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The Apollo or mountain Apollo (''Parnassius apollo''), is a
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprise ...
of the family
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 larg ...
.


Etymology

The species is named in the classical tradition for the deity
Apollo Apollo, grc, Ἀπόλλωνος, Apóllōnos, label=genitive , ; , grc-dor, Ἀπέλλων, Apéllōn, ; grc, Ἀπείλων, Apeílōn, label= Arcadocypriot Greek, ; grc-aeo, Ἄπλουν, Áploun, la, Apollō, la, Apollinis, label ...
.


Subspecies

Subspecies include: * ''Parnassius apollo apollo'' L. * ''Parnassius apollo alpherakyi'' Krulikowsky, 1906 * ''Parnassius apollo bartholomaeus'' Stichel, 1899 * ''Parnassius apollo democratus'' Kulikowsky, 1906 * ''Parnassius apollo filabricus'' Sagarra, 1933 * ''Parnassius apollo gadorensis'' Rougeot & Capdeville, 1969 ( Sierra de Gádor). Extinct. * ''Parnassius apollo geminus'' Schawerda, 1907 * ''Parnassius apollo graecus'' Ziegler, 1901 * ''Parnassius apollo hesebolus'' Nordmann, 1851 * ''Parnassius apollo hispanicus'' Oberthür, 1909 Central (
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
) * ''Parnassius apollo limicola'' Stichel, 1906 * ''Parnassius apollo merzbacheri'' Fruhstorfer, 1906 * ''Parnassius apollo nevadensis'' Oberthür, 1891 ( Sierra Nevada) * ''Parnassius apollo provincialis'' Kheil, 1905 * ''Parnassius apollo pyrenaica'' Harcourt-Bath, 1896 * ''Parnassius apollo rhodopensis'' Markowitsch, 1910 (
Greece Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders ...
,
Balkans The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...
) * ''Parnassius apollo rhaeticus'' Fruhstorfer, 1906 * ''Parnassius apollo rhea'' (Poda, 1761) * ''Parnassius apollo rubidus'' Fruhstorfer, 1906 * ''Parnassius apollo sibiricus'' Nordmann, 1851 * ''Parnassius apollo sicilae'' Oberthür, 1891 * ''Parnassius apollo valesiacus'' Fruhstorfer, 1906 * ''Parnassius apollo vinningensis'' Stichel, 1899 ( Moselle, Duitsland) For a more complete list of subspecies and type details of specimens in the British Museum (Natural History) see Ackery, P. R. (1973) A list of the type-specimens of Parnassius (Lepidoptera: Papilionidae) in the British Museum (Natural History). ''Bulletin of the British Museum (Natural History) Entomology'' 29 (1) (9.XI.1973): 1—35, 1 pl
online here


Distribution and habitat

This typically mountain species prefers hills and flowery alpine meadows and pastures of the continental European mountains, in Spain, Scandinavia and Central Europe, in the Balkans up to northern Greece and in the Alps between Italy and France.Funet-Taxonomy, distribution and images
/ref> It is also present in some areas of the central Asia (
Sakha Sakha, officially the Republic of Sakha (Yakutia),, is the largest republic of Russia, located in the Russian Far East, along the Arctic Ocean, with a population of roughly 1 million. Sakha comprises half of the area of its governing Far Ea ...
). Typical of high altitudes, its range is from up to , although it is far more present above .Captain's European Butterfly Guide
/ref> This species requires specific climatic conditions (cold winter, sunny summer). It also requires wide open spaces (with a cover of shrubs less than 5%) and a large surface of lawns (at least 50%). The presence of the host plant for the caterpillars is critical.


Description

''Parnassius apollo'' has a
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan o ...
of in males, of in females. The Apollo butterfly shows a great deal of individual variation in the appearance, with an evident colour polymorphism. These very very large, beautiful and conspicuous white butterflies are decorated with five large black eyespots on the forewing and two bright red or sometimes orange eyespots on the hindwing.Carter, D. (2000) Butterflies and Moths. Dorling Kindersley, London. These striking red eyespots can vary in size and form depending on the location of the Apollo butterfly, and the bright red colour often fades in the sun, causing the eyespots of older individuals to appear more orange. The wings are shiny, with slightly transparent edges,Still, J. (1996) Butterflies and Moths of Britain and Europe. Harper Collins, London. and some individuals are darker (sphragismelanistic); a general phenomenon common in many butterflies. The caterpillars of this species are velvety black with orange-red spots along the sides. Related species can be found all over the world. The clouded Apollo (''Parnassius mnemosyne'') lives in the valleys. while the small Apollo (''Parnassius phoebus'') is found in the high mountains. The latter one has strongly marked black and white antennae, with presence of two red spots near the apex of forewings.Euro Butterflies by Matt Rowlings
/ref> Parnassius apollo MHNT CUT 2013 3 3female Aragnouet dos.jpg, Female Parnassius apollo MHNT CUT 2013 3 3 female Aragnouet ventre.jpg, Female, bottom Parnassius apollo MHNT CUT 2013 3 3 male Porté-Puymorens dos.jpg, Male Parnassius apollo MHNT CUT 2013 3 3 male Porté-Puymorens ventre.jpg, Male, bottom


Distribution of colour variation

The drastic climate change of the Pleistocene era forced a separation of the red Apollo butterfly population. This in turn played a role in creating the distinct colour changes seen in the species. The ''Parnassius apollo'' became divided and isolated in the Eurasian region during the glacial period. The large glaciers created a physical barrier between the population, barring interaction between the groups. Still within these isolated populations the butterflies migrated westward into portions of southern Europe where they settled and reproduced. Within all of these particular isolated populations there is also variation in the wing colour allele. There is variation in size between the isolated populations. With the larger separated populations of butterflies these habitats are used to sustain populations with larger amount of resources. These larger populations are called metapopulations and with the smaller separated sub-populations they create a mainland-island system. The ''Parnassius apollo'' can migrate from habitats and thus create a variation seen in each isolated population.


Distribution pattern effects on survival rate

New environmental pressures lead to the selection of a better suited colour variant within these isolated populations. With this variety there is a correlation with extinction. In high variation environments there is cause for extinction in greater numbers of individuals. For example, there is high variation in the Swiss Alps and presently there is a high rate of individuals becoming extinct. One of the probable causes of extinction is the warming of the climate. It is said that the red Apollo is an "atypical glacial invader" and that with the warming of the climate in mountainous regions is causing the butterfly to not readily adapt to such an uncomfortable environment. Another possible cause of extinction is the interesting connection between nectar plant distribution and the ''Parnassius apollo''. If there was present a constraint of migration from nectar plant populations to another the red Apollo's population would slowly dissipate and reproduction might seize. This is because the outcrop of the nectar plants are the sight of reproduction and if the spatial structure is too far for the butterfly to migrate to the dynamics of the population is in danger.


Ecology

This species has a single brood. Adult Apollo butterflies are seen on the wing from May to September, feeding on nectar produced by flowers. During mating males deposit on the female's abdomen a gelatinous secretion called sphragis, that prevents the female mating a second time. The females lay eggs, which over-winter and hatch in spring the following year. The Apollo caterpillar a velvety blue black with small orange spots. These caterpillars feed on stonecrop (''
Sedum ''Sedum'' is a large genus of flowering plants in the family Crassulaceae, members of which are commonly known as stonecrops. The genus has been described as containing up to 600 species, subsequently reduced to 400–500. They are leaf succul ...
'' species, mainly '' Sedum telephium'', '' Sedum album'', '' Sedum rupestre'' and '' Sedum ropsea''), '' Hylotelephium caucasicum'' and houseleek (''
Sempervivum ''Sempervivum'' (Brit. , U.S. sɛ̃mpeɹ'vivũm is a genus of about 40 species of flowering plants in the family Crassulaceae, commonly known as houseleeks. Other common names include liveforever (the source of the taxonomical designation ''Sem ...
'' species). When the caterpillar is fully grown it will pupate on the ground, forming a loose cocoon from which the adult butterfly emerges following metamorphosis.


Predation and defensive strategy

The Apollo butterfly shares a variety of defensive strategies with quite a few species of butterflies. Even from a young age larva exhibit camouflage by being entirely black. This solid colour helps them avoid detection even at a close distance. However, as they mature, they lose this advantage by developing two rows of orange dots. These dots greatly decrease the amount of crypsis.Descimon, H., and M. Deschamps-Cottin. "A Possible Case of Mimicry in the Caterpillar of Parnassius Apollo (L.) (Lepidoptera: Papilionidae)." Linneana Belgica 15.8 (1996): 309-10. Print. In addition to this larval camouflage, the larva also shares in a form of
Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Mülleria ...
with a type of millipede, ''
Glomeris ''Glomeris'' is largely European genus of pill millipedes. It contains over 100 species, distributed across Europe, but also extending into the Canary Islands, Turkey, and eleven species from North Africa. Species * '' Glomeris albida'' * '' ...
( Glomeris guttata''). Both animals share the characteristic orange spots and black body and a common habitat. The millipedes and caterpillars secrete a foul smelling odour to repel predators. Once the butterfly completes its metamorphosis, it has a number of defensive mechanisms in place to avoid predation. One of the most easily identifiable traits is the bright eyespots found on the wings. These eyespots are essentially concentric circle of a wide variety of colours. Apart from the wide range of colours, eyespots are very limited in their plasticity. There are three main hypothesis to why these spots may have developed; they resemble the eyes of an enemy of the predator in order to intimidate them, they draw the attention of the predator to less vital components of the butterfly's body, or the spots are there simply to surprise the predator. The only disadvantage to these spots is that they cause the butterfly to be a great deal more conspicuous. Another form of defence is the taste of the butterfly. Similar to the monarch butterfly, the Apollo butterfly produces a repulsive taste to its predator. The butterfly seems to get this foul taste from its plant host, the '' Sedum stenopetalum''. There is a bitter tasting cyano glucoside, sarmentonsin, which is found in both the butterfly and the plant. There is a much higher concentration of sarmentonsin in the wings as opposed to the rest of the body. The high concentration in the wings indicates that the wings of the butterfly would taste much worse comparatively. A common predator, nesting
water pipit The water pipit (''Anthus spinoletta'') is a small passerine bird which breeds in the mountains of Southern Europe and the Palearctic eastwards to China. It is a short-distance migrant; many birds move to lower altitudes or wet open lowlands i ...
s, have evolved a strategy to avoid the poor taste of the butterfly; the bird will remove the wings before consuming the body. In theory, this will get rid of the poor tasting elements of the butterfly, leaving only the nutritious body.


Conservation

This species is of interest to entomologists due to the variety of subspecies, often only restricted to a specific valley in the
Alps The Alps () ; german: Alpen ; it, Alpi ; rm, Alps ; sl, Alpe . are the highest and most extensive mountain range system that lies entirely in Europe, stretching approximately across seven Alpine countries (from west to east): France, Swi ...
. The beautiful Apollo butterfly has long been prized by collectors, who aim to possess as many of the variants as possible. While over-collecting is believed to have caused populations to decline in some areas, such as in Spain and Italy, habitat change is thought to be a far more significant threat to this species' survival. Plantations of conifers, the succession of suitable habitat to scrubland, agriculture, and urbanization have all reduced the habitat of the Apollo butterfly.
Climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
and acid rain have also been implicated in this species decline in Fennoscandia. In addition, motor vehicles have been cited as a cause of Apollo butterfly mortalities; vehicles on a motorway system near
Bolzano Bolzano ( or ; german: Bozen, (formerly ); bar, Bozn; lld, Balsan or ) is the capital city of the province of South Tyrol in northern Italy. With a population of 108,245, Bolzano is also by far the largest city in South Tyrol and the third la ...
in South Tyrol, Italy, are said to have nearly wiped out a race of the Apollo. In
Finland Finland ( fi, Suomi ; sv, Finland ), officially the Republic of Finland (; ), is a Nordic country in Northern Europe. It shares land borders with Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of B ...
, the Apollo was one of the first species of insects declared endangered. The Apollo population in Finland and Sweden decreased drastically during the 1950s. The reason for this is not known, but it is commonly thought to be because of a disease. In Sweden, it is now restricted to areas that have
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
in the ground, suggesting that the decrease could hypothetically be related to acid rain. Laws exist to protect the Apollo butterfly in many countries. The Apollo is on the IUCN Red List of Threatened Species, in Appendix II in
CITES CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
, and is mentioned in annex IV of
Habitats Directive The Habitats Directive (more formally known as Council Directive 92/43/EEC on the Conservation of natural habitats and of wild fauna and flora) is a directive adopted by the European Community in 1992 as a response to the Berne Convention. The E ...
. It is protected in other states: the Principality of
Liechtenstein Liechtenstein (), officially the Principality of Liechtenstein (german: link=no, Fürstentum Liechtenstein), is a German-speaking microstate located in the Alps between Austria and Switzerland. Liechtenstein is a semi-constitutional monarch ...
,
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
(as critically threatened species in Czech code, Decree for implementation, No. 395/1992 Sb., and No. 175/2006 Sb.), Turkey and
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populou ...
. However, these laws focus on the protection of individuals, rather than their habitat, and so may do little to mitigate the greatest threat that populations face. Fortunately, there are a number of projects specifically working to save this vulnerable insect. A conservation programme in Pieniny National Park saved a subspecies of the Apollo butterfly that had declined to just 20 individuals in the early 1990s, through a combination of captive breeding and habitat protection. In south-west Germany, conservationists are working with shepherds to ensure favourable conditions for the butterflies, which share their grassland habitat with sheep. For example, grazing periods have been shifted to avoid the Apollo butterfly larvae stage, which is vulnerable to being trampled. The Apollo butterfly has many subspecies around the world, and some European subspecies are showing an alarming decline in numbers. This is mainly caused by
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
, air pollution affecting the insect's food plants, and butterfly collectors. The Apollo butterfly is also more vulnerable to predators as it spends two years as a caterpillar.


References


External links


Lepiforum.de''P. apollo'' images
at
Consortium for the Barcode of Life The Consortium for the Barcode of Life (CBOL) was an international initiative dedicated to supporting the development of DNA barcoding as a global standard for species identification. CBOL's Secretariat Office is hosted by the National Museum of ...
* *


Further reading

* Xavier Mérit and Véronique Mérit: ''Les'' Parnassius ''de France'', textes de Xavier Mérit-Véronique Mérit et Henri Descimon, cartes de répartition, planches, et photos en nature de Luc Manil, Xavier Mérit et Bernard Turlin, bibliographies, Bulletin des Lépidoptéristes parisiens, Volume 15 (2006), n°33 (numéro thématique), Paris, septembre 2006 (56 pages). * Pierre Capdeville, 1978–1980, Les races géographiques de ''Parnassius apollo'', 191 p - 26 tables - maps - 24 plates in colours, Sciences Nat, Venette. * Jean-Claude Weiss: ''The Parnassiinae of the World'', Pt. 4, 2005,
Hillside Books, Canterbury Hillside Books, Canterbury was a publisher specialising in books on entomology and small equipment associated with this science. Origin It was established in 1989, directed by Lydie Leforestier. Initially it was in Canterbury, England but then ...


* Edwin Möhn, 2005 ''Schmetterlinge der Erde, Butterflies of the world'' Part 23 Papilionidae XII. Parnassius apollo. Edited by Erich Bauer and Thomas Frankenbach Keltern: Goecke & Evers; Canterbury: Hillside Books. {{Authority control Parnassius Butterflies of Asia Butterflies of Europe Butterflies described in 1758 Taxa named by Carl Linnaeus