''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

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 ...

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'', 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 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 habitats. It occurs from sea level to more than with high mountain species. It is a genus of small,

trees with smooth, glabrous, or pubescent branchlets. The plants are generally slow-growing with some species growing to tall. The

is the European holly '' Ilex aquifolium'' 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, 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. 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 to humans, and can cause vomiting and diarrhea 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 ...

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''). 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'', 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 word for holly, ', derives from the Old Low Franconian (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 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 ', Breton ' 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 ', etc.

History

The phylogeography of this group provides examples of various speciation mechanisms at work. In this scenario ancestors of this group became isolated from the remaining ''Ilex'' when the Earth mass broke away into Gondwana and Laurasia 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 Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...

. 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 The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58mountain 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 The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

period; the earliest records of the distinctive pollen of ''Ilex'' are from the

Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...

of the Otway Basin of

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

. The earliest fossil holly fruit is known from the

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

central Europe Central Europe is a geographical region of Europe between Eastern Europe, Eastern, Southern Europe, Southern, Western Europe, Western and Northern Europe, Northern Europe. Central Europe is known for its cultural diversity; however, countries in ...

. Based on the molecular clock, the common ancestor of most of the extant species probably appeared during the

Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...

, about 50million years ago, suggesting that older representatives of the genus belong to now extinct branches. '' Ilex sinica'' seems to be the most basal extant species. The laurel forest covered great areas of the Earth during the Paleogene, when the genus was more prosperous. This type of forest extended during the

Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...

, 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 The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...

. 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 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'' – 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) (eastern China, Japan, Korea, Taiwan, and Sakhalin) *'' Ilex decidua'' Walter – possumhaw (eastern United States, northeastern

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

) *'' Ilex gardneriana'' (

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

: 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'' Torrey & 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'' ( L.) 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 The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...

and subtropics, with a worldwide distribution in

temperate zone 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 ra ...

s. The greatest diversity of species is found in the Americas and in Southeast Asia. '' Ilex mucronata'', formerly the type species of '' Nemopanthus'', is 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., Savolainen, V., 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'', and in continental

Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

by this species and '' Ilex mitis''. '' Ilex canariensis'', from Macaronesia, and ''Ilex aquifolium'' arose from a common ancestor in the laurel forests of the Mediterranean. Australia, isolated at an early period, has '' Ilex arnhemensis''. Of 204 species growing in China, 149 species are

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

. A species which stands out for its economic importance in Spanish-speaking countries and in Brazil is '' Ilex paraguariensis'' or ''Yerba mate''. Having evolved numerous species that are

to islands and small mountain ranges, and being highly useful plants, many hollies are now becoming rare.

Ecology

Often the

tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...

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,

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

s often take refuge in hollies, which provide shelter, protection from predators (by the spiny leaves), and

food Food is any substance consumed by an organism for Nutrient, nutritional support. Food is usually of plant, animal, or Fungus, fungal origin and contains essential nutrients such as carbohydrates, fats, protein (nutrient), proteins, vitamins, ...

. The flowers are sometimes eaten by the larva of the double-striped pug

moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...

(''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 Caffeine is a central nervous system (CNS) stimulant of the methylxanthine chemical classification, class and is the most commonly consumed Psychoactive drug, psychoactive substance globally. It is mainly used for its eugeroic (wakefulness pr ...

, 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 Yerba mate or yerba maté (), ''Ilex paraguariensis'', is a plant species of the holly genus native to South America. It was named by the French botanist Augustin Saint-Hilaire. The leaves of the plant can be steeped in hot water to make a bev ...

(''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: *'' I. aquifolium'' (common European holly) *'' I. crenata'' (box-leaved holly) *'' I. verticillata'' (winterberry) Hollies are often used for hedges; the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping. Many hundreds of hybrids and

cultivar A cultivar is a kind of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and which retains those traits when Plant propagation, propagated. Methods used to propagate cultivars include division, root a ...

s have been developed for garden use, among them the very popular " Highclere holly", ''Ilex'' × ''altaclerensis'' (''I. aquifolium'' × ''I. perado'') and the "blue holly", ''Ilex'' × ''meserveae'' (''I. aquifolium'' × '' 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 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 ...

'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

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 Mary was a first-century Jewish woman of Nazareth, the wife of Saint Joseph, Joseph and the mother of Jesus. She is an important figure of Christianity, venerated under titles of Mary, mother of Jesus, various titles such as Perpetual virginity ...

. Angie Mostellar discusses the Christian use of holly at Christmas, stating that: In

heraldry Heraldry is a discipline relating to the design, display and study of armorial bearings (known as armory), as well as related disciplines, such as vexillology, together with the study of ceremony, Imperial, royal and noble ranks, rank and genealo ...

, 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's wand. In some traditions of

Wicca Wicca (), also known as "The Craft", is a Modern paganism, modern pagan, syncretic, Earth religion, Earth-centred religion. Considered a new religious movement by Religious studies, scholars of religion, the path evolved from Western esote ...

, the Holly King is one of the faces of the Sun God. He is born at midsummer and rules from 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'' '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'' Image:Ilex-pernyi-leaves.JPG, '' Ilex pernyi'' from west China

References

External links

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Aquifoliaceae
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