Red Apollo the a. rubidus group on a thistle blossom (South Tyrol)
|( Linnaeus , 1758)|
The red Apollo or Apollo butterfly ( Parnassius apollo ) is a severely endangered and strictly protected butterfly ( butterfly ) from the family of the knight butterfly (Papilionidae) in Europe . The generic name is derived from Mount Parnassus in central Greece , which is considered the seat of the Muses and is dedicated to the god Apollon . The Apollo butterfly was the animal of the year in Germany in 1995 .
The moths reach a wingspan of 60 to 88 millimeters. The thorax is light gray to black and covered with fine hair-like scales . The wings are dominated by a whitish base tone. The outer edge is not scaled and looks glassy. One or more black spots can be seen on the forewing. The hind wing is characterized by black, red, sometimes yellowish eye spots (ocellae) with white mirrors. The ocels are particularly large in females. In addition, depending on the subspecies, more or less often dark pollinated individuals appear in the females. The darker color is used to camouflage and to increase the body temperature in higher altitudes in sunshine. With their few characteristics, the moths have a very variable appearance. Sometimes red spots can also be seen in the post-disk region of the forewings and on the inner angle of the hindwings. The upper and lower side of the wing are colored the same except for additional red spots in the basal area of the lower side of the wing.
The ocells do not imitate eyes, but serve as a warning of the toxicity of the moths that have received them through the caterpillar feed. In the rest position, with the wings folded, the red spots come into their own. Both birds and lizards avoid the moths as food.
The antennae are an important distinguishing feature from the Alpine Apollo ( Parnassius phoebus ) . In the Red Apollo they are monochrome gray with black pistons and in the Alpine Apollo they are black and white ringed with black pistons.
The size of the moths varies greatly and depends on the one hand on the height of the habitat and on the other hand with the food plant. The higher the habitat, the smaller the moths become. In subspecies that feed on purely succulent plants, the moths become smaller. The more succulent the plants are, the more water and fewer nutrients they contain. Thereby, the butterfly in Central Europe, where the succulent Sedum album ( Sedum album ), the preferred larval food is smaller than moths in the east and north of range, in the weakly succulent plants of genus Orostachys or subspecies of Great sedum ( Sedum telephium ) the preferred food of the caterpillars.
Features of the pre-imaginal stages
The eggs are white in color, rounded and have a grainy surface.
The black caterpillars are ringed dark gray at the segment boundaries and have short hairs. You have two or three red to yellow spots on each side of the thorax segment. Like the red ocella of the moth, these serve to warn of the toxicity of the caterpillars. Differences in the size and color of the mostly three spots can be used to differentiate between subspecies. Like all caterpillars of the knightly butterfly, these have a nuchal fork ( osmaterium ) between the head and the first thorax segment .
The mummy dolls are black-brown and have a bluish frosting.
Way of life
The caterpillar overwinters in the egg shell and leaves it in spring. It lives individually on the food plants, which vary depending on the region. In Europe, the food plants are white sedum ( Sedum album ) and large sedum ( Sedum telephium ssp.telephium ), more rarely also rock sedum ( Sedum rupestre ), or species of the genus Rhodiola from the family of thick-leaf plants (Crassulaceae) as well as the alpine apollo Parnassius phoebus . In the vicinity of Lake Baikal , the caterpillar feeds on the sedum plant or on species of the genus Orostachys (thick-leaf family) as well as the caterpillar of the closely related Parnassius nominon . With the expansion to the west, there was also a change from the lowlands to the mountainous areas, and this was accompanied by a change in the food plant from Sedum telephium subspecies to sedum sedum and closely related species. This change can be observed in the Alps , Turkey and the Beskids . Caterpillars from the eastern lowlands refuse the white sedum plant and die if they do not get their usual food plants, while the caterpillars from alpine regions also eat Sedum telephium subspecies.
The caterpillar likes to stay under stones and often sunbathes on the food plant to warm up. It pupates in a loose web in a mummy doll on the food plant or under stones. The duration of the pupal rest varies greatly, from eight to ten days to several weeks, which results in the butterfly's long flight time. Shortly after the moths hatch, they mate. In the female, the genital opening is closed with the so-called sphragis after the long-lasting copula ; which prevents further mating. The egg is laid soon after the copula. The females fly little while the males patrol their area. As a result, the spread to new habitats is slow. The moths prefer dry and warm, often rocky, habitats. The similar Alpine Apollo, on the other hand, can be found in spring meadows and in the vicinity of mountain streams.
The females lay up to 100 eggs individually on the food plants and the caterpillars overwinter fully developed in the egg shell.
As moths, the animals visit not only the white flowers of Sedum album , but preferably the red and purple flowers of thistles (e.g. Carduus nutans ), knapweeds ( Centaurea jacea, etc.) and oregano ( Origanum vulgare ). The moths often sit on stones with their wings open and bask in the sun.
distribution and habitat
The most important habitat of the Red Apollo are calcareous rubble corridors (over lime) and scree slopes (over silicate) in the high and low mountain ranges. It used to be seldom in vineyards; it is occasionally found in poor grasslands with sedum deposits. Its habitat are sunny, dry locations with stony subsoil, especially rocky slopes, scree slopes and rocky edges, leg stone walls, also railway and road embankments as well as spoil heaps from quarries, but hardly any with modern mining methods. Larval habitats with occurrence of the host plants (see above), on which also the oviposition takes place, determine its occurrence. In the Southern Alps, habitats from 1000 m above sea level are found. in Greece those over 1800 m above sea level. populated. In Scandinavia it occurs at sea level.
The flying season of the only annual generation begins at the end of May and lasts until around the end of August. The long flight time results from the very different rest of the pupae of the individual animals, because the life span of the moths is only about two to three weeks.
Its distribution ranges from the Iberian Peninsula across all European mountain regions, the Carpathians , the Caucasus and the Urals , the area of origin of the species on Lake Baikal, to Yakutia in the east, often in isolated populations . The northern distribution extends to Fennoscandinavia , the southern limit is the Sierra Nevada , Sicily , southern Turkey. It is completely absent in the British Isles and Denmark .
The vertical distribution begins at around 400 meters from the colline level and extends to around 2000 meters in the high-montane and subalpine areas.
The locally faithful moth often lives in clearly defined areas in isolated populations, which has led to the formation of many subspecies, semi-subspecies and semi-species (semispecies) due to the resulting lack of gene exchange. Within the species and the subspecies, the butterfly is extremely variable in its appearance and can therefore not be assigned to a subspecies solely on the basis of morphological differences. The location must also be used for allocation. About 290 subspecies have already been described, but many of them are now classified as synonyms. Möhn (2005) gives about 250 subspecies and expects a further reduction based on genetic studies. Even within the genus Parnassius , the classification of the species is not clear and varies between 38 (UNEP-WCMC, 2006) and 47 ( Weiss , 1991). These are in turn divided into eight subgenus, in which the red Apollo is the type for the subgenus Parnassius .
Selection of European subspecies:
- Parnassius apollo apollo Linnaeus , 1758, Sweden . The large butterfly with wingspan of 76 to 88 millimeters lives in the southern Swedish lowlands. The large eye-spots on the hind wings often have clear white mirrors. The caterpillar feeds mainly on Sedum telephium , but also eats Sedum album ( Sedum album ).
- Parnassius apollo nevadensis Oberthür 1891, Spain . The moth lives at an altitude of 1700 to 2500 meters in the Sierra Nevada ( Betic Cordillera ) and becomes significantly smaller with increasing altitude.
- Parnassius apollo filabricus de Sagarra y Castellarnau 1933, Spain, Betic Cordillera. The moth lives at an altitude of 2000 to 2100 meters in the Sierra de los Filabres
- Parnassius apollo gadorensis Rougeot & Capdeville 1969, Spain, Betic Cordillera. The butterfly lives at an altitude of 1900 to 2000 meters in the Sierra de Gádor
These three Spanish subspecies are very closely related and hardly differ from one another. As with all Spanish subspecies, the front wings of the fresh moths are clearly black and white piebald. In both sexes, the ocels are orange-yellow instead of red. The females are darkly pollinated and often have an orange-yellow front margin spot and anal spot. The caterpillars of the Spanish subspecies eat Sedum amplexicaule , Sedum micranthemum and stonecrop ( Sedum acre ).
- Parnassius apollo hispanicus Oberthür 1909, Spain. The butterfly lives at an altitude of 1600 to 1800 meters in the Sierra de Albarracín in Aragon . The males are bright and only have small ocells. The females are, like all in Spain, dark pollinated. In contrast to the previously mentioned subspecies, the ocells are rarely orange-yellow and have a large white mirror.
- Parnassius apollo testoutensis Eisner 1957, France . This small, light-colored subspecies, the females of which are only weakly pollinated, has a very large distribution area in the Savoy Alps with the Haute-Savoie department , the Graian Alps and the Dauphiné Alps .
- Parnassius apollo pumilus burin 1906, southern Italy . With a wingspan of 60 to 63 millimeters, the butterfly is the smallest of all subspecies and occurs in Aspromonte and Calabria . It is classified as a semi-species that splits off from Parnassius apollo . Its habitat has only sparse vegetation at altitudes of 1200 to 1900 meters. The caterpillars feed on Sedum tenuifolium . The males differ significantly from Parnassius apollo apollo and with their light-colored wings with small spots and light-colored ocels are more similar to the Alpine apollo ( Parnassius phoebus ), while the females are pollinated darkly.
- Parnassius apollo rhodopensis Markovic 1909, Bulgaria . This subspecies is the connection between the northern Turkish and the Balkan subspecies. It has distinctive spots and some of the pollinated females have a double mirror in the lower ocellus.
- Parnassius apollo vinningensis (Mosel Apollo) Prick 1899, Germany. The subspecies is named after Winningen and occurs in the lower Moselle valley from Güls near Koblenz to Traben-Trarbach . It differs significantly from Parnassius apollo meridionalis in the Vosges . The moth is chalk white and often has a white spot in the root pollination outside the cell . A special feature are the kidney-shaped lower ocells. The food plant Sedum album grows here on the natural steep banks of the Moselle and on the stone walls built by vintners in Roman times . The main flight time is from mid-June to mid-July.
- The Small Apollo Parnassius phoebus has in some authors the status of a Halbart and flies in the Central Alps at higher altitudes than Parnassius apollo (ecological allopatry ) and its caterpillars have to saxifrage ( Saxifraga changed).
The genus Parnassius originated in the Siberian-Mongolian region around Lake Baikal. Many species closely related to Parnassius apollo such as Parnassius nominon , Parnassius phoebus and Parnassius bremeri still live there today . With the vicarious species Parnassius nominon , Parnassius apollo repeatedly forms hybrids without these developing a zone where two areas of distribution of the species meet, as would be the case with speciation. From the area of origin, the genus spread to the east and west. Already in the Upper Pliocene ( Gelasian ) the species colonized large parts of today's distribution area. The climate there was similar to that of today, whereas in the previous Miocene it was still tropical to subtropical, making it unsuitable as a habitat for the Apollo butterfly.
Spread to the east
Parnassius apollo has its easternmost distribution in Yakutia , while the vicariate to Parnassius phoebus , Parnassius bremeri spread further to the northeast and colonized the North American continent in Alaska and the northern Yukon Territory via the Bering Strait . The Bering Strait, which is only 60 meters deep today, fell dry during the Old Quaternary and did not stand in the way of expansion. Other areas of North America are populated by the genus with Parnassius bremeri s vicariate Parnassius smintheus across the Rocky Mountains in the west to Tamaulipas in northeast Mexico in the south. Parnassius smintheus could arise during the Ice Ages when the southern populations were separated from the populations in the ice-free regions of Alaska by ice sheets.
Spread to the west
The main spread of Parnassius apollo goes west and finds its westernmost point in Spain. There are several lines of expansion to the west that follow mountain chains on the one hand or run over flat land with forest steppes.
The southernmost line of spread extends over many mountain ranges from the Changai Mountains in Mongolia via South Siberia ( Sajan Mountains , Tannu-ola Mountains ), the Altai , Salair Ridge , Tarbagatai Mountains , Saur Mountains , Djungarian Alatau , to Tianshan in Uzbekistan . This is where the southern-western distribution ends. On the southern slopes of the Tianshan Mountains, it goes back east over the Hissar Mountains and Kunlun Mountains to China in the Xinjiang Mountains with the subspecies Parnassius apollo khotanensis .
The way to the Alps
A second distribution to the west took place below the 60th parallel via the forest steppes to the southern Urals . With its low mountains and passes, this did not constitute an obstacle to further expansion to the west via the western taiga to southern Scandinavia . There southern Finland, Sweden and Norway were settled. Another line spread south across the Caucasus to Turkey and then went northwest to the Rhodope Mountains in Bulgaria and west across the then dry Aegean Sea to Greece . From the Rhodopes it went north over the Balkan Mountains , the Southern Carpathians and northwest over the Carpathians . From there the expansion followed the mountains of the High Tatras , Giant Mountains , Ore Mountains , Fichtel Mountains , Franconian Alb , Swabian Alb to the west until it ended in the Black Forest . The Upper Rhine Plain was an unbridgeable obstacle and the line ended here. From the High Tatras to the southwest over the Little Carpathians and the Vienna Woods were Ostalpen reached. The northern Alps were then settled up to the Bregenz Forest . The route to the Southern Alps ran from the Rhodopes across the Western Balkans. The main Alpine ridge was an insurmountable obstacle and the two lines along the Alps only met again in the western Swiss Alps.
The Moselle, Italy and Spain
The French Alps or the Jura were the starting point for conquering other habitats in the north, west and south.
From the Jura to the north via the Burgundian Gate , the Vosges were populated, the mountains that run almost parallel to the Black Forest along the Upper Rhine Plain and were not populated from there. From here the species reached the Moselle region, in which a protected and stable population still exists today due to the rich occurrence of the forage plant sedum plant - especially in the slopes of the vineyards between the villages of Valwig and Bruttig-Fankel .
The settlement of Spain took place from the Jura to the west across the Massif Central . From there over the southern Cevennes to the eastern Pyrenees. In Spain, the spread was along the north coast over the Cantabrian Mountains and then south to the Castilian Scheidegebirge in central Spain. Finally, the Betic Cordilleras in the south were settled via a route that is still unknown today.
The influence of the ice ages on species development
The Red Apollo is a glacial immigrant from Asia who between 100,000 and 70,000 per cent, with the opening of the habitats after the Riss-Würm interglacial, rapidly expanded its area in a westerly direction. Phylogenetic signals can also suggest primary immigration during the Rift Age. At the glacial maxima, the new area in the west was separated from the original area in West Asia by inland freezing in Central and Northern Europe. At the same time, differentiation began in the southern populations. A second wave of area expansion began in southern Europe with the onset of Ice Age glacier maxima under fully glacial climatic and habitat conditions. The species could have been around 30,000 BC. H. penetrate into cold-arid landscapes that had resulted from the deterioration of the climate.
The Red Apollo expanded its range to southern regions during the Ice Ages. In the interglacials, this fragmented into mountain-island occurrences after its previous habitats were narrowed with the return of trees and the gradual expansion of the forest. The southernmost and most isolated populations are therefore most at risk today as small populations are prone to genetic impoverishment (bottleneck effect) and negative demographic trends. In addition, climatic changes are more pronounced in the southernmost distribution areas and the niches in which it still occurs today could completely disappear in the future.
After a similar distribution of the butterfly as today existed in the Upper Pliocene, the ice ages in the subsequent Pleistocene had wiped out the populations there through the glaciers in the Alps and the ice sheets in Northern Europe and Northern Asia. Southern Europe and the east of the distribution area were not affected by the four ice ages Elbe (Günz), Elster (Mindel), Saale (Riss) and Vistula ice ages (northern Germany), or the Würme ice age (Alpine region). There they could survive the entire period and differentiate themselves more strongly and form consolidated subspecies. The strongest glaciation took place during the Saale or Riss Ice Age, when the entire Alps were glaciated into the foreland and western Russia and the Urals. In the subsequent Würm Ice Age around 115,000 to 10,000 years ago, the valleys of the Eastern Alps were free of ice. In the following warm period, in which we still live today, ideal conditions for the spread of the species formed through forest steppes with the food plants of the caterpillars. However, this period is too short for established new subspecies to develop, so these must be in the affected Classify areas as semi-subspecies.
Hazard and protection
The endangerment situation of this species is shown in some Red Lists of Endangered Species . The International Union for Conservation of Nature IUCN lists them as endangered (Vulnerable). The Red List of Germany sees it as threatened with extinction (Cat. 1), the Red List of Switzerland and also that of Austria shows the species as endangered (Cat. 3); the endangerment situation in the federal states of Austria is stated very differently and ranges from not endangered to extinct. The Apollo butterfly is critically endangered or critically endangered in many European regions.
The Red Apollo has been a nature reserve in Germany since 1936 . According to the Washington Convention on the Protection of Species , Appendix II, it is considered to be protected worldwide and is the only globally protected non-tropical butterfly species (as of 1990). In addition, this species is also listed in Appendix II of the Bern Convention , is also one of the few butterfly species in Appendix IV of the Fauna-Flora-Habitat Directive and is therefore designated as strictly protected. In Germany it is also classified as a national type of responsibility within the national strategy for biological diversity of the federal government.
In Germany, the Red Apollo is only found outside the Alps on the Moselle, the Swabian Alb and the Franconian Alb. In Baden-Württemberg , it lost all of over 60 sites around 1900 except for one in the Swabian Alb in 1988. After a decline to around a dozen moths at the end of the 1980s, the population of this occurrence has now recovered and stabilized, and a second population has even formed. It is no longer found in the Black Forest. Parnassius apollo ancile Fruhstorfer has been extinct in the Fichtel Mountains since 1909. Parnassius apollo posthumus Fruhstorfer 1925 has been extinct in the northern Franconian Forest and Saale Valley since 1905. In Bavaria it can be found in the Franconian Alb and there mainly in the Altmühltal and in the Bavarian Alps with a focus on the Berchtesgaden and Chiemgau Alps .
The Moselapollo was on the verge of extinction in the 1970s to the early 1980s due to the use of insecticides that were used in viticulture and sprayed with helicopters. The insecticides fell not only on the vineyards, but also on distant natural areas and killed the caterpillars. Land consolidation measures were also carried out during the same period , during which many of the old vineyard walls were removed with Sedum album . Unused vineyards fell into bush and the food plant disappeared through the shadows. Today it is forbidden to apply insecticides by helicopter. The habitats have been maintained since 1987 and kept free of bushes and perennials. The use of insecticides has declined sharply since the 1980s, because spraying is more targeted or, in the case of organic wine, no longer spraying at all. The populations have recovered and at airfields where only five to ten butterflies were seen in the early 1980s, 20 years later the figure was over 100. The deposits are advertised today in tourism and the approximately 7.5 kilometer long Apolloweg Valwig was set up for pedestrians in the local community of Valwig .
In Austria, Parnassius apollo cetius Fruhstorfer in particular is threatened as a subspecies in the lowlands in 1909, while the subspecies Parnassius apollo brittingeri Rebel & Rogenhofer 1893 in the mountains is significantly less endangered. The population on the Hohe Wand near Vienna is one of the populations of Parnassius apollo cetius that are still richest in individuals today . Occurrences outside the Alps, for example in Strudengau, have largely been extinct for many decades.
In France, Parnassius apollo meridionalis is extinct in the Vosges. Resettlement attempts failed here as well as in Forez and in the Massif de la Sainte-Baume . In contrast, a resettlement at the Puy de Dôme in the Massif Central was successful.
In Sweden the Red Apollo was widespread in the southern flatlands, today it can only be found on the east coast. In Finland, the decline began in the 1930s and 30 years later it had disappeared in many of the original areas. In recent years, the moth has spread again in the south-west of the country and has repopulated original areas with the islands between the island of Kemiö and the Hankoniemi peninsula . There seems to be a connection between the heavy metal contamination of the food crops and the extinction of the populations, since the caterpillars die when the food is contaminated. This has decreased in recent years and the caterpillars have better chances of survival again.
In Spain, the population of Parnassius apollo filabricus in the Sierra de los Filabres fell very sharply within 20 years, despite protection until 2005. The habitats in which the moths once flew by the thousands were either reforested with pine trees with the support of the nature conservation authorities or destroyed through intensive sheep grazing.
The main causes for the decline of the species are the destruction of the habitats by bushes or afforestation and the use of herbicides in agriculture and viticulture . Road and rail traffic also claims many victims, for example on the Moselle. Weak populations can be further weakened by natural predators such as tits and lizards, especially if additional nesting aids are offered for tits. The influence of foragers on weakened populations is not clear, but could have weakened threatened ones further. Today the danger posed by collectors is hardly relevant anymore. The species seems to be very sensitive to minor climatic changes.
- Edwin Möhn: Papilionidae XII: Parnassius apollo. Text . In: Erich Bauer and Thomas Frankenbach (eds.): Butterflies of the earth, day butterflies . tape 23 . Goecke & Evers, Keltern 2005, ISBN 3-937783-16-4 .
- Thomas C. Emmel: Wonderful and mysterious world of butterflies . Bertelsmann Lexikon-Verlag, Gütersloh and Berlin 1976, ISBN 3-570-00893-2 , p. 192 f .
- Wolfgang Wagner 2005–2019 Parnassius apollo
- ibid. Wolfgang Wagner 2005–2019
- Ibid. Wolfgang Wagner 2005–2019
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- VALENTINA TODISCO *, PAOLO GRATTON, DONATELLA CESARONI and VALERIO SBORDONI 2010: Phylogeography of Parnassius apollo: hints on taxonomy and conservation of a vulnerable glacial butterfly invader. Biological Journal of the Linnean Society, 2010, 101, 169-183. (PDF)
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- Naturschutzbund Österreich ( Memento of the original from September 22, 2007 in the Internet Archive ) Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice.
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- Karl Puchberger, Thoughts on the desertification of the butterfly fauna of the Strudengau , Steyr Entomologenrunde - Contributions to the knowledge of the insect fauna of Upper Austria, 1984
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- Species protection. (No longer available online.) In: Butterflies and Wine, 5/2007. NABU, archived from the original on May 15, 2007 ; Retrieved October 27, 2007 . Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice.
- Ministry for the Environment and Forests Rhineland-Palatinate (ed.): Apollofalter in Rhineland-Palatinate . 2003 ( PDF, 5 MB [accessed October 27, 2007]).
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