Dinaric Columbine ( Aquilegia dinarica in the wild in the sub-Adriatic Orjen)
The columbines ( Aquilegia ) form a genus of plants in the buttercup family (Ranunculaceae). The 70 to 75 species are mainly found in the temperate areas of the northern hemisphere . Varieties of some Aquilegia species are used as ornamental plants .
Columbine species are perennial (usually three to five years old) to perennial herbaceous plants . The richly branched root system forms slender, slightly woody rhizomes with a permanent taproot as a permanent organ. Over time, the plant expands to include the hypocotyl region above the root neck in the form of a thickened shoot base or corm that remains on or below the soil surface. This structure helps the stem axis during the winter. With constant growth over several seasons, side buds form next to the primary leaf crown, which form new growth axes. Several upright, mostly branched stems stand together on a plant .
The seedlings have two seed leaves ( cotyledons ). The leaves stand together in basal leaf rosettes. In addition, the somewhat smaller leaves are alternate and distributed in a spiral on the stem. However, these can also be completely absent as an adaptation to drier habitats or high mountain locations. The leaves are divided into a long petiole and a leaf blade. The leaf blades, which are pinnate to three parts, consist of lobed to divided leaflets . The edge of the leaflets is notched.
With the transition to flowering, the apical meristem transforms into an inflorescence . The flowers are terminal, sometimes solitary, but usually in two to ten zymous or doldigen monochasial or dichasile inflorescences together with foliage -like bracts . The hermaphroditic, radially symmetrical flowers have five leaf organs and are also arranged in five-hosted flowers. In the first whorl standing kronblattartigen sepals (sepals) that have a protruding feature in the attraction of pollinators. In the second whorl are arranged the petals, which are strongly differentiated by a backward nectar spur and contain nectaries in the spur . The length of these spurs varies enormously from 9 to 15 centimeters in Aquilegia longissima and the spurless Aquilegia ecalcarata . The species also vary in the length of the petal blade and the curvature of the spur. The colors of the bracts range from white to blue and yellow to red. The five free, short nailed sepals are spread out and 0.7 to 5.1 inches long. The five more or less upright, free petals are usually shorter than the sepals with less than 30 millimeters.
The many stamens are arranged in ten orthostiches , each with four to nine whorls. The flowers following the first bloom each have gradually decreasing numbers of stamen whorls. At the apical end of each orthostitch there is a new type of flower organ, the approximately seven scale-shaped, membranous staminodes . These sterile, flattened organs are found in all flowers regardless of their number of stamens. The staminodes consist of a central filament with a lateral lamina and are typically colorless. The ecological function of these organs is still debated, but it is evident that they will remain on the flower even after the other flower organs have fallen off; they remain as a surrounding ring on the carpel . One hypothesis is that these organs are equipped with mixtures of defensive substances against herbivores to provide protection in the early stages of fruiting. All Aquilegia species with the exception of Aquilegia jonesii have such staminodes. There are four to six free carpels in the center of the flower. The style is about half as long as the ovary.
The style is clearly recognizable from the cylindrical follicles , which are 3 to 26 millimeters long and narrow . Each follicle contains 10 to 36 seeds. The black, smooth seeds are narrow and obovate.
Ecology and evolution
The columbines belong to the original flowering plants and therefore have a relatively simple morphological blueprint.
The original Aquilegia species emerged from a central Asian distribution center around 6.18 to 6.51 million years ago. They form a monophyletic group ( monophyly ) both as a whole and in the individual areas of distribution .
The origin of the Aquilegia species is given for Europe to a period of 1.25 to 3.96 million years ago, for North America to 1.42 to 5.01 million years ago. Since there are no fossil remains of Aquilegia spec. are found, these dates are based on molecular genetic data ( molecular clock ). North America was settled only once via the land connection from Beringia, which was opened in the Pliocene (Bering Strait as a land connection opened in the geological period from 5.5 to 3.1 million years ago).
In Aquilegia TYPES there are usually hemicryptophytes .
The plant genus Aquilegia has long been important to the science of botany. The genus Aquilegia has turned out to be one of the most outstanding models for understanding the evolutionary history of the origins of flower organs and morphology in the parallel development of plant species and animal pollinators. While having Aquilegia TYPES directional adjustments completed their nectar spurs to different pollinators like hummingbird moth and bumblebees. Therefore, the lengths of the nectar spurs vary between 1 and 2 millimeters and from 10 to 12 centimeters, but the flower colors and the orientation have also been shown to be directly dependent on pollinating animals (bumblebee flowers are blue-violet, hummingbird flowers are red, hawk flowers are white or yellow) . They have adapted to a variety of different pollinators: hoverflies , bumblebees , swarms and hummingbirds .
In Eurasia and North America, have Aquilegia TYPES evolutionarily developed specifically different in a relatively short time, however: while in Eurasia Aquilegia - subspecies by adaptive radiation to different habitats advanced (forest, grassland, alpine locations), but the flowers modification remained relatively insignificant, In the new world, for example, the flower morphological adaptation to different pollinators took place. This is why the European Aquilegia species formed predominantly allopatric through relictic isolation (local endemics ), while the American species also formed sympatricly through barriers in the pollination mechanism. The flowers of the Eurasian columbines are still fixated on bumblebees, while the American species developed greater diversity and, in addition to species with bumblebee pollination, also developed forms that were wholly or predominantly based on hummingbirds ( Aquilegia flavescens , Aquilegia skinneri , Aquilegia formosa , Aquilegia canadensis , Aquilegia elegantula ) or swarm pollination are created.
The 70 to 75 Aquilegia species have their areas in the temperate areas of the northern hemisphere (circumboreal): in Eurasia and North America . The genus area extends north into the boreal zone and south into the mountains of northern Mexico and North Africa. The main distribution area is the Central Asian mountains in southern Siberia with around ten species. The species are distributed approximately one third each on the continents North America, Asia and Europe.
The Aquilegia species inhabit a multitude of different habitats , from oases in arid areas to alpine grasslands, rocky heaths or temperate forests, from the seashore to the slopes of the Himalayas , the Rocky Mountains or the Alps . They thrive from the desert ( Aquilegia skinneri or Aquilegia chrysanta ) to the high mountains ( Aquilegia dinarica or Aquilegia jonesii ). As generalists, certain species colonize a wide variety of habitats; so there is Aquilegia vulgaris in both rock, forest and grass vegetation. Specialists are then often adapted to rocky or mountainous locations, which applies to some of the rare endemic species of southern Europe and the Alps in the case of European columbines (for example Aquilegia alpina , Aquilegia dinarica , Aquilegia kitaibelii ).
European species complexes and sets of chromosomes
There are six Aquilegia species in Central Europe . On the basis of their morphological properties, they are divided into the groups of the Vulgaris complex and the Alpina complex. A combination of flower characteristics as well as leaf and stem hair are used to differentiate the European species taxonomically. The common columbine ( Aquilegia vulgaris ) and dark columbine ( Aquilegia nigricans ), which are difficult to separate from each other in southern Central Europe, can usually only be identified by the protrusion of the stamens, which either protrude from the petals or are enclosed by them, and the presence or absence of glandular hairiness can be clearly distinguished on the stem. In addition, the flower color of the dark columbine is purple-violet, and that of the common columbine is blue-violet. In southern Europe, however, the feature complexes are also clearer through features of the leaf segments: the dark columbine here has deeply incised lobes of the leaf segments. The Pleasant Columbine ( Aquilegia grata ), which is probably closely related to the Dark Columbine, has the shortest petals of all European Columbines . With 6 mm, the stamens also protrude far from the petals in the Pleasant Columbine. The black-violet columbine ( Aquilegia atrata ) with the 5 mm stamens protruding from the petals and the very dark flower color is also striking. The alpine columbine ( Aquilegia alpina ) with its particularly large blue flowers and the small-flowered columbine ( Aquilegia einseliana ), which is already part of the alpine complex of columbines, are also easily accessible . Since columbines generally have high demands on the water supply, they are represented in subtropical locations in southern Europe by special mountain clans. They are the Dinaric Columbine ( Aquilegia dinarica ), Aquilegia nikolicii , Aquilegia ottonis and the Kitaibel Columbine ( Aquilegia kitaibelliana ) that grow in limestone mountains above the tree line on moist spring niches or on shady rock ledges. However, they remain rare in the arid climates and mostly occur very locally.
To this day it is not possible to distinguish the more than twenty European columbines by genetic sequences, but genetic discriminants have been found for the American and Asian clans. Because of the close relationship of all Aquilegia species, even the most geographically distant species always remained fertile in infraspecific crossings . The genus therefore has no polyploid representatives. This means that all Aquilegia species and even infraspecific hybrids always remain diploid in their chromosome set. Because of these described as "religious" Diploidie all were Aquilegia - taxa ( "Species Flock" English.) With respect to a species flock set. For example, 2n = mostly 14, less often 16, 18 or 20 were found.
Systematics and distribution
The genus Aquilegia was established in 1753 by Carl von Linné in Species Plantarum , 1, p. 533. The scientific generic name Aquilegia is made up of two Latin word elements: aqua for water and legere for collect, i.e. water collector and refers to the nectar accumulated in the spurs with which pollinating insects are attracted.
There are about 70 to 75 species in the genus Aquilegia :
- Aquilegia afghanica (Brühl) T.Durand & BDJacks. : It occurs in Pakistan.
- Alpine Columbine ( Aquilegia alpina L. ): It thrives in Europe in the western Alps and in the Apennines .
- Aquilegia amurensis Kom . : It occurs in Siberia and in Russia's Far East .
- Black-violet Columbine ( Aquilegia atrata W.DJKoch ), Syn .: Aquilegia vulgaris subsp. atrata ( WDJKoch & Gaudin ): It thrives in Europe in the Western Alps and the Apennines .
- Aquilegia atrovinosa Popov & Gamajun. : It occurs in Kazakhstan and in the northern part of the Uighur autonomous region of Xinjiang .
- Gold Columbine ( Aquilegia aurea Janka ): It occurs in Bulgaria and North Macedonia .
- Aquilegia baluchistanica Qureshi & Chaudhri : It occurs in Pakistan.
- Aquilegia barbaricina Arrigoni & E. Nardi : This endemic occurs only in Sardinia .
- Aquilegia barnebyi Munz : It occurs in the US states of Utah and Colorado .
- Corsican Columbine ( Aquilegia bernardii Gren. & Godr. ): This endemic occurs only in Corsica .
- Bertoloni Columbine ( Aquilegia bertolonii Schott ): It occurs in southeastern France and northwestern Italy .
- Aquilegia borodinii Shishk . : It occurs in Siberia.
- Aquilegia brachyceras fish. & CAMey. ex Turcz. : It occurs in Siberia.
- Aquilegia brevistyla Hook. : It occurs in North America in Canada and in the northern United States.
- Japanese Columbine ( Aquilegia buergeriana Sieb. & Zucc. ): It occurs in Japan.
- Red Columbine ( Aquilegia canadensis L. ): It is common in North America in Canada and the USA.
- Aquilegia champagnatii Moraldo, Nardi & la Valva : It occurs in Italy.
- Aquilegia chaplinei Standl. ex Payson : It occurs in Texas and New Mexico.
- Aquilegia chitralensis Qureshi & Chaudhri : It occurs in Pakistan.
- Goldspur Columbine ( Aquilegia chrysantha A. Gray ): It is distributed from the USA to northern Mexico.
- Rocky Mountains Columbine ( Aquilegia caerulea E. James ): It thrives in the Rocky Mountains .
- Aquilegia colchica Kem.-Nath. : It only occurs in Georgia.
- Aquilegia × cultorum Bergmans
- Solitary Columbine ( Aquilegia desertorum ( MEJones ), A.Heller ): It thrives at altitudes of 2000 to 2500 meters in the US states of Arizona , New Mexico and Utah .
- Dinaric Columbine ( Aquilegia dinarica Beck ): It occurs on the Balkan Peninsula .
- Aquilegia discolor Levier & Leresche : This endemic occurs only in northern Spain.
- Aquilegia dumeticola Jord. : It occurs in Italy and Corsica, but is alsocountedby some authors to the common columbine ( Aquilegia vulgaris ).
- Spurless Columbine ( Aquilegia ecalcarata Maxim. ): It is common in China.
- Small-flowered Columbine ( Aquilegia einseleana F.W.Schultz ): It thrives in Europe in the Alps.
- Aquilegia elegantula Greene : It is distributed from the USA to Mexico.
- Aquilegia euchroma Rech. F. : It occurs in Afghanistan.
- Serpentine Columbine ( Aquilegia eximia Van Houtte ex Planch. ): It occurs only in California.
- Kuril Columbine ( Aquilegia flabellata Sieb. & Zucc. , Syn .: Aquilegia akitensis Huth ), home: Japan, Sakhalin , Kuril Islands , with the varieties:
- Yellowish Columbine ( Aquilegia flavescens S.Watson ), native to Canada, USA, at altitudes between 1300 and 3500 meters
- Beautiful Columbine ( Aquilegia formosa fish. Ex DC. ), Native to: Alaska, Canada, USA
- Fragrant Columbine ( Aquilegia fragrans Benth. ), Native to the Himalayas (Pakistan, western India), at altitudes between 2,400 and 3,600 meters
- Glandular Columbine ( Aquilegia glandulosa Fisch. Ex Link ): It is common in Siberia, Central Asia, Mongolia and in the Chinese Xinjiang .
- Aquilegia gracillima Rech. F. : It occurs in Afghanistan.
- Pleasant Columbine ( Aquilegia grata Zimmeter ): This endemic occurs in Montenegro only in the Orjen .
- Aquilegia × helenae Arends (= Aquilegia flabellata × Aquilegia coerulea )
- Aquilegia hinckleyana Munz : It occurs in Texas.
- Aquilegia incurvata P.K.Hsiao : It occurs in China in the provinces of Gansu , Shaanxi and Sichuan .
- Lime Columbine or Jones Columbine ( Aquilegia jonesii Parry ): It is common in Canada and the USA.
- Aquilegia kareliniana C.A.Mey. ex Trautv. : It occurs in Kazakhstan, in Kyrgyzstan and in India in Uttar Pradesh .
- Kitaibel Columbine ( Aquilegia kitaibelii Schott ): It occurs in the former Yugoslavia .
- Aquilegia kurramensis Qureshi & Chaudhri : It occurs in Pakustan.
- Aquilegia lactiflora Kar. & Kir. : It occurs in Kazakhstan, Kyrgyzstan, Turkmenistan and the Chinese province of Xinjiang.
- Wyoming Columbine ( Aquilegia laramiensis A.Nelson ): It only thrives at altitudes of 2000 to 2500 meters in the US state of Wyoming.
- Aquilegia litardierei Briq. : This endemic occurs only in Corsica .
- Long-tailed Columbine ( Aquilegia longissima A.Gray ex S.Watson ), native to USA, Mexico
- Aquilegia maimanica Rech. F. : It occurs in Afghanistan.
- Aquilegia micrantha Eastw. : It occurs in the US states of Utah, Colorado and Arizona.
- Aquilegia microcentra Rech. F. : It occurs in Afghanistan.
- Aquilegia moorcroftiana Wall. ex Royle : It occurs in Afghanistan, India, Nepal, Pakistan and Xizang .
- Dark Columbine ( Aquilegia nigricans Baumg. , Syn .: Aquilegia vulgaris subsp. Nigricans ( Baumg. ) Domin ): It occurs in Europe mainly in the southeast.
- Aquilegia nikolicii Niketić & Cikovac : It occurs in Serbia , Bosnia and Herzegovina and Montenegro .
- Snow Columbine ( Aquilegia nivalis Falc. Ex BDJacks. ): It occurs only in Kashmir .
- Aquilegia nugorensis Arrigoni & E. Nardi : It only occurs in Sardinia .
- Aquilegia nuragica Arrigoni & E. Nardi : It only occurs in Sardinia.
- Caucasian Columbine ( Aquilegia olympica Boiss. ), Native to: Caucasus, Transcaucasia, Turkey, northern Iran
Aquilegia ottonis Orph. ex Boiss. : The three subspecies occur in Europe in the Apennines and on the Balkan Peninsula:
- Balkan Columbine ( Aquilegia ottonis subsp. Amaliae ( Heldr. Ex Boiss. ) Strid ): It occurs in Albania , in the former Yugoslavia and in Greece.
- Aquilegia ottonis subsp. Ottonis : It occurs in central Italy and Greece.
- Aquilegia ottonis subsp. taygetea ( Orph. ) Strid : It occurs only in southern Greece.
- Aquilegia oxysepala Trautv. & CAMey. , Home: Eastern Siberia, Northern China, Manchuria , Korea, Japan
- Aquilegia pancicii sword : It occurs in the former Yugoslavia.
- Aquilegia parviflora Ledeb. : It occurs in Asiatic Russia, Mongolia and China.
- Californian Columbine ( Aquilegia pubescens Coville ): It only thrives on rocks at altitudes of 3000 to 4000 meters in California.
- Aquilegia pubiflora Wall. ex Royle : It occurs in Afghanistan, Pakistan, India and Nepal.
Pyrenees Columbine ( Aquilegia pyrenaica DC. ), Europe: Spain and France, with four subspecies:
- subsp. cazorlensis ( Heywood ) Galeano & Rivas Mart. : This endemic occurs in southeastern Spain only in the province of Caén .
- subsp. discolor ( Levier & Leresche) Pereda & Laínz : It occurs in Spain.
- subsp. guarensis (Losa) Rivas Mart. (Syn .: Aquilegia aragonensis Willk. ): This endemic occurs in southeastern Spain only in the Sierra de Guara near Huesca.
- subsp. pyrenaica : It occurs in Spain and France.
- Aquilegia rockii Munz : It thrives in mixed forests and on roadsides at altitudes of 2500 to 3500 meters in southeastern Tibet and in the Chinese provinces of southwestern Sichuan and northeastern Yunnan.
- Low Columbine ( Aquilegia saximontana Rydb. ): It thrives at altitudes of 3300 to 4400 meters only in the US state of Colorado.
- Rock Columbine ( Aquilegia scopulorum Tidestr. ): It thrives at altitudes of 2000 to 3500 meters in the US states of Wyoming, Nevada and Utah.
- Shockley Columbine ( Aquilegia shockleyi Eastw. ): It thrives at altitudes of 1200 to 2700 meters in the US states of California and Nevada.
- Siberian Columbine ( Aquilegia sibirica Lam. ): It occurs in Kazakhstan , Siberia, Mongolia and in the Uighur autonomous region of Xinjiang .
- Aquilegia skinneri Hook. : It occurs in the US state of New Mexico.
- Aquilegia × stuartii Balf. f. (= Aquilegia glandulosa × Aquilegia olympica )
- Meadow rue-leaved Columbine ( Aquilegia thalictrifolia Schott & Kotschy ): It occurs under overhanging limestone cliffs on damp limestone mulm together with other endemic species in northern Italy (Lake Garda area, Vicentine Alps ).
- Aquilegia transsilvanica Schur : It occurs in Europe in the Carpathian Mountains , Romania and the Ukraine.
- Aquilegia tuvinica I.M.Vassiljeva
- Aquilegia turczaninovii Kamelin & Gubanov
- Aquilegia vestinae pfenn. & DMMoser : It was first described in 2002 and only occurs on Lake Garda.
- Aquilegia vicaria Nevski
Green-flowered Columbine ( Aquilegia viridiflora Pall. ): There are at least two varieties:
- Aquilegia viridiflora var. Atropurpurea (Willd.) Trevir. (Syn .: Aquilegia atropurpurea Willd. ): It occurs in Siberia, Mongolia , Inner Mongolia and the Chinese provinces in Hebei, southern Liaoning, eastern Qinghai, eastern Shandong and Shanxi.
- Aquilegia viridiflora var. Viridiflora Pall. : It occurs in Japan, Siberia, Mongolia , Inner Mongolia and the Chinese provinces of Gansu, Hebei, Heilongjiang, Hubei, Jilin, Liaoning, Ningxia, Shaanxi, Shandong and Shanxi.
- Aquilegia viscosa Gouan : There are three subspecies in Spain and France:
- Common Columbine ( Aquilegia vulgaris L. ): It occurs in Europe, Macaronesia and North Africa .
- Aquilegia yabeana Kitag. (Syn .: Aquilegia oxysepala var. Yabeana (Kitag.) Munz ): It thrives on the edges of forests and on grass slopes in Inner Mongolia and in the Chinese provinces of Hebei, Henan, Hubei, western Liaoning, southern Shaanxi and Shanxi.
The following species are no longer included in Aquilegia :
- Aquilegia adoxoides (DC.) Ohwi → Semiaquilegia adoxoides (DC. ) Makino
- Aquilegia anemonoides Willd. → Paraquilegia anemonoides (Willd. ) OEUlbr.
The derivation of the common German name Akelei is unclear. But probably the German common name Akelei is borrowed from the Latin aquilegia. In the Old High German glosses , forms such as agaleia or ageleia (since the 10th century) can be found. In Hildegard of Bingen is the plant acoleia, ackeleia, agleia, akuleye the Middle Low German. In the vernacular the word has been changed many times, e.g. B. in Akelchen (Thuringia), Aggerlei, Aggerleine (Palatinate), Aglije (Lucerne, Zurich), Hagleie (Schaffhausen), Hakeleden, Hakelehnen (Mecklenburg), Gakeilei (based on Gaggel 'Ei' in children's language (Lower Hesse, Rhenish) ) or Klei (e) (Niederrheinisch).
Many folk names refer to the shape of the nodding flowers, such as bells, bells, bells, blue bells (common), gypsy bells (Gailtal / Carinthia), devil bells (Lenggries / Upper Bavaria), imperial bells (Giant Mountains), sugar bells (Thurgau), bellflower ( widespread), bell piece (Swabian Alb) or bell pink (Anhalt).
Other popular names related to the shape of the flower are Pausewängel (Sächs, Felsengebirge), Stellhäfele (actually an earthen cooking vessel with feet) (Aachern / Baden), Kessel (Mittenwald / Upper Bavaria), Stanitzelblume (Bavarian Stanizl 'paper bag') (Knittelfeld / Styria), Manselblume (Swiss. Manse 'rock sleeves with lace') (Aargau), fool's hats (e.g. Lörrach / Baden, Canton St. Gallen), Kapuzinerchappe (s), - Hüetli (Canton St. Gallen), Pfaffenkäpple (Achkarren / Baden), Plumphose (Kt. Schaffhausen), Schlotterhose (St. Gallen), Schwizerhose (Aargau), Pants (n) lätzli (Aargau), Frae (n) schüehli (Küsnacht / Schwyz), Fünf Vögerl zsam (Eastern Styria), Tauberln (South Moravia) or Gugerschen (Schönhengstgau, Sudetenland).
References to the dark flower color can be found in the names ink bell (Thuringian Forest, Thurgau) and Truarbliemli (funeral flowers because they are also planted in rural cemeteries) (Grindelwald / Bern).
The columbine is also called Kaiserblume (Albendorf / Riesengebirge), Hernblume (Eifel), Zaniggele, Zinäggele or Süniggele (based on 'Sanikel') (Schaffhausen).
Elf shoe, gypsy bells, devil bells, imperial bells and fool's caps are also common names of the Columbine.
Another name, Agelblume , was used by the noble sisterhood of the Agelblume in Königsberg in Bavaria , which existed until the Reformation and was located in Königsberg. The Columbine stood for the modesty that it should remind the Sisters of the Agelblume.
Symbolism and cultural meaning
The Columbine has probably been an ornamental plant in European gardens since the late Middle Ages. The common columbine was used in various forms in medicine in the Middle Ages and early modern times. Due to the symbolism attributed to it, it can also be found on numerous medieval panel paintings. In ancient times the Columbine was consecrated to the fertility goddess Freya , later it was assigned to the Virgin Mary. The seeds have been used as an aphrodisiac since the Middle Ages. It was even said that the aphrodisiac powers of the columbine seeds could be transmitted with mere touch. Rätsch wrote that if you ground the seeds into a fine powder that you smeared on your palms and then touched a woman, she was immediately sexually aroused. In symbolism, the Columbine stands on the one hand for humility, and it symbolizes the Holy Spirit, life force, overcoming earthly limitations, comprehensive salvation, triumph, redemption, trinity and the praise of God. On the other hand, it stands for the sexual power of men, for seduction and love. It was and is still used as a grave plant.
Varieties of some Aquilegia TYPES (eg Aquilegia alpina , Aquilegia atrata , Aquilegia caerulea , Aquilegia canadensis , Aquilegia chrysantha , Aquilegia elegantula , Aquilegia flabellata , Aquilegia formosa , Aquilegia longissima , Aquilegia saximontana , Aquilegia skinneri , Aquilegia viridiflora and Aquilegia vulgaris ) and hybrids (for example, McKana hybrids) are used as ornamental plants . Depending on the type and variety, they are used very differently as bedding plants, in rock gardens or as cut flowers .
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