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Ilex aquifolium
Species of flowering plant in the family Aquifoliaceae From Wikipedia, the free encyclopedia
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Ilex aquifolium, the holly, common holly, English holly, European holly, or occasionally Christmas holly, is a species of flowering plant in the family Aquifoliaceae, native to western and southern Europe, northwest Africa, and southwest Asia.[2][3][4][5] It is regarded as the type species of the genus Ilex, which by association is also called "holly". It is an evergreen tree or shrub found, for example, in shady areas of forests of oak and in beech hedges. In the British Isles it is one of very few native hardwood evergreen trees. It has a great capacity to adapt to different conditions and is a pioneer species that repopulates the margins of forests or clearcuts.
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I. aquifolium can exceed 10 m in height, but is often found at much smaller heights, typically 2–3 m (6.6–9.8 ft) tall and broad, with a straight trunk and pyramidal crown, branching from the base. It grows slowly and does not usually fully mature due to cutting or fire. It can live 500 years, but usually does not reach 100.
Ilex aquifolium is the species of holly long associated with Christmas, and previously the Roman festival of Saturnalia. Its glossy green prickly leaves and bright red berries (produced only by the female plant) are represented in wreaths, garlands and cards wherever Christmas is celebrated. It is a subject of music and folklore, especially in the British tradition. It is also a popular ornamental shrub or hedge, with numerous cultivars in a range of colours.
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Description
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In exceptional cases Ilex aquifolium can grow to 25 m (82 ft) in height,[6] but more typically is 3 to 15 m (10 to 49 ft) tall and often only grows into a shrub rather than reaching tree stature.[7] It has a woody stem as wide as 40–80 cm (16–31 in), rarely 100 cm (39 in) or more, in diameter.[4] The bark is smooth and silver-grey, but with age will develop small fissures.[7]
The leaves are 5–12 cm long and 2–6 cm broad; they are evergreen, lasting about five years, and are dark green on the upper surface and lighter on the underside, oval, leathery, shiny, and about 5 to 9 cm long. In the young and in the lower limbs of mature trees, the leaves have three to five sharp spines on each side, pointing alternately upward and downward, while leaves of the upper branches in mature trees lack spines.[4][5] In rare cases, a tree may lack spines even on its lower leaves.[8]
The flowers are white, four-lobed, and pollinated by bees. Holly is dioecious, meaning that there are male plants and female plants. The sex cannot be determined until the plants begin flowering, usually between 4 and 12 years of age. In male specimens, the flowers are yellowish and appear in axillary groups. In the female, flowers are isolated or in groups of three and are small and white or slightly pink, and consist of four petals and four sepals partially fused at the base.
The fruit only appears on female plants, which require male plants nearby to fertilise them. The fruit is a drupe (stone fruit), about 6–10 mm in diameter, a bright red or bright yellow, which matures around October or November; at this time they are very bitter due to the ilicin content[9] and so are rarely eaten until late winter after frost has made them softer and more palatable. They are eaten by rodents, birds and larger herbivores. Each fruit contains 3 to 4 seeds which do not germinate until the second or third spring.[10]
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Taxonomy
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Ilex aquifolium was given its scientific name in 1753 by Linnaeus. He placed it in the genus Ilex which is classified in the family Aquifoliaceae. More than 100 botanical varieties of Ilex aquifolium have been described by botanists, but none of them are accepted names according to Plants of the World Online. Together with 23 species and other names it has a total of 145 synonyms. Though for brevity none of the varieties or forms of I. aquifolium are presented here.[11]
Names
The name holly can be used to mean any species in the genus Ilex, but originally referred to Ilex aquifolium and continues to be used to mean this species particularly in addition to names such as European holly. Holly is a shortend form of the Old English holegn or holen.[12] In Scotland holly is still sometimes called by the older names hollin or hollen.[13] It is also known by the common names common holly and English holly.[14][15]
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Distribution and habitat
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Today, holly is found in western Asia and Europe in the undergrowth of oak forest and beech forest in particular, although at times it can form a dense thicket as the dominant species. It requires moist, shady environments, found within forests or in shady slopes, cliffs, and mountain gorges.[2][3][4] The species is shifting northwards in Norway and towards to the northeast in Denmark and Germany. Holly grows from sea level to elevations as high as 600 m (2,000 ft).[1]
Holly is a common species in its range and widespread. Because of this it is a species of least-concern according to the IUCN, however there is a decline in mature individuals and its population is severely fragmented. Also, local populations may face threats. In the country of Sweden only one native tree remains.[1]
Along the west coast of the United States and Canada, from California to British Columbia,[16] non-native English Holly has proved very invasive, quickly spreading into native forest habitat, where it thrives in shade and crowds out native species. It has been placed on the Washington State Noxious Weed Control Board's monitor list, and is a Class C invasive plant in Portland.[17][18][19]
During the Cenozoic Era, the Mediterranean region, Europe, and northwest Africa had a wetter climate and were largely covered by laurel forests. Holly was a typical representative species of this biome, where many current species of the genus Ilex were present. With the drying of the Mediterranean Basin during the Pliocene, the laurel forests gradually retreated, replaced by more drought-tolerant sclerophyll plant communities. The modern Ilex aquifolium resulted from this change.[clarification needed][citation needed] Most of the last remaining laurel forests around the Mediterranean are believed to have died out approximately 10,000 years ago at the end of the Pleistocene.
Ecology
Holly is a rugged pioneer species that prefers relatively moist areas, and tolerates frost as well as summer drought. The plant is common in the garrigue and maquis and is also found in deciduous forest and oak forest.
Pure stands of hollies can grow into a labyrinth of vaults in which thrushes and deer take refuge, while smaller birds are protected among their spiny leaves. After the first frost of the season, holly fruits become soft and fall to the ground serving as important food in its native regions for winter birds at a time of scarce resources.
The flowers are attractive as nectar sources for insects such as bees, wasps, flies, and small butterflies. The commonly-encountered pale patches on leaves are due to the leaf-mine insect Phytomyza ilicis.
It is an invasive species on the West Coast of Canada[16] and the United States as well as in Hawaii.[19][20]
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Epigenetics
Holly is well known in epigenetics. Some cultivars have smooth leaf edges, or both smooth and prickly leaf edges on the same plant. In response to stress these cultivars can produce leaves with more prickles.[21]
Cultivation
Ilex aquifolium is widely grown in parks and gardens in temperate regions.[22] Hollies are often used for hedges; the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping.[23]
AGM cultivars
Numerous cultivars have been selected,[24] of which the following have gained the Royal Horticultural Society's Award of Garden Merit:[25]
Ilex × altaclerensis
The hybrid Ilex × altaclerensis was developed at Highclere Castle in Hampshire, England, in 1835, a cross between I. aquifolium and the tender species I. perado. The following cultivars have gained the RHS AGM:
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Chemistry and toxicity
Holly berries contain alkaloids, theobromine, saponins, caffeic acid, and a yellow pigment, ilixanthin.[9][40][41] The berries are generally regarded as toxic to humans.[40][41]
Uses
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Between the thirteenth and eighteenth centuries, before the introduction of turnips, Ilex aquifolium was cultivated for use as winter fodder for cattle and sheep. Less spiny varieties of holly were preferred, and in practice the leaves growing near the top of the tree have far fewer spines, making them more suitable for fodder.
Ilex aquifolium was once among the traditional woods for Great Highland bagpipes before tastes turned to imported dense tropical woods such as cocuswood, ebony, and African blackwood.
In culture
In the British Isles common holly has been used in rituals and as a symbol both before and after the coming of Christianity in similar ways to the use of evergreen plants during midwinter in many cultures.[42] Branches of holly continue to be used in place of palm fronds on Palm Sunday in some areas of Europe. It is also often used in Christmas decorations, particularly in Britain, its evergreen leaves a symbol of life continuing during the winter season.[43] Though there is no traditional prohibition against cutting boughs or twigs, cutting down a holly tree is believed to bring bad luck.[44] Like other spiny plants, the common holly is believed to protect against malicious witchcraft.[45]
The name of the hamlet of Hulver in Suffolk is the same at the Middle English name of the species, though in 1997 there were only 18 holly trees there. There is a similar lack of the namesake trees in Hollybush near the border between Worcestershire and Herefordshire.[46]
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References
External links
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