Aquilegia

Balkan columbine (Aquilegia ottonis subsp. Amaliae )

The columbines (Aquilegia ) are a genus of flowering plants in the family Ranunculaceae ( Ranunculaceae ). The approximately 70 to 75 species are distributed mainly in the temperate regions of the northern hemisphere. Varieties of some Aquilegia species are used as ornamental plants.

• 8.1 Literature
• 8.2 Notes and references

Description

Vegetative characteristics

Aquilegia species are perennial (usually three to five years ) to perennial herbaceous plants. The richly branched root system forms slender, slightly woody rhizomes with permanent taproot as Überdauerungsorgan. Over time, the plant expanded to the hypocotyl region above the root collar in the form of a thickened shoot base or corms, which remains at or below the soil surface. This structure helps the stem axis in the winter. With steady growth over several seasons away, the new axes of growth form in addition to the primary leaf crown lateral buds form. At one plant, several erect, usually branched stems together.

The seedlings have two seed leaves ( cotyledons ). The leaves are borne in basal rosettes. In addition, some smaller leaves are alternate and distributed spirally on the stem. However, this can be used as adaptation to drier habitats or high altitude locations also completely absent. The leaves are divided into a long petiole and leaf blade one. The one -to three- fold in three parts feathery leaf blades consist of lobed to divided leaflets. The edge of the leaflets is notched.

Generative features

With the transition to flowering, the apical meristem is transformed into an inflorescence. The flowers are terminal, sometimes individually but usually zymösen in two to ten or doldigen monochasialen or dichasilen inflorescences together with foliage leaf-like bracts. The hermaphroditic, flowers have radial symmetry five leaf organs and are ordered in five economic time flowers. In the first whorl are the kronblattartigen sepals ( sepals ) having a protruding feature in the attraction of pollinators. In the second whorl strongly differentiated by a backward nectar spur the petals ( petals ) are arranged, which included the spur nectary. The length of these spurs varies enormously 9-15 centimeters in Aquilegia longissima and the spur -free Aquilegia ecalcarata. However, the species also vary in length of Kronblattspreite 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 and from 0.7 to 5.1 inches long. The five more or less upright, free petals are usually shorter than the sepals less than 30 millimeters.

The many stamens are arranged in ten Ortho stitches to four to nine whorls. The first flowering following flowers have in each case gradually decreasing numbers of Staubblattwirteln. At the apical end of each ortho stitches there is a novel floral organ, the flaky about seven, membranous staminodes. These sterile, flattened bodies found in all flowers regardless of their stamen number. The staminodes consist of a central filament with lateral lamina and are typically colorless. The ecological function of these organs is still being discussed before, but it is obvious that they will still remain on the flower when the other floral organs have fallen; they remain as enclosing wreath at the carpel. One hypothesis is that these institutions are equipped with mixtures of defense substances against herbivores in order to provide protection in the early stage of fruit development. All Aquilegia species with the exception of Aquilegia jonesii possess such staminodes. There are four to six free carpels in the center of the flower. The stylus is about half as long as the ovary.

At the at a length of 3 to 26 mm narrow, cylindrical Balgfrüchten the pen is clearly visible. Each pod contains 10 to 36 seeds. The smooth black seeds are narrow and obovate.

Ecology and Evolution

The columbines are among the original flowering plants and therefore have a relative simple morphological blueprint.

The original Aquilegia species have emerged from a Central Asian distribution center million years ago, about 6.18 to 6.51. They form a monophyletic group ( monophyly ) both overall and in individual distribution priorities.

The emergence of Aquilegia species is thereby given to Europe on a period of 1.25 to 3.96 million years ago, before today, for North America at 1.42 to 5.01 million years ago before today. Since no fossil remains of Aquilegia sp. be found, these datings based on molecular genetic data (molecular clock ). The colonization of North America took place only once via the open in the Pliocene land bridge of Beringia ( Bering Strait land bridge as in the geological period from 5.5 to 3.1 million years ago opened before today).

In Aquilegia species are mostly to hemicryptophytes.

The wettability of the sheet surface is small. Water beads in drops from, how it can be also observed in lotus flowers, and does, on the surface of particles of dirt with (lotus effect).

For a long time the plant genus Aquilegia in the science of botany is important. The genus Aquilegia has been found for the understanding of evolution of historical lineage of floral organs and morphology in the parallel development of plant species and animal pollinators as one of the outstanding models. This Aquilegia species have undergone directional adjustments to their nectar spurs to different pollinators such as Hummingbird moth and bumble bees. Therefore vary the lengths of nectar spurs between 1 to 2 millimeters and 10 to 12 inches, but the flower color as the orientation have been found to be directly dependent on pollinating animals ( Hummel flowers are blue-violet, hummingbird flowers red, enthusiast flowers white or yellow). They have thereby adapted to a variety of different pollinators: hoverflies, bumblebees, and hummingbirds enthusiast.

In Eurasia and North America, the Aquilegia species have evolutionarily but it is different in different in a relatively short time: While in Eurasia Aquilegia subtypes through adaptive radiation in different habitats (forest, grassland, alpine locations) further developed, but the flowers modification remained relatively insignificant, so the blütenmorphologische adaptation to different pollinators was held in the new world. Therefore, the European Aquilegia species have predominantly allopatric formed by relict insulation (local endemics ), the U.S., however, also sympatric through barriers in the pollination mechanism. So the flowers of Eurasian columbines are still fixated on bumblebees, while the American species evolved greater variety and next species with Hummel pollination also developed forms which wholly or mainly on Hummingbird (Aquilegia flavescens, Aquilegia skinneri, Aquilegia formosa, Aquilegia canadensis, Aquilegia elegantula ) are created or moth - pollination.

Aquilegia species can not propagate due to the lack of certain promotional propagation mechanisms of small seeds over longer distances. They occur thereby often on only locally common.

Occurrence

The 70 to 75 Aquilegia species have their areas in the temperate regions of the northern hemisphere ( zirkumboreal ): in Eurasia and North America. Here, the generic area extends north into the boreal zone and south to the mountains of northern Mexico and North Africa. Distribution area are the Central Asian Mountains in southern Siberia with about ten species. The species are distributed each about one-third of the continents North America, Asia and Europe.

The Aquilegia species colonize a variety of habitats, from oases in arid areas to alpine grasslands, rocky pagans or temperate forests, from the coast to the slopes of the Himalayas, the Rockies or the Alps. They thrive from the desert (Aquilegia Aquilegia skinneri or Chrysanta ) to high mountains (Aquilegia Aquilegia Dinarica or jonesii ). Certain species inhabit a variety of habitats than generalists; Aquilegia vulgaris then found both in rock, forest and grass vegetation. Specialists are then often adapted to rocky or mountainous locations, which is particularly true for the European columbines on some of the rare endemic species of southern Europe and the Alps ( Aquilegia alpina example, Dinarica Aquilegia, Aquilegia kitaibelii ).

European species complexes and sets of chromosomes

In Central Europe, six Aquilegia species occur. They are divided into the groups of the vulgaris complex as well as the Alpina complex based on morphological characteristics.

However, it is still not possible to distinguish more than twenty European columbines on genetic sequences to the American and Asian families but genetic discriminants were found. Due to the close relationship of all Aquilegia species even the geographically most distant species were always fertile in infraspecific crosses. Thus, the genus has no plyoploiden representative. That is, all Aquilegia species and infraspecific hybrids themselves remain in their diploid set of chromosomes. Because of this diploidy "religious" described as well as all Aquilegia taxa ( " species flock " engl.) were related to a species flock. For example, were mostly 2n = 14, rarely 16, 18 or 20 have been found.

System

The genus Aquilegia was erected in 1753 by Carl Linnaeus in Species Plantarum, 1, p 533. The scientific genus name Aquilegia is made up of two Latin word elements: aqua for water and for casual gathering, ie water collector ( in ) and refers to the accumulated nectar in the spurs, are lured with the pollinating insects.

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

In the genus Aquilegia, there are about 70 to 75 species. Species found in Europe, are marked with (Europe).

• Aquilegia afghanica (Brühl ) T.Durand & BDJacks.
• Alpine Columbine (Aquilegia alpina L.) (Europe: Western Alps and Apennines )
• Aquilegia amurensis Kom
• Black and Purple Columbine (Aquilegia atrata WDJKoch; Syn: . A. vulgaris subsp atrata ( WDJKoch ) Gaudin ) (Europe: Alps and Apennines )
• Aquilegia atrovinosa Popov & Gamajun.
• Gold columbine (Aquilegia aurea Janka ) (Europe: Bulgaria, Macedonia )
• Aquilegia baluchistanica Qureshi & Chaudhri
• Aquilegia barbaricina Arrigoni & E.Nardi (Europe: Sardinia)
• Aquilegia barnebyi Munz, Origin: USA (Utah, Colorado)
• Corsican Columbine (. . Aquilegia bernardii Gren & Godr ) (Europe: Corsica)
• Bertoloni columbine (Aquilegia bertolonii Schott ) (Europe: South East France, northwestern Italy )
• Aquilegia borodinii Schischk.
• Aquilegia Brachyceras fish. & C.A.Mey. ex Turcz.
• Aquilegia brevistyla Hook.
• Japanese Columbine (. Aquilegia Sieb & Zucc buergeriana. ), Origin: Japan
• Red columbine (Aquilegia canadensis L.), Origin: USA, Canada
• Aquilegia champagnatii Moraldo et al (Europe: Italy)
• Aquilegia chaplinei Standlschmaus. ex Payson
• Aquilegia chitralensis Qureshi & Chaudhri
• Gold Spur Columbine (Aquilegia chrysantha A. Gray ), Origin: USA, Northern Mexico
• Rocky Mountain columbine (Aquilegia coerulea E.James ), Origin: Rocky Mountains
• Aquilegia colchica Kem. - Nath.
• Aquilegia × cultorum Bergmans
• Desert columbine (Aquilegia desertorum ( MEJones ) A.Heller ), Origin: USA ( Arizone, New Mexico, Utah, at altitudes 2000-2500 meters )
• Dinaric columbine (Aquilegia Dinarica Beck) (Europe: Balkan Peninsula)
• Aquilegia discolor Levier & Laresche, Origin: North Spain
• Aquilegia dumeticola Jord. , Occurs in Italy and Corsica, but is also expected by some authors for ordinary columbine (A. vulgaris)
• Spur lots columbine (Aquilegia ecalcarata Maxim. ), Origin: China
• Small-flowered Columbine (Aquilegia einseleana FWSchultz ) (Europe: Alps)
• Aquilegia elegantula Greene, Origin: USA, Mexico
• Aquilegia euchromatic Rech f
• Serpentine columbine (Aquilegia eximia Van Houtte ex Planch. )
• Kuril columbine (Aquilegia Sieb & Zucc flabellata, Syn: .. Aquilegia akitensis Huth ), Origin: Japan, Sakhalin, Kuril Islands, with the varieties: var flabellata
• Dwarf columbine (Aquilegia flabellata var pumila ( Huth ) Kudo )

• Balkan columbine (Aquilegia ottonis subsp. Amaliae ( Heldr. ex Boiss. ) Strid ), comes only in Albania, the former Yugoslavia and in Greece
• Subsp. ottonis, occurs only in central Italy and in Greece
• Subsp. Taygetea ( Orph. ) Strid, occurs in southern Greece

• Subsp. cazorlensis ( Heywood ) Galiano & Rivas- Martínez, occurs only in southeast Spain in the province of Caen ago
• Subsp. discolor ( Levier & Laresche ) Pereda % Lainz, occurs only in Spain
• Subsp. guarensis ( Losa ) Rivas- Martínez ( syn.. Aquilegia aragonensis Willk ), occurs only in southeast Spain in the Sierra de Guara in Huesca ago
• Subsp. pyrenaica, is found in Spain and France

• Subsp. hirsutissima ( Timb. - Lagr. ) Breistr. , occurs only in the south of France
• Subsp. montsicciana ( font cross ) O.Bolòs & Vigo, occurs only in Spain
• Subsp. viscosa, occurs only in the south of France

Are no longer expected to Aquilegia species:

• Aquilegia adoxoides ( DC.) Ohwi → Semiaquilegia adoxoides ( DC.) Makino
• Aquilegia anemonoides Willd. → Paraquilegia anemonoides ( Willd.) O.E.Ulbr.

Trivial names

The derivation of the German name is unclear. But probably the German Common name Columbine is borrowed from the Latin aquilegia. In the Old High German glosses forms such agaleia or ageleia are to be found ( since the 10 century). In Hildegard of Bingen called the plant acoleia, ackeleia, agleia, akuleye in Middle Low German. In the vernacular, the word has often been converted, eg in Akelchen ( Thuringia), Aggerlei, Agger leash ( Pfalz), Aglije ( Lucerne, Zurich ), Hagleie (Schaffhausen), Hakeleden, Hake backrest (Mecklenburg), Gakeilei (based on Gaggel ' egg ' in child language ( Lower Hesse, Rhine) ) or clay ( e) (Lower Rhine ).

Many common names refer to the shape of the nodding blooms, as bells, Glöckerl, Bells, Blue Bells ( widespread), Zigeunderglocken ( Gail Valley / Carinthia ), Devil bells ( Lenggries / Upper Bavaria), Emperor Bell (Giant Mountains ), sugar bells ( Thurgau ), Bellflower ( widely used ), bells piece ( Swabian Alb) or bells Rosa ( Anhalt).

Other common names the flower shape in question are Pausewängel ( Saxon, Rocky Mountains ), Stllhäfele (actually an earthenware cooking vessel with feet ) ( Aachern / Baden), boiler ( Mittenwald / Upper Bavaria), Stanitzelblume ( Bavarian Stanizl ' paper bag ') ( Knittel Field / Styria), Manselblume ( schweiz. Manse ' skirt with lace sleeves ') ( Aargau ), dunce caps (eg Lörrach / Baden, Canton of St. Gallen), Kapuzinerchappe (s) - Hüetli (Canton St. Gallen), Pfaffenkäpple ( Achkarren / Baden), Plump pants (canton Schaffhausen ), Schlotter pants (St. Gallen), Schwizerstrasse pants (Aargau), pants ( n ) lätzli (Aargau), Frae (s) schüehli ( Küsnacht / Schwyz ), Five Vögerl zsam ( Styria ) or Tauberln (South Moravia).

References to the dark flower color can be found in the name of ink cap ( Thuringian Forest, Thurgau ) and Truarbliemli ( Trauerblümlein since planted on rural cemeteries ) (Grindelwald / Bern ).

In addition, the columbine flower still Emperor ( Albendorf / Giant Mountains ), Hernblume (Eifel ), Zaniggele, Zinäggele or Süniggele is (based on ' Sanikel ') ( Schaffhausen ) called.

Elf Shoe, gypsy bells, bells Devil, Emperor bells and dunce caps are also common names of Columbine.

Another name Agelblume, used the nobles Sisterhood of the Agelblume in Königsberg in Bayern, which existed until the Reformation and was a resident of Königsberg. The Columbine stand for modesty, to which they should remember the Sisters of Agelblume.

Use

Varieties of some Aquilegia species ( for example, 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. They are widely used depending on the type and variety as bedding plant, in the rock garden or as a cut flower.

Swell