Ilex (/ˈaɪlɛks/) orholly[3] is agenus of over 570 species of flowering plants in thefamilyAquifoliaceae, and the only living genus in that family.[4]Ilex has the most species of any woodydioeciousangiosperm genus.[5] The species areevergreen ordeciduous trees, shrubs, and climbers from tropics to temperate zones worldwide. Thetype species isIlex aquifolium, the common European holly used inChristmas decorations and cards.
The genus is widespread throughout thetemperate andsubtropical 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 theTertiary period and many species are adapted tolaurel forest habitats. It occurs from sea level to more than 2,000 m (6,600 ft) with high mountain species. It is a genus of small,evergreen trees with smooth, glabrous, or pubescentbranchlets. The plants are generally slow-growing with some species growing to 25 m (82 ft) tall. Thetype species is the European hollyIlex aquifolium described by Linnaeus.[2] 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 generallydioecious, with male and female flowers on different plants.[6]
The smallfruits ofIlex, although often referred to asberries, are technicallydrupes.[7] They range in color from red to brown to black, and rarely green or yellow. The "bones" contain up to tenseeds each. Some species produce fruitsparthenogenetically, 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 ofI. aquifolium, are widely used in Christmas decoration. The fruits are generally slightlytoxic to humans, and can causevomiting anddiarrhea 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, fromCalifornia toBritish Columbia, 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.[8][9][10][11]
Ilex in Latin means the holm-oak or evergreen oak (Quercus ilex). Despite theLinnaean classification ofIlex as holly, as late as the 19th century in Britain, the termIlex was still being applied to the oak as well as the holly – possibly due to the superficial similarity of the leaves.[12][13] The name "holly" in common speech refers toIlex 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 ofOld Englishhole(ġ)n,[14] Middle EnglishHolin, laterHollen.[15][16]TheFrench word for holly,houx, derives from theOld Low Franconian*hulis (Middle Dutchhuls).[17] Both are related toOld High Germanhulis,huls,[18] as areLow German/Low Franconian terms likeHülse orhulst. TheseGermanic words appear to be related to words for holly inCeltic languages, such asWelshcelyn,Bretonkelen(n) andIrishcuileann.[19]
SeveralRomance languages use the Latin wordacrifolium, literally "sharp leaf" (turned intoaquifolium in modern time), hence Italianagrifoglio,Occitangrefuèlh, etc.[20]
Hollies (here,Ilex aquifolium) are dioecious: (above) shoot with flowers from male plant; (top right) male flower enlarged from female plant; (lower right) female flower enlarged, showing ovary and reduced, sterile stamens with no pollen.
Thephylogeography of this group provides examples of variousspeciation mechanisms at work. In this scenario ancestors of this group became isolated from the remainingIlex when the Earth mass broke away intoGondwana andLaurasia 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 calledallopatric speciation. Over time, survivor species of the holly genus adapted to different ecological niches. This led toreproductive isolation, an example ofecological speciation. In thePliocene, around five million years ago,mountain formation diversified the landscape and provided new opportunities for speciation within the genus.
Thefossil record indicates that theIlex lineage was already widespread prior to the end of theCretaceous period; the earliest records of the distinctive pollen ofIlex are from theTuronian of theOtway Basin ofAustralia. The earliest fossil holly fruit is known from theMaastrichtian ofcentral Europe.[21] Based on themolecular clock, the common ancestor of most of the extant species probably appeared during theEocene, about 50million years ago, suggesting that older representatives of the genus belong to now extinct branches.[22]Ilex sinica seems to be the mostbasal extant species.[23] Thelaurel forest covered great areas of the Earth during thePaleogene, when the genus was more prosperous. This type of forest extended during theNeogene, 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 thePleistocene. 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 speciesrelict in coastal enclaves, archipelagos, and coastal mountains sufficiently far from areas of extreme cold and aridity and protected by the oceanic influence.
The genus is distributed throughout the world's different climates. Most species make their home in thetropics andsubtropics, with a worldwide distribution intemperate zones. The greatest diversity of species is found in the Americas and in Southeast Asia.
In Europe the genus is represented by a single species, the classically named hollyIlex aquifolium, and in continentalAfrica by this species andIlex mitis.Ilex canariensis, fromMacaronesia, andIlex aquifolium arose from a common ancestor in thelaurel forests of the Mediterranean. Australia, isolated at an early period, hasIlex arnhemensis. Of 204 species growing in China, 149 species areendemic. A species which stands out for its economic importance in Spanish-speaking countries and in Brazil isIlex paraguariensis orYerba mate. Havingevolved numerous species that areendemic to islands and small mountain ranges, and being highly useful plants, many hollies are now becoming rare.
Often thetropical species are especially threatened byhabitat destruction andoverexploitation. At least two species ofIlex have become extinct recently, and many others are barely surviving.[28]
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 winterstorms,birds often take refuge in hollies, which provide shelter, protection frompredators (by the spiny leaves), andfood. The flowers are sometimes eaten by thelarva of thedouble-striped pugmoth (Gymnoscelis rufifasciata). OtherLepidoptera whose larvae feed on holly includeBucculatrix ilecella, which feeds exclusively on hollies, andthe engrailed (Ectropis crepuscularia).
Holly berries can cause vomiting and diarrhea. They are especially dangerous in cases involving accidental consumption by children attracted to the bright red berries.[31] Ingestion of over 20 berries may be fatal to children.[30][31]
Holly leaves, if eaten, might cause diarrhea, nausea, vomiting, and stomach and intestinal problems.[31]
Holly plants might be toxic to pets and livestock.[32]
Leaves of some holly species are used by some cultures to make daily tea. These species areYerba 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.[33] In general little is known about inter-species variation in constituents or toxicity of hollies.
Holly berries are fermented and distilled to produce aneau de vie.[34][35]
Hollies are often used forhedges; the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping.[37] Many hundreds ofhybrids andcultivars 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).[38] The cultivarsI. ×meserveaeBlue Prince = 'Conablu' andBlue Princess = 'Conapri'[39] have gained theRoyal Horticultural Society'sAward of Garden Merit.[40][41] Another hybrid isIlex ×koehneana, with the cultivar 'Chestnut Leaf'.[42]
Traditional Christmas card with holly andmistletoe. Circa 1880s
Holly – specifically the European holly,Ilex aquifolium – is commonly referenced atChristmas time, and is often referred to by the nameChrist's thorn.[43][44] In manyWestern Christian cultures, holly is used as a traditionalChristmas decoration,[45] used especially inwreaths and illustrations, for instance onChristmas cards. Since medieval times the plant has carried aChristian symbolism,[46] as expressed in the traditional Christmas carol "The Holly and the Ivy", in which the holly represents Jesus and the ivy represents theVirgin Mary.[44] Angie Mostellar discusses the Christian use of holly at Christmas, stating that:[44]
Christians have identified a wealth of symbolism in its form. The sharpness of the leaves help to recall thecrown of thorns worn by Jesus; the red berries serve as a reminder of the drops ofblood that were shed forsalvation; and the shape of the leaves, which resemble flames, can serve to reveal God's burning love for His people. Combined with the fact that holly maintains its bright colors during theChristmas season, it naturally came to be associated with the Christian holiday.[44]
Inheraldry, holly is used to symbolize truth. The Norwegian municipality ofStord has a yellow twig of holly in its Coat-of-arms.[citation needed]
TheDruids held that "leaves of holly offered protection against evil spirits" and thus "wore holly in their hair".[44]
In some traditions ofWicca, the Holly King is one of the faces of the Sun God. He is born at midsummer and rules fromMabon toOstara.[citation needed]
In the Irish language, the wordsmac cuilinn 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.[citation needed]
Gan Mao Ling (simplified Chinese: 感冒灵; traditional Chinese: 感冒靈) is a Chinese herbal medication alleged to be effective in the early stages of cold or flu, and one main ingredient is the root of the Ilex. It’s a commonly used and widely known medication in Traditional Chinese Medicine.The ingredients of the remedy are:Ilex root, Gang Mei GenEuodia leaf, San Cha KuChrysanthemum flower, Ju HuaVitex herb, Huang Jing CaoIsatis root, Ban Lan GenLonicera flower, Jin Yin Hua
^Loizeau, P.-A.; Barriera G.; Manen J.-F.; Broennimann O. (2005)."Towards an understanding ofIlex L. (Aquifoliaceae) on a World-wide scale". In Friis I.; Balslev H. (eds.).Plant diversity and complexity patterns: local, regional, and global dimensions: proceedings of an international symposium held at the Royal Danish Academy of Sciences and Letters in Copenhagen, Denmark, 25–28 May 2003. Biologiske skrifter. Vol. 55. Kongelige Danske videnskabernes selskab. pp. 507–517.ISBN978-87-7304-304-2. Retrieved9 April 2012.
^Bailes, Christopher (December 2022). "Plant profile: Hollies".The Garden.147 (12):40–45.
^Sanford Niles, ed. (1895).Christmas History and Customs. School Education Company. p. 24.In Germany and Scandinavia the holly, or holy tree, is called "Christ's thorn," from its use in church decorations, and because it bears berries at Christmas-tide.
^Ciesla, William M. (2002).Non-wood Forest Products from Temperate Broad-leaved Trees. Food and Agriculture Organization of the United Nations. p. 13.ISBN978-92-5-104855-9.Holly is still a popular Christmas decoration among Christian cultures.
^Struthers, Jane (4 October 2010).The Book of Christmas. Ebury Publishing. p. 218.ISBN978-1-4481-4893-6.The British native holly (Ilex aquifolium) has tremendous religious significance at Christmas. Its prickly leaves are evocative of the crown of thorns that was placed on Jesus Christ's head at His crucifixion, and its scarlet berries are synonymous with drops of His blood.
^Rowling, J. K.; Rowling, J. K. (1999).Harry Potter. 1: Harry Potter and the sorcerer's stone (1. American ed., [Nachdr.]. 1999 ed.). New York: Levine Books.ISBN978-0-590-35340-3.
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