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Ilex aquifolium

Ilex aquifolium
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Aquifoliales
Family: Aquifoliaceae
Genus: Ilex
Species:
I. aquifolium
Binomial name
Ilex aquifolium
Distribution map of Ilex aquifolium
Ilex aquifolium

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.

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.

Description

Frosted foliage and berries
European holly flowers; male above, female below (leaves cut to show flowers more clearly)

Ilex aquifolium grows to 10–25 m (33–82 ft) tall with a woody stem as wide as 40–80 cm (16–31 in), rarely 100 cm (39 in) or more, in diameter. 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]

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[6] 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.[7]

Distribution

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]

Along the west coast of the United States and Canada, from California to British Columbia,[8] 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.[9][10][11]

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[8] and the United States as well as in Hawaii.[11][12]

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.[13]

Cultivation

One of several variegated cultivars

Ilex aquifolium is widely grown in parks and gardens in temperate regions.[14] Hollies are often used for hedges; the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping.[15]

AGM cultivars

Numerous cultivars have been selected,[16] of which the following have gained the Royal Horticultural Society's Award of Garden Merit:[17]

  • I. aquifolium[18]
  • 'Amber' (female)[19]
  • 'Argentea Marginata'[20]
  • 'Ferox Argentea'[21]
  • 'Golden Queen'[22]
  • 'Handsworth New Silver'[23]
  • 'J.C. van Tol'[24]
  • 'Madame Briot'[25]
  • 'Pyramidalis'[26]
  • 'Silver Queen'[27]

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:

Chemistry, toxicity, and uses

Holly berries contain alkaloids, theobromine, saponins, caffeic acid, and a yellow pigment, ilixanthin.[6][32][33] The berries are generally regarded as toxic to humans.[32][33] Accidental consumption may occur by children or pets attracted to the bright red berries. The berries are emetic, possibly due to the compound ilicin. The leaves of yerba mate, also in the genus Ilex, are used to make a caffeinated beverage[34] called mate or Paraguayan tea.[35]

Other uses

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.

References

  1. ^ Barstow, M.; Khela, S. (2018). "Ilex aquifolium". IUCN Red List of Threatened Species. 2018: e.T202963A68067360. doi:10.2305/IUCN.UK.2018-1.RLTS.T202963A68067360.en. Retrieved 19 November 2021.
  2. ^ a b Flora Europaea: Ilex aquifolium
  3. ^ a b Med-Checklist: Ilex aquifolium
  4. ^ a b c Rushforth, K. (1999). Trees of Britain and Europe. Collins ISBN 0-00-220013-9.
  5. ^ a b Flora of NW Europe: Ilex aquifolium Archived 2009-06-09 at the Wayback Machine
  6. ^ a b Hoppe, Heinz, A. (1975). Drogenkunde. W. de Gruyter. ISBN 3-13-566001-X.((cite book)): CS1 maint: multiple names: authors list (link)
  7. ^ Peterken, G. F.; Lloyd, P. S. (November 1967). "Ilex aquifolium L.". Journal of Ecology. 55 (3): 841–858. Bibcode:1967JEcol..55..841P. doi:10.2307/2258429. JSTOR 2258429.
  8. ^ a b "Ring in the holiday season and reduce the spread of invasive species! -". Invasive Species Council of British Columbia | ISCBC Plants & Animals. 2020-02-10. Retrieved 2020-12-27.
  9. ^ "Washington State Noxious Weed Control Board".
  10. ^ "English Holly | Invasive Species Council of British Columbia | ISCBC Plants & Animals". Archived from the original on 2015-09-06. Retrieved 2015-08-14.
  11. ^ a b "Ilex aquifolium (English holly)". California Invasive Plant Council. Archived from the original on 2016-08-16. Retrieved 2016-10-02.
  12. ^ "English Holly - Ilex aquifolium". King County, Washington. Retrieved 2011-01-30.
  13. ^ "#bioPGH blog – to Prickle or Not to Prickle: Holly Leaves and Epigenetics | Phipps Conservatory and Botanical Gardens | Pittsburgh PA".
  14. ^ RHS A-Z encyclopedia of garden plants. United Kingdom: Dorling Kindersley. 2008. p. 1136. ISBN 978-1405332965.
  15. ^ Northumbria Police: Security starts at the Garden Gate
  16. ^ Huxley, A., ed. (1992). New RHS Dictionary of Gardening. Macmillan ISBN 0-333-47494-5.
  17. ^ "AGM Plants - Ornamental" (PDF). Royal Horticultural Society. July 2017. p. 43. Retrieved 3 March 2018.
  18. ^ "RHS Plant Selector Ilex aquifolium AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-07-04.
  19. ^ "RHS Plant Selector Ilex aquifolium 'Amber' (f) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-07-04.
  20. ^ "RHS Plant Selector Ilex aquifolium 'Argentea Marginata' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-07-04.
  21. ^ "RHS Plant Selector Ilex aquifolium 'Ferox Argentea' (m/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-10.
  22. ^ "RHS Plant Selector Ilex aquifolium 'Golden Queen' (m/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  23. ^ "RHS Plant Selector Ilex aquifolium 'Handsworth New Silver' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  24. ^ "RHS Plant Selector Ilex aquifolium 'J.C. van Tol' (f) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  25. ^ "RHS Plant Selector Ilex aquifolium 'Madame Briot' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  26. ^ "RHS Plant Selector Ilex aquifolium 'Pyramidalis' (f) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  27. ^ "RHS Plant Selector Ilex aquifolium 'Silver Queen' (m/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  28. ^ "RHS Plant Selector Ilex × altaclerensis 'Belgica Aurea' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  29. ^ "RHS Plant Selector Ilex × altaclerensis 'Camelliifolia' (f) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  30. ^ "RHS Plant Selector Ilex × altaclerensis 'Golden King' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  31. ^ "RHS Plant Selector Ilex × altaclerensis 'Lawsoniana' (f/v) AGM / RHS Gardening". Apps.rhs.org.uk. Retrieved 2020-09-16.
  32. ^ a b Leikin, Jerrold Blair; Frank P. Paloucek (2002). Poisoning & Toxicology Handbook, Third Edition. Hudson, Ohio USA: Lexi-Comp Inc. p. 80. ISBN 978-1-930598-77-5.
  33. ^ a b Turner, Nancy J.; P. von Aderkas (2009). The North American Guide to Common Poisonous Plants and Mushrooms. Timberpress. p. 210. ISBN 978-0-88192-929-4.
  34. ^ Dellacassa, Cesio et al. Departamento de Farmacognosia, Facultad de Química, Universidad de la República, Uruguay, Noviembre: 2007, pages 1–15
  35. ^ Grieve, M. "Holly". Botanical.com. Retrieved 2014-02-26.
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Ilex aquifolium
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