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''Ilex'' () or holly is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of over 570 species of flowering plants in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Aquifoliaceae, and the only living genus in that family. ''Ilex'' has the most species of any woody
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
genus. The species are
evergreen In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
or
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
trees, shrubs, and climbers from tropics to temperate zones worldwide. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
is ''
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 As ...
'', the common European holly used in
Christmas Christmas is an annual festival commemorating Nativity of Jesus, the birth of Jesus Christ, observed primarily on December 25 as a Religion, religious and Culture, cultural celebration among billions of people Observance of Christmas by coun ...
decorations and cards.


Description

The genus is widespread throughout the
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
regions of the world. It includes species of trees, shrubs, and climbers, with evergreen or deciduous foliage and inconspicuous flowers. Its range was more extended in the
Tertiary period The Tertiary ( ) is an obsolete Period (geology), geologic period spanning 66 million to 2.6 or 1.8 million years ago. The period began with the extinction of the non-bird, avian dinosaurs in the Cretaceous–Paleogene extinction event, at t ...
and many species are adapted to
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 el ...
habitats. It occurs from sea level to more than with high mountain species. It is a genus of small,
evergreen In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
trees with smooth, glabrous, or pubescent
branchlet A branch, also called a ramus in botany, is a stem that grows off from another stem, or when structures like veins in leaves are divided into smaller veins. History and etymology In Old English, there are numerous words for branch, includ ...
s. The plants are generally slow-growing with some species growing to tall. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
is the European holly ''
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 As ...
'' described by Linnaeus. Plants in this genus have simple, alternate glossy leaves, frequently with a spiny leaf margin. The inconspicuous flower is greenish white, with four petals. They are generally
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
, with male and female flowers on different plants. The small
fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
s of ''Ilex'', although often referred to as
berries A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone fruit, stone or pit (fruit), pit although many wikt:pip#Etymology 2, pips or seeds may be p ...
, are technically
drupes In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part ( exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or '' pyrena'') of hardened endocarp with a s ...
. They range in color from red to brown to black, and rarely green or yellow. The "bones" contain up to ten
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s each. Some species produce fruits
parthenogenetic Parthenogenesis (; from the Greek + ) is a natural form of asexual reproduction in which the embryo develops directly from an egg without need for fertilization. In animals, parthenogenesis means the development of an embryo from an unfertiliz ...
ally, such as the cultivar 'Nellie R. Stevens'. The fruits ripen in winter and thus provide winter colour contrast between the bright red of the fruits and the glossy green evergreen leaves. Hence the cut branches, especially of ''I. aquifolium'', are widely used in Christmas decoration. The fruits are generally slightly
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, and can cause
vomiting Vomiting (also known as emesis, puking and throwing up) is the 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, gastroenteritis, pre ...
and
diarrhea Diarrhea (American English), also spelled diarrhoea or diarrhœa (British English), is the condition of having at least three loose, liquid, or watery bowel movements in a day. It often lasts for a few days and can result in dehydration d ...
when ingested. However, they are a food source for certain birds and other animals, which help disperse the seeds. Unfortunately this can have negative impacts as well. Along the west coast of North America, from
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
to
British Columbia British Columbia is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...
, English holly (''Ilex aquifolium''), which is grown commercially, is 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.


Etymology

''Ilex'' in Latin means the holm-oak or evergreen oak (''
Quercus ilex ''Quercus ilex'', the holly oak, also (ambiguously, as many oaks are evergreen) evergreen oak, is a large evergreen oak native to the Mediterranean region. It is a member of the section (botany), section ''List of Quercus species#Section Ilex, Il ...
''). Despite the Linnaean classification of ''Ilex'' as holly, as late as the 19th century in Britain, the term ''Ilex'' was still being applied to the oak as well as the holly – possibly due to the superficial similarity of the leaves. The name "holly" in common speech refers to ''
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 As ...
'', specifically stems with berries used in Christmas decoration. By extension, "holly" is also applied to the whole genus. The origin of the word "holly" is considered a reduced form of
Old English Old English ( or , or ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the Early Middle Ages. It developed from the languages brought to Great Britain by Anglo-S ...
',Middle English ''Holin'', later ''Hollen''. Middle English ''Holin'', later ''Hollen''. The
French French may refer to: * Something of, from, or related to France ** French language, which originated in France ** French people, a nation and ethnic group ** French cuisine, cooking traditions and practices Arts and media * The French (band), ...
word for holly, ', derives from the
Old Low Franconian In linguistics, Old Dutch ( Modern Dutch: ') or Old Low Franconian (Modern Dutch: ') is the set of dialects that evolved from Frankish spoken in the Low Countries during the Early Middle Ages, from around the 6th Page 55: "''Uit de zesde eeuw ...
(Middle Dutch ). Both are related to
Old High German Old High German (OHG; ) is the earliest stage of the German language, conventionally identified as the period from around 500/750 to 1050. Rather than representing a single supra-regional form of German, Old High German encompasses the numerous ...
', ''huls'', as are
Low German Low German is a West Germanic languages, West Germanic language variety, language spoken mainly in Northern Germany and the northeastern Netherlands. The dialect of Plautdietsch is also spoken in the Russian Mennonite diaspora worldwide. "Low" ...
/
Low Franconian In historical linguistics, historical and comparative linguistics, Low Franconian is a linguistic category used to classify a number of historical and contemporary West Germanic languages, West Germanic Variety (linguistics), varieties closely r ...
terms like ' or '. These Germanic words appear to be related to words for holly in
Celtic languages The Celtic languages ( ) are a branch of the Indo-European language family, descended from the hypothetical Proto-Celtic language. The term "Celtic" was first used to describe this language group by Edward Lhuyd in 1707, following Paul-Yve ...
, such as
Welsh Welsh may refer to: Related to Wales * Welsh, of or about Wales * Welsh language, spoken in Wales * Welsh people, an ethnic group native to Wales Places * Welsh, Arkansas, U.S. * Welsh, Louisiana, U.S. * Welsh, Ohio, U.S. * Welsh Basin, during t ...
',
Breton Breton most often refers to: *anything associated with Brittany, and generally **Breton people **Breton language, a Southwestern Brittonic Celtic language of the Indo-European language family, spoken in Brittany ** Breton (horse), a breed **Gale ...
' and Irish '. Several
Romance languages The Romance languages, also known as the Latin or Neo-Latin languages, are the languages that are Language family, directly descended from Vulgar Latin. They are the only extant subgroup of the Italic languages, Italic branch of the Indo-E ...
use the Latin word ''acrifolium'', literally "sharp leaf" (turned into ''aquifolium'' in modern time), hence Italian ',
Occitan Occitan may refer to: * Something of, from, or related to the Occitania territory in parts of France, Italy, Monaco and Spain. * Something of, from, or related to the Occitania administrative region of France. * Occitan language, spoken in parts o ...
', etc.


History

The
phylogeography Phylogeography is the study of the historical processes that may be responsible for the past to present geographic distributions of genealogical lineages. This is accomplished by considering the geographic distribution of individuals in light of ge ...
of this group provides examples of various
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
mechanisms at work. In this scenario ancestors of this group became isolated from the remaining ''Ilex'' when the Earth mass broke away into
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
and
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
about 82million years ago, resulting in a physical separation of the groups and beginning a process of change to adapt to new conditions. This mechanism is called allopatric speciation. Over time, survivor species of the holly genus adapted to different ecological niches. This led to reproductive isolation, an example of ecological speciation. In the Pliocene, around five million years ago, orogeny, mountain formation diversified the landscape and provided new opportunities for speciation within the genus. The fossil record indicates that the ''Ilex'' lineage was already widespread prior to the end of the Cretaceous period; the earliest records of the distinctive pollen of ''Ilex'' are from the Turonian of the Otway Basin of Australia. The earliest fossil holly fruit is known from the Maastrichtian of central Europe. Based on the molecular clock, the common ancestor of most of the extant species probably appeared during the Eocene, about 50million years ago, suggesting that older representatives of the genus belong to now extinct branches. ''Ilex sinica'' seems to be the most Basal (phylogenetics), basal extant species. The
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 el ...
covered great areas of the Earth during the Paleogene, when the genus was more prosperous. This type of forest extended during the Neogene, more than 20million years ago. Most of the last remaining temperate broadleaf evergreen forests are believed to have disappeared about 10,000 years ago at the end of the Pleistocene. Many of the then-existing species with the strictest ecological requirements became extinct because they could not cross the barriers imposed by the geography, but others found refuge as a species Relict (biology), relict in coastal enclaves, archipelagos, and coastal mountains sufficiently far from areas of extreme cold and aridity and protected by the oceanic influence.


Selected species

*''Ilex ambigua'' - Southeastern USA *''Ilex amelanchier'' - Southeastern USA *''Ilex anomala'' (Hawaii) *''
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 As ...
'' – European holly, English holly, Christ's thorn (western and southern Europe, northwest Africa, and southwest Asia) *''Ilex canariensis'' (Macaronesian islands) *''Ilex cassine'' – dahoon holly, cassena (Virginia to southeast Texas of US, Veracruz of Mexico, Bahamas, Cuba, and Puerto Rico) *''Ilex coriacea'' – gallberry (Virginia to Texas of United States) *''Ilex cornuta'' – Chinese holly, horned holly (eastern China and Korea) *''Ilex crenata'' – Japanese holly, box-leaved holly, ''inutsuge'' (Japanese (language), Japanese) (eastern China, Japan, Korea, Taiwan, and Sakhalin) *''Ilex decidua'' Walter – possumhaw (eastern United States, northeastern Mexico) *''Ilex gardneriana'' (extinct: 20th century?) (India) *''Ilex glabra'' L. A.Gray – evergreen winterberry, bitter gallberry, inkberry (eastern North America) *''Ilex guayusa'' (Amazon rainforest) *''Ilex integra'' – mochi tree, Nepal holly (Korea; Taiwan; the mid-southern regions of China; and Honshu, Shikoku and Kyushu in Japan) *''Ilex kaushue'' (China) *''Ilex khasiana'' (India) *''Ilex latifolia'' – tarajo holly, ''tarayō'' (Japanese) (southern Japan and eastern and southern China ) *''Ilex mitis'' (southern Africa) *''Ilex montana'' John Torrey, Torrey & Asa Gray, A.Gray – mountain winterberry (Eastern United States) *''Ilex mucronata'' (L.) M.Powell, Savol., & S.Andrews – mountain holly, catberry (Eastern North America) *''Ilex opaca'' – American holly (Eastern United States) *''Ilex paraguariensis'' – yerba mate (mate, erva-mate) *''Ilex pedunculosa'' – longstalked holly *''Ilex perado'' – Macaronesian holly *''Ilex quercetorum'' (Mexico and Guatemala) *''Ilex rotunda'' — Kurogane holly (east Asia) *''Ilex rugosa'' – Tsuru Holly (mountains of Japan, Sakhalin, Khabarovsk Krai and Kuril Islands Siberia) *''Ilex serrata'' – Japanese winterberry *''Ilex verticillata'' (Carl Linnaeus, L.) Asa Gray, A.Gray American winterberry (Eastern North America) *''Ilex vomitoria'' – yaupon holly (southeastern United States)


Range

The genus is distributed throughout the world's different climates. Most species make their home in the tropics and subtropics, with a worldwide distribution in temperate zones. The greatest diversity of species is found in the Americas and in Southeast Asia. ''Ilex mucronata'', formerly the type species of ''Nemopanthus'', is native plant, native to eastern North America. ''Nemopanthus'' was treated as a separate genus with eight species. of the family Aquifoliaceae, now transferred to ''Ilex'' on molecular data;Powell, M., Vincent Savolainen, Savolainen, V., Philippe Cuénoud, Cuénoud, P., Manen, J. F., & Andrews, S. (2000). The mountain holly (''Nemopanthus mucronatus'': Aquifoliaceae) revisited with molecular data. ''Kew Bulletin'' 55: 341–347. it is closely related to ''Ilex amelanchier''. In Europe the genus is represented by a single species, the classically named holly ''
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 As ...
'', and in continental Africa by this species and ''Ilex mitis''. ''Ilex canariensis'', from Macaronesia, and ''Ilex aquifolium'' arose from a common ancestor in the laurisilva, laurel forests of the Mediterranean. Australia, isolated at an early period, has ''Ilex arnhemensis''. Of 204 species growing in China, 149 species are endemism, endemic. A species which stands out for its economic importance in Spanish-speaking countries and in Brazil is ''Ilex paraguariensis'' or ''Yerba mate''. Having evolution, evolved numerous species that are endemism, endemic to islands and small mountain ranges, and being highly useful plants, many hollies are now becoming rare.


Ecology

Often the tropical species are especially threatened by habitat destruction and overexploitation. At least two species of ''Ilex'' have become extinct recently, and many others are barely surviving.International Union for Conservation of Nature and Natural Resources (IUCN) (2007): 2007 ''IUCN Red List of Threatened Species'': ''Ilex''] They are an extremely important food for numerous species of birds, and also are eaten by other wild animals. In the autumn and early winter the fruits are hard and apparently unpalatable. After being frozen or frosted several times, the fruits soften, and become milder in taste. During winter storms, birds often take refuge in hollies, which provide shelter, protection from predators (by the spiny leaves), and food. The flowers are sometimes eaten by the larva of the double-striped pug moth (''Gymnoscelis rufifasciata''). Other Lepidoptera whose larvae feed on holly include ''Bucculatrix ilecella'', which feeds exclusively on hollies, and the engrailed (''Ectropis crepuscularia'').


Toxicity

Holly can contain caffeic acid, caffeoyl derivatives, chlorogenic acid, quercetin, quinic acid, kaempferol, tannins, rutin, caffeine, theobromine, and ilicin. Holly berries can cause vomiting and diarrhea. They are especially dangerous in cases involving accidental consumption by children attracted to the bright red berries. Ingestion of over 20 berries may be fatal to children. Holly leaves, if eaten, might cause diarrhea, nausea, vomiting, and stomach and intestinal problems. Holly plants might be toxic to pets and livestock.


Uses


Culinary use

Leaves of some holly species are used by some cultures to make daily tea. These species are Yerba mate (''I. paraguariensis''), ''Ilex guayusa'', Kuding (''Ilex kaushue''), Yaupon (''I. vomitoria'') and others. Leaves of other species, such as gallberry (''I. glabra'') are bitter and emetic. In general little is known about inter-species variation in constituents or toxicity of hollies. Holly berries are fermented and distilled to produce an ''eau de vie''.


Ornamental use

Many of the holly species are widely used as ornamental plants in temperate/European gardens and parks, notably: *''Ilex aquifolium, I. aquifolium'' (common European holly) *''Ilex crenata, I. crenata'' (box-leaved holly) *''Ilex verticillata, I. verticillata'' (winterberry) Hollies are often used for hedge (barrier), hedges; the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping. Many hundreds of Hybrid (biology), hybrids and cultivars have been developed for garden use, among them the very popular "Highclere Castle, Highclere holly", ''Ilex'' × ''altaclerensis'' (''I. aquifolium'' × ''I. perado'') and the "blue holly", ''Ilex'' × ''meserveae'' (''I. aquifolium'' × ''Ilex rugosa, I. rugosa'').Huxley, A., ed. (1992). ''New RHS Dictionary of Gardening''. Macmillan . The cultivars ''I.'' × ''meserveae'' = 'Conablu' and = 'Conapri' have gained the Royal Horticultural Society's Award of Garden Merit. Another hybrid is ''Ilex'' × ''koehneana'', with the cultivar 'Chestnut Leaf'.


Culture

Holly – specifically the European holly, ''Ilex aquifolium'' – is commonly referenced at
Christmas Christmas is an annual festival commemorating Nativity of Jesus, the birth of Jesus Christ, observed primarily on December 25 as a Religion, religious and Culture, cultural celebration among billions of people Observance of Christmas by coun ...
time, and is often referred to by the name ''Christ's thorn''. In many Western Christian cultures, holly is used as a traditional Christmas decoration, used especially in wreaths and illustrations, for instance on Christmas cards. Since medieval times the plant has carried a Christian symbolism, as expressed in the traditional Christmas carol "The Holly and the Ivy", in which the holly represents Jesus and the ivy represents the Virgin Mary. Angie Mostellar discusses the Christian use of holly at Christmas, stating that: In heraldry, holly is used to symbolize truth. The Norwegian municipality of Stord has a yellow twig of holly in its Coat-of-arms. The Druids held that "leaves of holly offered protection against evil spirits" and thus "wore holly in their hair". In the ''Harry Potter'' novels, holly is used as the wood in Harry Potter (character), Harry's List of wands in Harry Potter, wand. In some traditions of Wicca, the Holly King is one of the faces of the Sun God. He is born at midsummer and rules from Wheel of the Year#Autumn Equinox (Mabon), Mabon to Ostara. In the Irish language, the words mean 'son of holly'. Common anglicized forms of this arose; last names such as McCullen, McCullion, McQuillan, and MacCullion, which are quite common surnames in some areas.


Gallery

Image:Starr_020925-0058_Ilex_anomala.jpg, ''Ilex anomala'' Image:Contorted Hedgehog Holly.JPG, A contorted hedgehog holly ''
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 As ...
'' 'Ferox' Image:Ilex chinensis1.jpg, ''Ilex chinensis'' Image:Ilex pedunculosa5.jpg, ''Ilex pedunculosa'' Image:Ilex rotunda5.jpg, ''Ilex rotunda'' Image:Yunnan Holly Ilex yunnanensis Sprig 3008px.jpg, ''Ilex yunnanensis'' Image:Beautifulflowers6.jpg, Ilex × meserveae, ''Ilex'' × ''meserveae'' Image:Ilex-pernyi-leaves.JPG, ''Ilex pernyi'' from west China


References


External links

*
Aquifoliaceae
i

{{Authority control Ilex, Medicinal plants Winter traditions Christmas plants Dioecious plants