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| status = LC
| status_system = IUCN3.1
| status_ref = <ref name="iucn status 19 November 2021">{{cite iucn |author=Barstow, M. |author2=Khela, S. |
|genus = Ilex
|species = aquifolium
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|range_map_caption = Distribution map of ''Ilex aquifolium''
}}
[[File:Ilex aquifolium in the Tashkent Botanical Garden.jpg|thumb|''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 (botany)|family]] Aquifoliaceae, [[native plant|native]] to western and southern Europe, northwest Africa, and southwest Asia.<ref name=fe>Flora Europaea: [http://rbg-web2.rbge.org.uk/cgi-bin/nph-readbtree.pl/feout?FAMILY_XREF=&GENUS_XREF=Ilex&SPECIES_XREF=aquifolium&TAXON_NAME_XREF=&RANK= ''Ilex aquifolium'']</ref><ref name=mc>Med-Checklist: [http://ww2.bgbm.org/mcl/PTaxonDetail.asp?NameId=1263&PTRefFK=1273 ''Ilex aquifolium'']</ref><ref name=rushforth>Rushforth, K. (1999). ''Trees of Britain and Europe''. Collins {{ISBN|0-00-220013-9}}.</ref><ref name=fnwe>Flora of NW Europe: [http://ip30.eti.uva.nl/BIS/flora.php?selected=beschrijving&menuentry=soorten&id=3088 ''Ilex aquifolium''] {{webarchive|url=https://web.archive.org/web/20090609025123/http://ip30.eti.uva.nl/BIS/flora.php?selected=beschrijving&menuentry=soorten&id=3088 |date=2009-06-09 }}</ref> It is regarded as the [[type species]] of the genus ''[[Holly|Ilex]]'', which by association is also called "holly".
''I. aquifolium'' can exceed 10 m in height, but is often found at much smaller heights, typically {{convert|2-3 |m|ft|abbr=on}}
▲'''''Ilex aquifolium''''', the '''holly''', '''common holly''', '''English holly''', '''European holly''', or occasionally '''Christmas holly''', is a [[species]] of [[flowering plant]] in the [[family (botany)|family]] Aquifoliaceae, [[native plant|native]] to western and southern Europe, northwest Africa, and southwest Asia.<ref name=fe>Flora Europaea: [http://rbg-web2.rbge.org.uk/cgi-bin/nph-readbtree.pl/feout?FAMILY_XREF=&GENUS_XREF=Ilex&SPECIES_XREF=aquifolium&TAXON_NAME_XREF=&RANK= ''Ilex aquifolium'']</ref><ref name=mc>Med-Checklist: [http://ww2.bgbm.org/mcl/PTaxonDetail.asp?NameId=1263&PTRefFK=1273 ''Ilex aquifolium'']</ref><ref name=rushforth>Rushforth, K. (1999). ''Trees of Britain and Europe''. Collins {{ISBN|0-00-220013-9}}.</ref><ref name=fnwe>Flora of NW Europe: [http://ip30.eti.uva.nl/BIS/flora.php?selected=beschrijving&menuentry=soorten&id=3088 ''Ilex aquifolium''] {{webarchive|url=https://web.archive.org/web/20090609025123/http://ip30.eti.uva.nl/BIS/flora.php?selected=beschrijving&menuentry=soorten&id=3088 |date=2009-06-09 }}</ref> It is regarded as the [[type species]] of the genus ''[[Holly|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]] [[hedge]]s. In the British Isles it is one of very few native 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 {{convert|2-3 |m|ft|abbr=on}} tall and broad, with a straight [[Trunk (botany)|trunk]] and pyramidal [[crown (botany)|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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The [[flower]]s are white, four-lobed, and pollinated by [[bee]]s. Holly is [[Glossary of botanical terms#dioecious|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 bud|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<ref name=Hoppe>{{cite book | first=Heinz, A. |last=Hoppe |date=1975 |title=Drogenkunde |publisher=W. de Gruyter |isbn=3-13-566001-X }}</ref> 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 [[seed]]s
==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.<ref name = fe/><ref name=mc/><ref name=rushforth/>
Along the west coast of the United States and Canada, from California to British Columbia,<ref name="BC-ISC-holiday" /> 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.<ref>{{Cite web | url=http://www.nwcb.wa.gov/weeds/english-holly | title=Washington State Noxious Weed Control Board}}</ref><ref>{{cite web |url=http://bcinvasives.ca/invasive-species/identify/invasive-species/invasive-plants/english-holly/ |title=
During the [[Cenozoic Era]], the Mediterranean region, Europe, and northwest Africa had a wetter climate and were largely covered by [[laurel forest]]s. Holly was a typical representative species of this [[biome]], where many current species of the genus ''Ilex'' were present. With the [[Messinian salinity crisis|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.{{clarify|reason=does this refer to the modern distribution or the modern species?|date=February 2017}}{{citation needed|date=February 2017}} 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]].
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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 shrubland|maquis]] and is also found in deciduous forest and oak forest.
Pure stands of hollies can grow into a labyrinth of vaults in which [[thrush (bird)|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.
It is an [[invasive species]] on the West Coast of Canada<ref name="BC-ISC-holiday" /> and the United States as well as in Hawaii.<ref name="Cal-IPC"/><ref>{{cite web| url=http://www.kingcounty.gov/environment/animalsAndPlants/noxious-weeds/weed-identification/english-holly.aspx| title=English Holly - ''Ilex aquifolium''| publisher=[[King County, Washington]]|
==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.<ref>{{cite web | url=https://www.phipps.conservatory.org/blog/detail/biopgh-blog-to-prickle-or-not-to-prickle-holly-leaves-and-epigenetics | title=#bioPGH blog – to Prickle or Not to Prickle: Holly Leaves and Epigenetics | Phipps Conservatory and Botanical Gardens | Pittsburgh PA }}</ref>
==Cultivation==
[[File:2014-08-29 13 46 38 Variegated English Holly at the Pinelands Preservation Alliance headquarters in Southampton Township, New Jersey.JPG|thumb|One of several variegated cultivars]]
''Ilex aquifolium'' is widely grown in parks and gardens in [[temperateness|temperate]] regions.<ref>{{cite book|title=RHS A-Z encyclopedia of garden plants|year=2008|publisher=Dorling Kindersley|location=United Kingdom|isbn=978-1405332965|page=1136}}</ref> Hollies are often used for [[hedge (barrier)|hedges]]; the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping.<ref>Northumbria Police: [http://ww2.northumbria.police.uk/ePolicing/Web/WMS.nsf/CrimeReductionContentDocs/CRD000301 Security starts at the Garden Gate]</ref>
===AGM cultivars=== Numerous cultivars have been selected,<ref name=rhs>Huxley, A., ed. (1992). ''New RHS Dictionary of Gardening''. Macmillan {{ISBN|0-333-47494-5}}.</ref> {{div col|colwidth=
*''I. aquifolium''<ref name = RHSPF>{{cite web|url=https://www.rhs.org.uk/Plants/9087/Ilex-aquifolium/Details |title=RHS Plant Selector Ilex aquifolium AGM / RHS Gardening |publisher=Apps.rhs.org.uk |
*'Amber' (female)<ref>{{cite web|url=https://www.rhs.org.uk/Plants/97772/Ilex-aquifolium-Amber-(f)/Details |title=RHS Plant Selector Ilex aquifolium 'Amber' (f) AGM / RHS Gardening |publisher=Apps.rhs.org.uk |
*'Argentea Marginata'<ref>{{cite web|url=https://www.rhs.org.uk/Plants/91514/Ilex-aquifolium-Argentea-Marginata-(f-v)/Details |title=RHS Plant Selector Ilex aquifolium 'Argentea Marginata' (f/v) AGM / RHS Gardening |publisher=Apps.rhs.org.uk |
*'Ferox Argentea'<ref>{{cite web|url=
*'Golden Queen'<ref>{{cite web|url=
*'Handsworth New Silver'<ref>{{cite web|url=
*'J.C. van Tol'<ref>{{cite web|url=
*'Madame Briot'<ref>{{cite web|url=
<!-- AGM withdrawn*'Myrtifolia Aurea Maculata'<ref>{{cite web|url=http://apps.rhs.org.uk/plantselector/plant?plantid=3133 |title=RHS Plant Selector Ilex aquifolium 'Myrtifolia Aurea Maculata' (m/v) AGM / RHS Gardening |publisher=Apps.rhs.org.uk |
*'Pyramidalis'<ref>{{cite web|url=
*'Silver Queen'<ref>{{cite web|url=
{{div col end}}
===''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 ''[[Ilex perado|I. perado]]''. {{div col|colwidth=30em}}
*'Belgica Aurea'<ref>{{cite web|url=
*'Camelliifolia'<ref>{{cite web|url=
*'Golden King'<ref>{{cite web|url=
*'Lawsoniana'<ref>{{cite web|url=
{{div col end}}
Holly berries contain [[alkaloid]]s,
▲==Chemistry, toxicity, tea and traditional uses==
▲Holly berries contain [[alkaloid]]s, [[caffeine]], and [[theobromine]], [[saponins]], [[caffeic acid]], and a yellow [[pigment]], ilixanthin.<ref name=Hoppe/><ref name=handbook>{{cite book
|last = Leikin
|first = Jerrold Blair
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|url = https://books.google.com/books?id=bmGY5APFfFQC&pg=RA1-PA210
}}
</ref> The berries are generally regarded as [[toxicity|toxic]] to humans.<ref name=handbook/><ref name=guide/> Accidental consumption may occur by children or pets attracted to the bright red berries. The berries are [[emetic]], possibly due to the compound
==Other uses==
Between the thirteenth and eighteenth centuries, before the introduction of [[turnip]]s, ''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
==References==
{{Reflist|30em
<ref name="BC-ISC-holiday">{{cite web | title=Ring in the holiday season and reduce the spread of invasive species! - | website=Invasive Species Council of British Columbia {{!}} ISCBC Plants & Animals | date=2020-02-10 | url=http://bcinvasives.ca/news-events/news-from-iscbc/ring-in-the-holiday-season | access-date=2020-12-27}}</ref>
}}
==External links==
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[[Category:Garden plants of Europe]]
[[Category:Ornamental trees]]
[[Category:Plants described in 1753]]
[[Category:Taxa named by Carl Linnaeus]]
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