Columbines


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Columbines
Dinaric Columbine (Aquilegia dinarica in the wild in the sub-Adriatic Orjen)

Dinaric Columbine ( Aquilegia dinarica in the wild in the sub-Adriatic Orjen)

Systematics
Eudicotyledons
Order : Buttercups (Ranunculales)
Family : Buttercup Family (Ranunculaceae)
Subfamily : Isopyroideae
Tribe : Isopyreae
Genre : Columbines
Scientific name
Aquilegia
L.

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 .

description

Illustration of the Alpine Columbine ( Aquilegia alpina )
Bloom of a columbine in detail
Columbine pollen (400 ×)

Vegetative characteristics

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.

Generative characteristics

Flower diagram of the genus Aquilegia . Two innovations of the genus as well as a closely related one of the buttercup family are given by the five-organ flowers. The development of staminodes and nectar spurs are more recent developments in evolutionary terms.
Five immature follicles of the common columbine ( Aquilegia vulgaris )

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.

Bumblebees are the characteristic pollinators of the Aquilegia vulgaris group in Eurasia.
Numerous North American columbines specialize in pollinating hawks.
Aquilegia shockleyi belongs to the group of North American species that specialize in hummingbird pollination.

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 wettability of the leaf surface is poor. Water rolls off in droplets, as can also be observed with lotus flowers , and it takes dirt particles adhering to the surface with it ( lotus effect ).

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.

Aquilegia species can not spread over greater distances due to the lack of certain promoting mechanisms of propagation of the small seeds . As a result, they often only occur more frequently locally.

Occurrence

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.

Alpine Columbine ( Aquilegia alpina )

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

The Pleasant Columbine is an endemic European forest acre from western Montenegro with particularly prominent stamens . The stamina protrude 6 mm from the petals in the species. To date, it has not been adequately investigated whether this characteristic is due to the biological background of the flower.

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.

Black-violet Columbine
( Aquilegia atrata )
Bertoloni Columbine ( Aquilegia bertolonii )
Red Columbine
( Aquilegia canadensis )
Goldspur Columbine ( Aquilegia chrysantha )
Rocky Mountains Columbine
( Aquilegia coerulea )
Flowers and follicles of the Dinaric Columbine ( Aquilegia dinarica )
Dwarf Columbine ( Aquilegia flabellata var. Pumila )
Beautiful Columbine
( Aquilegia formosa )
The spurs are best seen from the back: Fragrant Columbine ( Aquilegia fragrans )
Pyrenees Columbine ( Aquilegia pyrenaica )
Siberian Columbine ( Aquilegia sibirica )

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.

The genus Aquilegia belongs to the subtribe Isopyrinae from the tribe Isopyreae in the subfamily Isopyroideae within the family Ranunculaceae .

There are about 70 to 75 species in the genus Aquilegia :

Illustration from Curtis's Magazine , 1935, plate 9405 from Aquilegia nikolicii (as Aquilegia grata ). Previously, the species shown was mistakenly held as Aquilegia grata . It is managed as an independent species. The specimen shown was propagated in the Royal Botanical Garden in 1929 by William Bertram Turrill from seeds collected in ex-Yugoslavia.

The following species are no longer included in Aquilegia :

The hybrid Aquilegia × maruyamana

Common names

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.

Flower of the cultivar 'Red Star'

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.

use

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 .

swell

literature

  • Robert Nold: Columbines: Aquilegia, Paraquilegia, and Semiaquilegia. Timber Press, Portland 2003, ISBN 0-88192-588-8 .
  • Jaakko Jalas, Juha Suominen: Atlas florae europaeae. Volume 8: Nymphaeaceae to Ranunculaceae. Helsinki 1989, ISBN 951-9108-07-6 , pp. 225-235.
  • Eckehart J. Jäger, Friedrich Ebel, Peter Hanelt, Gerd K. Müller: Excursion flora from Germany. Volume 5: Herbaceous ornamental and useful plants. Spectrum Academic Publishing House. Berlin / Heidelberg 2008, ISBN 978-3-8274-0918-8 .
  • Werner Greuter , HM Burdet, G. Long: MED Checklist. Volume 4, Conservatoire et Jardin botaniques, Genève 1989, ISBN 2-8277-0154-5 , pp. 395-397.
  • Harald Riedl: The Aquilegia vulgaris group in Austria. In: Austrian Botanical Journal. Volume 114, 1967, ISSN  0029-8948 , pp. 94-100.
  • Harald Riedl: Aquilegia at Tropicos.org. In: Flora of Pakistan . Missouri Botanical Garden, St. Louis, pp. 12-23. In: YJ Nasir: Ranunculaceae. Volume 193, Department of Botany, University of Karachi, Karachi 1991.
  • Alan T. Whittemore: Aquilegia - the same text online as the printed work , In: Flora of North America Editorial Committee (Ed.): Flora of North America North of Mexico , Volume 3 - Magnoliidae and Hamamelidae , Oxford University Press, New York and Oxford, 1997, ISBN 0-19-511246-6 .
  • Fu Dezhi, Orbélia R. Robinson: Aquilegia , p. 278 - online with the same text as the printed work , In: Wu Zhengyi, Peter H. Raven & Deyuan Hong (Eds.): Flora of China , Volume 6 - Caryophyllaceae through Lardizabalaceae , Science Press and Missouri Botanical Garden Press, Beijing and St. Louis, 2001, ISBN 1-930723-05-9 .

Individual evidence

  1. ^ Plate (lithograph) and text by Aquilegia alpina in Curtis's Botanical Magazine .
  2. ^ A b Elena M. Kramer: Aquilegia : A New Model for Plant Development, Exology, and Evolution. In: Annual Review of Plant Biology. Volume 60, 2009, pp. 261-278.
  3. a b c Jesus M. Bastida, Julio M. Alcantara, Pedro J. Rey, Pablo Vargas, Carlos M. Herrera: Extended phylogeny of Aquilegia: the biogeographical and ecological patterns of two simultaneous but contrasting radiations. In: Plant Systematics and Evolution. Volume 284, 2010, pp. 171–185 (online, PDF, 630 kB)
  4. Rolf Froböse: When frogs fall from the sky. The craziest natural phenomena. Wiley-VCH Verlag, Weinheim 2007, ISBN 978-3-527-31659-5 , p. 170.
  5. a b Elena M. Kramer, Scott A. Hodges: Aquilegia as a model system for the evolution and ecology of petals. In: Philosophical Transaction of the Royal Society, Biological Sciences. Volume 365, No. 1539, February 12, 2010, pp. 477-490. (abstract) doi: 10.1098 / rstb.2009.0230
  6. Aquilegia species: distribution, length of the flower spurs and trunk length of the pollinators .
  7. ^ A b Alan T. Whittemore: Aquilegia - the same text online as the printed work , In: Flora of North America Editorial Committee (Ed.): Flora of North America North of Mexico , Volume 3 - Magnoliidae and Hamamelidae , Oxford University Press, New York and Oxford, 1997. ISBN 0-19-511246-6
  8. Gustav Hegi (Ed.): Illustrated flora of Central Europe. Volume 3.1., 1958, pp. 128-140.
  9. Harald Riedel 1967: The Aquilegia vulgaris group in Austria. Austrian Botanical Journal, 114/1 (1967), 94-100. (PDF)
  10. Harald Riedel 1967: p. 96
  11. Marjan Niketić, Pavle Cikovac, Vladimir Stevanović 2013: Taxonomic and nomenclature notes on Balkan columbines ( Aquilegia L., Ranunculaceae). In: Bulletin of the Natural History Museum Belgrade, 6: 33-42. (PDF)
  12. Harald Riedel 1967: p. 95
  13. Marjan Niketić, Pavle Cikovac, Vladimir Stevanović 2013: Here pp. 37–40 (PDF)
  14. ^ S. Fior, M. Li, B. Oxelman, R. Viola, SA Hodges, L. Ometto, C. Varotto: Spatiotemporal reconstruction of the Aquilegia rapid radiation through next-generation sequencing of rapidly evolving cpDNA regions. In: New Phytologist. Volume 198, Issue 2, 2013, pp. 579-592.
  15. ^ Aquilegia at Tropicos.org. In: IPCN Chromosome Reports . Missouri Botanical Garden, St. Louis
  16. First publication scanned at biodiversitylibrary.org
  17. ^ Aquilegia at Tropicos.org. Missouri Botanical Garden, St. Louis, accessed January 6, 2014.
  18. Explanations of the generic name by the University of Duisburg.
  19. a b c d e f g h i j k l m n o p q r s t u v w x Aquilegia in the Germplasm Resources Information Network (GRIN), USDA , ARS , National Genetic Resources Program. National Germplasm Resources Laboratory, Beltsville, Maryland. Retrieved February 11, 2017.
  20. a b c d e f g h Fu Dezhi, Orbélia R. Robinson: Aquilegia , p. 278 - online with the same text as the printed work , In: Wu Zhengyi, Peter H. Raven & Deyuan Hong (eds.): Flora of China , Volume 6 - Caryophyllaceae through Lardizabalaceae , Science Press and Missouri Botanical Garden Press, Beijing and St. Louis, 2001, ISBN 1-930723-05-9 .
  21. Flora Europaea online : Aquilegia , last viewed on September 26, 2011
  22. a b c d e f g h i j Walter Erhardt , Erich Götz, Nils Bödeker, Siegmund Seybold: The great pikeperch. Encyclopedia of Plant Names. Volume 2: Types and Varieties. Eugen Ulmer, Stuttgart (Hohenheim) 2008, ISBN 978-3-8001-5406-7 .
  23. a b M. Niketić, P. Cikovac, V. Stevanovic: Taxonomic and nomenclature notes on Balkan columbines (Aquilegia L., Ranunculaceae). In: Bulletin of the Natural History Museum Belgrade. Volume 6, 2013, pp. 33-42. PDF
  24. Hans Pitschmann, Herbert Reisigl, Hugo Schiechtl: Flora of the Southern Alps from Lake Garda to Lake Como. 2nd Edition. Verlag Gustav Fischer, Stuttgart 1965, p. 84.
  25. Gustav Hegi (Ed.): Illustrated flora of Central Europe. Volume 3, Part 1, 1958, pp. 134-135.
  26. http://blumenssprache.blogspot.de/2013/01/akelei.html
  27. http://www.gabryon.de/de/blog/blumen-bedeutung-sprache
  28. Gordon Cheers (Ed.): Botanica. The ABC of plants. 10,000 species in text and images . Könemann Verlagsgesellschaft, 2003, ISBN 3-8331-1600-5 , p. 101-103 .

Web links

Commons : Akeleien ( Aquilegia )  - album with pictures, videos and audio files
Wiktionary: Columbine  - explanations of meanings, word origins, synonyms, translations