Ilex Aquifolium
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''Ilex aquifolium'', the holly, common holly, English holly, European holly, or occasionally Christmas holly, is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
in the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Aquifoliaceae,
native Native may refer to: People * Jus soli, citizenship by right of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Native Americans (disambiguation) In arts and entert ...
to western and southern Europe, northwest Africa, and southwest Asia.Flora Europaea
''Ilex aquifolium''
/ref>Med-Checklist
''Ilex aquifolium''
/ref>Rushforth, K. (1999). ''Trees of Britain and Europe''. Collins .Flora of NW Europe
''Ilex aquifolium''
It is regarded as the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the genus ''
Ilex ''Ilex'' (), or holly, is a genus of over 570 species of flowering plants in the family (biology), family Aquifoliaceae, and the only living genus in that family. ''Ilex'' has the most species of any woody dioecious angiosperm genus. The speci ...
'', which by association is also called "holly". It is an
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
or
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
found, for example, in shady areas of forests of
oak An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ''L ...
and in
beech Beech (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engle ...
hedge A hedge or hedgerow is a line of closely spaced shrubs and sometimes trees, planted and trained to form a barrier or to mark the boundary of an area, such as between neighbouring properties. Hedges that are used to separate a road from adjoini ...
s. 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 tall and broad, with a straight trunk and pyramidal
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, partic ...
, 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 Saturnalia is an ancient Roman festival and holiday in honour of the god Saturn, held on 17 December of the Julian calendar and later expanded with festivities through to 23 December. The holiday was celebrated with a sacrifice at the Temple ...
. 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

''Ilex aquifolium'' grows to tall with a woody stem as wide as , rarely or more, in diameter. The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
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. The
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s are white, four-lobed, and pollinated by bees. Holly is
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
, 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 In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
is a
drupe In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or '' pyrena'') of hardened endocarp with a seed (''kernel'') ...
(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 and so are rarely eaten until late winter after
frost Frost is a thin layer of ice on a solid surface, which forms from water vapor in an above-freezing atmosphere coming in contact with a solid surface whose temperature is below freezing, and resulting in a phase change from water vapor (a gas) ...
has made them softer and more palatable. They are eaten by rodents, birds and larger herbivores. Each fruit contains 3 to 4
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s which do not germinate until the second or third spring.


Distribution

Today, holly is found in western Asia and Europe in the undergrowth of
oak An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ''L ...
forest and
beech Beech (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engle ...
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. Along the west coast of the United States and Canada, from California to British Columbia, 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. During the
Cenozoic Era The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configu ...
, the Mediterranean region, Europe, and northwest Africa had a wetter climate and were largely covered by
laurel forest Laurel forest, also called laurisilva or laurissilva, is a type of subtropical forest found in areas with high humidity and relatively stable, mild temperatures. The forest is characterized by broadleaf tree species with evergreen, glossy and elo ...
s. Holly was a typical representative species of this
biome A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...
, where many current species of the genus ''Ilex'' were present. With the drying of the Mediterranean Basin during the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58drought A drought is defined as drier than normal conditions.Douville, H., K. Raghavan, J. Renwick, R.P. Allan, P.A. Arias, M. Barlow, R. Cerezo-Mota, A. Cherchi, T.Y. Gan, J. Gergis, D.  Jiang, A.  Khan, W.  Pokam Mba, D.  Rosenfeld, J. Tierney, an ...
-tolerant
sclerophyll Sclerophyll is a type of vegetation that is adapted to long periods of dryness and heat. The plants feature hard leaf, leaves, short Internode (botany), internodes (the distance between leaves along the stem) and leaf orientation which is paral ...
plant communities. The modern ''Ilex aquifolium'' resulted from this change. 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 The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
.


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 Garrigue or garigue ( ), also known as phrygana ( el, φρύγανα , n. pl.), is a type of low scrubland ecoregion and plant community in the Mediterranean forests, woodlands, and scrub biome. It is found on limestone soils in southern Fra ...
and
maquis Maquis may refer to: Resistance groups * Maquis (World War II), predominantly rural guerrilla bands of the French Resistance * Spanish Maquis, guerrillas who fought against Francoist Spain in the aftermath of the Spanish Civil War * The network ...
and is also found in deciduous forest and oak forest. Pure stands of hollies can grow into a labyrinth of vaults in which
thrushes The thrushes are a passerine bird family, Turdidae, with a worldwide distribution. The family was once much larger before biologists reclassified the former subfamily Saxicolinae, which includes the chats and European robins, as Old World flycat ...
and
deer Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...
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 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 ''Phytomyza ilicis'', the holly leaf miner, is a leaf mining fly in the family Agromyzidae, whose larvae burrow into leaves of the holly tree (''Ilex aquifolium'') leaving characteristic pale trails or leaf mines. Description Adults of ''P. ili ...
''. It is an
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
on the West Coast of Canada and the United States as well as in Hawaii.


Cultivation

''Ilex aquifolium'' is widely grown in parks and gardens in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
regions. Hollies are often used for hedges; the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping.


AGM cultivars

Numerous cultivars have been selected,Huxley, A., ed. (1992). ''New RHS Dictionary of Gardening''. Macmillan . of which the following have gained the
Royal Horticultural Society The Royal Horticultural Society (RHS), founded in 1804 as the Horticultural Society of London, is the UK's leading gardening charity. The RHS promotes horticulture through its five gardens at Wisley (Surrey), Hyde Hall (Essex), Harlow Carr (Nort ...
's
Award of Garden Merit The Award of Garden Merit (AGM) is a long-established annual award for plants by the British Royal Horticultural Society (RHS). It is based on assessment of the plants' performance under UK growing conditions. History The Award of Garden Merit ...
:- *''I. aquifolium'' *'Amber' (female) *'Argentea Marginata' *'Ferox Argentea' *'Golden Queen' *'Handsworth New Silver' *'J.C. van Tol' *'Madame Briot' *'Pyramidalis' *'Silver Queen'


''Ilex'' × ''altaclerensis''

The hybrid ''Ilex'' × ''altaclerensis'' was developed at
Highclere Castle Highclere Castle is a Grade I listed country house built in 1679 and largely renovated in the 1840s, with a park designed by Capability Brown in the 18th century. The estate is in Highclere in Hampshire, England, about south of Newbury, ...
in Hampshire, England, in 1835, a cross between ''I. aquifolium'' and the tender species '' I. perado''. The following cultivars have gained the RHS AGM:- *'Belgica Aurea' *'Camelliifolia' *'Golden King' *'Lawsoniana'


Chemistry, toxicity, tea and traditional uses

Holly berries contain
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
s,
caffeine Caffeine is a central nervous system (CNS) stimulant of the methylxanthine class. It is mainly used recreationally as a cognitive enhancer, increasing alertness and attentional performance. Caffeine acts by blocking binding of adenosine t ...
,
theobromine Theobromine, also known as xantheose, is the principal alkaloid of ''Theobroma cacao'' (cacao plant). Theobromine is slightly water-soluble (330 mg/L) with a bitter taste. In industry, theobromine is used as an additive and precursor to s ...
,
saponins Saponins (Latin "sapon", soap + "-in", one of), also selectively referred to as triterpene glycosides, are bitter-tasting usually toxic plant-derived organic chemicals that have a foamy quality when agitated in water. They are widely distributed ...
,
caffeic acid Caffeic acid is an organic compound that is classified as a hydroxycinnamic acid. This yellow solid consists of both phenolic and acrylic functional groups. It is found in all plants because it is an intermediate in the biosynthesis of lignin, one ...
, and a yellow
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
, ilixanthin. The berries are generally regarded as
toxic Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subst ...
to humans. Accidental consumption may occur by children or pets attracted to the bright red berries. The berries are
emetic Vomiting (also known as emesis and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteri ...
, possibly due to the compound ilicin. Caffeine and theobromine found in the plant are generally toxic to dogs and cats. The leaves of
yerba mate Yerba mate or yerba-maté (''Ilex paraguariensis''; from Spanish ; pt, erva-mate, or ; gn, ka'a, ) is a plant species of the holly genus ''Ilex'' native to South America. It was named by the French botanist Augustin Saint-Hilaire. The leave ...
, also in the genus ''Ilex'', are used to make a caffeinated beverage called mate or Paraguayan tea. In
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
, holly is supposed to be
diuretic A diuretic () is any substance that promotes diuresis, the increased production of urine. This includes forced diuresis. A diuretic tablet is sometimes colloquially called a water tablet. There are several categories of diuretics. All diuretics in ...
, a relief from
fever Fever, also referred to as pyrexia, is defined as having a body temperature, temperature above the human body temperature, normal range due to an increase in the body's temperature Human body temperature#Fever, set point. There is not a single ...
, and a
laxative Laxatives, purgatives, or aperients are substances that loosen stools and increase bowel movements. They are used to treat and prevent constipation. Laxatives vary as to how they work and the side effects they may have. Certain stimulant, lubri ...
.Wren, R.C. (1988). ''Potter's New Cyclopedia of Botanical Drugs and Preparations''. C.W. Daniel .


Other uses

Between the thirteenth and eighteenth centuries, before the introduction of
turnip The turnip or white turnip (''Brassica rapa'' subsp. ''rapa'') is a root vegetable commonly grown in temperate climates worldwide for its white, fleshy taproot. The word ''turnip'' is a compound of ''turn'' as in turned/rounded on a lathe and ' ...
s, ''Ilex aquifolium'' was cultivated for use as winter
fodder Fodder (), also called provender (), is any agriculture, agricultural foodstuff used specifically to feed domesticated livestock, such as cattle, domestic rabbit, rabbits, sheep, horses, chickens and pigs. "Fodder" refers particularly to food g ...
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 Bagpipes are a woodwind instrument using enclosed reeds fed from a constant reservoir of air in the form of a bag. The Great Highland bagpipes are well known, but people have played bagpipes for centuries throughout large parts of Europe, No ...
before tastes turned to imported dense tropical woods such as
cocuswood ''Brya ebenus'', also known as espino de sabana, granadillo, cocus wood, cocuswood, and coccuswood, is a species of flowering tree in the pea family, Fabaceae, that is native to the Caribbean islands of Cuba and Jamaica. Horticulturally it is ...
,
ebony Ebony is a dense black/brown hardwood, coming from several species in the genus ''Diospyros'', which also contains the persimmons. Unlike most woods, ebony is dense enough to sink in water. It is finely textured and has a mirror finish when pol ...
, and
African blackwood ''Dalbergia melanoxylon'' (African blackwood, grenadilla, or mpingo) is a flowering plant in the family Fabaceae, native to seasonally dry regions of Africa from Senegal east to Eritrea and south to the north-eastern parts of South Africa. The ...
.


References


External links


''Ilex aquifolium''
- information, genetic conservation units and related resources.
European Forest Genetic Resources Programme European Forest Genetic Resources Programme (EUFORGEN) is an international network that supports the conservation and sustainable use of forest genetic resources in Europe. The programme’s tasks include to coordinate and promote ''in situ ' ...
(EUFORGEN) {{Taxonbar, from=Q192190 aquifolium Trees of Africa Trees of Asia Trees of Europe Trees of mild maritime climate Medicinal plants of Africa Medicinal plants of Asia Medicinal plants of Europe Bird food plants Garden plants of Asia Garden plants of Europe Ornamental trees Plants described in 1753 Taxa named by Carl Linnaeus