginal area mostly yellow; inner margin of the hindwing yellow; underside of subanal area of hindwing with a deep groove; body black, with yellow hairs laterally and beneath.

Variability:
The females usually are bigger and lighter in ground color than the males, with broader fasciae. Generally, the yellow ground colour is subject to variation concerning tone and brightness, as is the width of the postdiscal black bands. Some forms, especillay ssp. hippocrates and various north American subspecies tend to darkened forms regularly or occasionally. On the other side, in ssp. syriacus for example, black spots, especially along the forewing costa, are reduced. The tails of the hindwings vary in length.

Distribution:

P. machaon is distributed in a variety of subspecies and forms from northwest Africa throughout Europe, the middle East and temperate Asia including northern India to Japan, as well as in North America (Higgins & Riley 1978, Tolman & Lewington 1998, Hodges et al. 1983, Kurz & Kurz 2011, GBIF 2011, anonymous 2011a, anonymous 2011b). Some authors have even raised some subspecies to specific rank (e.g. ssp. annae and verityi from China by Chuan-Lung 1980). In the mountains of Europe, the species reaches 2000 m, animals that have gone astray have been found exceptionally up to 3000 m. In central Asia, the species climbs even higher than 4000 m a.s.l. (Kurz & Kurz 2011).

Biology:

P. machaon machaon (race britannicus): UK, Strumpshaw Fen, 2009.06.04 P. machaon machaon (race sphyrus): Portugal, Lissabon, Mercês, Jardim Botânico de Lisboa, 2006.11.18 P. machaon machaon (race gorganus): Austria, Salzburg, Osterhorn group, Schlenken, summit, 2006.06.15 P. machaon syriacus: Turkey, Mugla province
Picture from: wikimedia commons: LittleHow (creative commons)
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Picture from: www.flickr.com: Emerging Birder (Creative Commons)
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Picture from: Kurz Michael
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Picture from: www.flickr.com: © Nichols Karen
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P. machaon syriacus: Armenia, Kotayk province, Garni, Khosrov State Reserve, 2010.06.26 P. machaon chinensis ?: China, Shaanxi, near Xián, 2008.05.02 P. machaon hippocrates: Japan, Osaka, 2009.09.02 P. machaon oregonius: US, Oregon, Redmond, 2008.08.28
Picture from: www.flickr.com: © lepidopterologist
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Picture from: www.flickr.com: © jf_berlin
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Picture from: wikimedia commons: Kenpei (creative commons)
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Picture from: www.flickr.com: Sobkoviak Lauren (creative commons)
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The species inhabits open areas, which are not utilized too intensive. In northern Europe, this is mostly moist land, like moors and marshes, whereas further south, primarily dry biotops like poor meadows or even hot and sunny places are preferred (Tolman & Lewington 1998, Kurz & Kurz 2011). Here, the butterflies have 1 - 3 generations per year (in central Europe mostly April/May and July/August, in southern Europe from late March till early November, see Kurz & Kurz 2011). In summer, the animals often fly together on the summits of hills or mountains (hilltopping). The butterflies have been found on the wing from 9 a.m. till 17 p.m., exceptionally also till 19 p.m., with nectaring activity from 11 a.m. till 17 p.m. (Kurz & Kurz 2011). They are nectaring on a variety of different plants, prefering open accessible blossoms.

Stages in development:

Egg: without data Italy, Abruzzo, Bucchianico first instar: without data young larva: Spain, Galicia, Capela, 2008.08.21
Picture from: wikimedia commons: Chai (GNU Free Documentation License)
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full grown larva: Spain, Galicia, Coirós, 2008.08.26 aberrant form: Italy, Abruzzo, Bucchianico larva, exhibiting its osmaterium: Spain, Castellon, Peñiscola larva, preparing for pupation
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Picture from: www.flickr.com: Angelini Remo (creative commons)
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Picture from: www.flickr.com: Martínez Isidro (creative commons)
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Picture from: www.flickr.com: Morley Dean (creative commons)
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Picture from: www.flickr.com: Morley Dean (creative commons)
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Picture from: www.flickr.com: Morley Dean (creative commons)
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Picture from: www.flickr.com: Morley Dean (creative commons)
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Picture from: www.flickr.com: Morley Dean (creative commons)
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Pupa: Spain, Galicia, Oza dos ríos      
Picture from: wikimedia commons: Bugallo Sánchez Luis Miguel (creative commons)
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Description of early stages following Forster & Wohlfahrt (1976), Tolman & Lewington (1998) and personal observations.

Egg:
The eggs are laid on the leaves of the foodplants. The egg is globular, with weak sculptures, at first light yellowish-green, later becoming darker.

Larva:
In its youth, the larva is black with red papilles and a big white spot on the back. Fully grown, it is green with a black transverse band on each segment, each band carrying 6 orange-red dots, being dissolved into black spots only laterally. Transverse black bands are found also between the segments, but without orange-red dots. For defense purposes, the larva can turn inside out a yellow-orange fork from its neck, from which is excreted a caustic secrete. In very hot mediterranean areas, the larvae are not green, but black and white, the orange-red dots remaining unchanged. As foodplants utilized are different Apiaceae and Rutaceae (Tolman & Lewington 1998, own observations): Aegopodium podagraria, Ammi visnaga, Anethum graveolens, Angelica archangelica, Angelica sylvestris, Bupleurum flacatum, Bupleurum fruticosum, Carum carvi, Crithmum maritimum, Daucus carota, Dictamnus albus, Falcaria vulgaris, Ferula communis, Foeniculum vulgare, Haplophyllum linifolium, Haplophyllum tuberculatum, Heracleum sphondylium, Laserpitium halleri, Laserpitium latifolium, Meum athamanticum, Petroselinum crispum, Peucedanum cervaria, Peucedanum officinale, Peucedanum oreoselinum, Peucedanum palustre, Pimpinella saxifraga, Ridolfia segetum, Ruta angustifolia, Ruta chalepensis, Selinum carvifolia, Seseli varium, Silaum silaus and Trinia glauca. It is not known, if all these plants are also utilized in the wild, or are only suitable for cultivation of the larvae. The larvae develop within a few weeks, pupating on robust stems or on withered woody plants.

Pupa:
The pupa is green, yellow or brown, ventrally with two rows of blunt knobs on the segments. The colour seems to be influenced directly by the environment. The pupa overwinters.

Anatomy:

     
-genitalia: Switzerland      
Picture from: © Higgins 1975, modified
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genitalia (Higgins 1975, based on ssp. machaon). Uncus well formed with slender pointed apex; scaphium vestigal, depressed, with vestiges of tegumen; valve broad, sacculus with densely chitinized, elongate row of many dorsal teeth; penis bowed, short and massive.

Diagnosis:

P. machaon machaon: Austria, Salzburg, Osterhorn group, Ochsenberg, 2001.08.12 P. hospiton: Sardegna, Capoterra, 2008.04.06 P. alexanor: Greece, Peloponnesos, Kambos-ravine, 1999.05.29 P. xuthus: Japan, Osaka-fu, 2007.06.09
Picture from: Kurz Michael
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Picture from: Kurz Michael
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Picture from: wikimedia commons: Kenpei (creative commons)
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P. zelicaon: US, California, Campell, 2009.06.13      
Picture from: www.flickr.com: Zelenke Eugene (creative commons)
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In North Africa and the Middle East, the desert swallowtail (Papilio saharae Oberthür, 1879) cannot be distinguished externally, but its antenna has only 30 - 31 segments (in contrast to 33 - 36 for P. machaon). Papilio hospiton Gené, 1839 (Tyrrhenian Archipelago) and Papilio alexanor Esper, 1799 (southern Europe and Middle East to central Asia) can be distinguished by their wing pattern: In P. hospiton, the black drawings are more widespread, especially along the veins. The eyelike spot in the anal angle of the hind wing is crescent-shaped and not round, the tails are distinctly shorter than in P. machaon. In P. alexanor, the basal black area of the forewing is reduced to a transverse fascia at about 1/4, which is extending also onto the hindwing. The submarginal spots on the forewing mostly are united to form a non interrupted band. In eastern Asia, the most similar species to P. machaon is Papilio xuthus Linné, 1767, but it can readily be distinguished by the broadly black lined veins on the upperside and by the basal part of the forewing upperside, which shows longitudinal black and yellow stripes, in contrast to P. machaon, where the basal part of the forewing is black with yellow suffusion. In North America, there are several somewhat similar species, the most similar being Papilio zelicaon Lucas, 1852. In this species, the big black costal spots on the forewing upperside are much broader and often confluent. Furthermore, P. zelicaon often shows extensive black colouring in the basal half of both wings on the upperside. Such black colouring on the other hand, may occur also in syntopically flying supspecies of P. machaon and also in hybrids of both species. Nevertheless, P. zelicaon can easily be distinguished by the red eyespot in the anal angle of the hindwing, which is centered black in that species, whereas in all forms of P. machaon, the small black dot does not lie in the center, but at the edge of the eyespot, close to the wing margin.

Genetics:

Chromosome number (haploid): 30 - 32 (!?, Higgins, 1975)

Worth knowing:

Ethymology:
Following Spuler (1908), Machaon, the brother of Podalirius, has been the name of a physician in greek mythology.

Sources:

anonymous 2011a. Butterflies of Japan. Yamaha corporation.- available at: http://www.asahi-net.or.jp/~ak5t-kmn/ (online 21 March 2011).
anonymous 2011b. Papilio machaon LINNAEUS, 1758 (Lepidoptera, Papilionidae). 7 pp. - available at: www.actias.de/index.php?page=Attachment&attachmentID=16992.
Bisby F. A., Y. R. Roskov, T. M. Orrell, D. Nicolson, L. E. Paglinawan, N. Bailly, P. M. Kirk, T. Bourgoin, G. Baillargeon, & D. Ouvrard eds (2011). Species 2000 & ITIS Catalogue of Life: 2011 Annual Checklist. Digital resource at www.catalogueoflife.org/annual-checklist/2011/. Species 2000: Reading, UK.
Chuan-Lung, L. 1980. A Revision of the Chinese species of Papilio machaon L. and their geographical distribution. Acta Entomologica Sinica 23 (4): 427-431.
Forster W. & Th. A. Wohlfahrt (1976): Die Schmetterlinge Mitteleuropas, 2, 2. Auflage, Franckh´sche Verlagshandlung, Stuttgart
GBIF 2011. Biodiversity occurrence data accessed through GBIF Data Portal, www.gbif.net, [online 22 March 2011].
Higgins, L. (1975): The Classification of European Butterflies, Collins, London
Higgins, L.G. & N.D. Riley (1978): Die Tagfalter Europas und Nordwestafrikas, übersetzt von W. Forster, 2. Auflage, Verlag Paul Parey, Hamburg und Berlin
Hodges, R. W. et al. 1983. Check List of the Lepidoptera of America North of Mexico. E. W. Classey, London, UK. Updated by Jean-François Landry, Don Lafontaine and Jim Troubridge [available on http://www.lepbarcoding.org/cl_nth_am.php]
Kurz, M. A. & M. E. Kurz 2000–2011. Naturkundliches Informationssystem. – URL: http://www.nkis.info [online 31 March 2011].
Leraut P. (1992): Les papillons dans leur milieu, Bordas
Linnaeus, C. 1758. Systema Naturae, Ed. X. (Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.) Holmiae. Systema Nat. ed. 10 i-ii + 1-824
Savela, M. 2011. Lepidoptera and some other life forms. Papilio Linnaeus, 1758. - available at: http://ftp.funet.fi/index/Tree_of_life/insecta/lepidoptera/ditrysia/papilionoidea/papilionidae/papilioninae/papilio/index.html [online 28 March 2011].
Spuler, A. 1908/1910. Die Schmetterlinge Europas. 4 vol., Stuttgart – E. Schweizerbart´sche Verlagsbuchhandlung (E. Nägele).
The Linnean Society of London 2011. The Linnean Collections. - available at: http://www.linnean-online.org/. (online 21 March 2011).
Tolman T. & R. Lewington (1998): Die Tagfalter Europas und Nordwestafrikas, übersetzt von M. Nuß, Franckh-Kosmos Verlags-GmbH & Co., Stuttgart.

Publication data:

history:
Kurz Michael: 2011.02.17
Kurz Michael: 2011.03.23
Kurz Michael: 2011.03.29
Kurz Michael: 2011.03.31
Kurz Michael: 2011.04.01
Kurz Michael: 2011.04.06
Kurz Michael: 2011.04.07
Kurz Michael: 2011.04.12
Kurz Michael: 2011.04.26
Kurz Michael: 2011.05.11
Kurz Michael: 2014.05.14
not reviewed

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