''Ilex'' (), or holly, is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

of over 570 species of flowering plants 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, and the only living genus in that family. ''Ilex'' has the most species of any woody dioecious angiosperm genus. The species are

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 ...

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 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 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 ...

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 (botany), family Aquifoliaceae, native plant, native to western and southern Europe, nort ...

'', the common European holly used in

Christmas Christmas is an annual festival commemorating Nativity of Jesus, the birth of Jesus, Jesus Christ, observed primarily on December 25 as a religious and cultural celebration among billions of people Observance of Christmas by country, around t ...

decorations and cards.

Description

Ilex paraguariensis - Köhler–s Medizinal-Pflanzen-074

The genus ''Ilex'' is divided into three subgenera: *''Ilex'' subg. ''Byronia'', with the type species ''

Ilex polypyrena ''Ilex'' (), or holly, is a genus of over 570 species of flowering plants in the family Aquifoliaceae, and the only living genus in that family. ''Ilex'' has the most species of any woody dioecious angiosperm genus. The species are evergreen o ...

'' *''Ilex'' subg. ''Prinos'', with 12 species *''Ilex'' subg. ''Ilex'', with the rest of the species The genus is widespread throughout the

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 ...

and

subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...

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 Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...

period 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 elo ...

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 ''

'' 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 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 ...

, with male and female flowers on different plants. The small

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 ...

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 or pit, although many pips or seeds may be present. Common examples are strawberries, raspb ...

, are technically

drupes 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'') ...

. They range in color from red to brown to black, and rarely green or yellow. The "bones" contain up to ten

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 each. Some species produce fruits

parthenogenetic Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...

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 and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the Human nose, nose. Vomiting can be the result of ailments like Food-poisoning, foo ...

and

diarrhea Diarrhea, also spelled diarrhoea, is the condition of having at least three loose, liquid, or watery bowel movements each day. It often lasts for a few days and can result in dehydration due to fluid loss. Signs of dehydration often begin wi ...

when ingested. However, they are an important food source for 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, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...

British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...

, 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 evergreen oak, holly oak or holm oak is a large evergreen oak native to the Mediterranean region. It is a member of the ''Ilex'' section of the genus, with acorns that mature in a single summer. Description An evergreen tr ...

''). 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 ''

'', 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 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 was brought to Great Britain by Anglo-Saxon settlement of Britain, Anglo ...

',Middle English ''Holin'', later ''Hollen''. Middle English ''Holin'', later ''Hollen''. The

French French (french: français(e), link=no) may refer to: * Something of, from, or related to France ** French language, which originated in France, and its various dialects and accents ** French people, a nation and ethnic group identified with Franc ...

word for holly, ', derives from the Old Low Franconian (Middle Dutch ). Both are related to

Old High German Old High German (OHG; german: Althochdeutsch (Ahd.)) is the earliest stage of the German language, conventionally covering the period from around 750 to 1050. There is no standardised or supra-regional form of German at this period, and Old High ...

', ''huls'', as are

Low German : : : : : (70,000) (30,000) (8,000) , familycolor = Indo-European , fam2 = Germanic , fam3 = West Germanic , fam4 = North Sea Germanic , ancestor = Old Saxon , ancestor2 = Middle L ...

/ Low Franconian terms like ' or '. These Germanic words appear to be related to words for holly in

Celtic languages The Celtic languages ( usually , but sometimes ) are a group of related languages descended from Proto-Celtic. They form a branch of the Indo-European language family. The term "Celtic" was first used to describe this language group by Edward ...

, such as

Welsh Welsh may refer to: Related to Wales * Welsh, referring or related to Wales * Welsh language, a Brittonic Celtic language spoken in Wales * Welsh people People * Welsh (surname) * Sometimes used as a synonym for the ancient Britons (Celtic peop ...

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 **Ga ...

' and Irish '. Several

Romance languages The Romance languages, sometimes referred to as Latin languages or Neo-Latin languages, are the various modern languages that evolved from Vulgar Latin. They are the only extant subgroup of the Italic languages in the Indo-European language fam ...

use the Latin word ''acrifolium'', literally "sharp leaf" (turned into ''aquifolium'' in modern time), so 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 Occitan (; o ...

', etc.

History

The phylogeography 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, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...

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 Pan ...

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 The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offspring ...

, an example of ecological speciation. In the

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million 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 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...

period. Based on the

molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleoti ...

, the common ancestor of most of the extant species probably appeared during the

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

, about 50million years ago, suggesting that older representatives of the genus belong to now extinct branches. ''

Ilex sinica ''Ilex sinica'' is a species of flowering plant in the holly family Aquifoliaceae, native to Yunnan and Guangxi Guangxi (; ; Chinese postal romanization, alternately romanized as Kwanghsi; ; za, Gvangjsih, italics=yes), officially the Gu ...

'' seems to be the most

basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...

extant species. The

covered great areas of the Earth during the

Paleogene The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...

, when the genus was more prosperous. This type of forest extended during the

Neogene The Neogene ( ), informally Upper Tertiary or Late Tertiary, 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 Mya. ...

, 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 ( , 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 ...

. 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'' *'' Ilex amelanchier'' *'' Ilex anomala'' *''

'' – European holly, English holly, Christ's thorn (western and southern Europe, northwest Africa, and southwest Asia) *''

Ilex canariensis ''Ilex canariensis'', the small-leaved holly, is an endemic species of holly native to Macaronesian islands. It is a species of plant in the family Aquifoliaceae. It is found in the Macaronesian islands of Madeira (Portugal) and Canary Island ...

'' (Macaronesian islands) *''

Ilex cassine ''Ilex cassine'' is a holly native to the southeastern coast of North America, in the United States from Virginia to southeast Texas, in Mexico in Veracruz, and in the Caribbean on the Bahamas, Cuba, and Puerto Rico. It is commonly known as dahoo ...

'' – 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 ''Ilex cornuta'', commonly known as Chinese holly or horned holly, is a slow-growing, densely foliaged evergreen shrub in the Aquifoliaceae plant family. It is native to eastern China and Korea and attains a height of about . The leaves are usu ...

, 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 (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...

) *''

Ilex gardneriana ''Ilex gardneriana'' is a critically endangered species of plant in the family Aquifoliaceae. It is endemic to the Nilgiri Hills of India. References External links Specimenat Kew Gardens Kew Gardens is a botanical garden, botanic ga ...

'' (

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

: 20th century?) (India) *'' Ilex glabra'' L. A.Gray – evergreen winterberry, bitter gallberry, inkberry (eastern North America) *''

Ilex guayusa ''Ilex guayusa'' ( or ) is a species of tree of the holly genus, native to the Amazon Rainforest. One of four known caffeinated holly trees, the leaves of the guayusa tree are harvested fresh and brewed like a tea for their stimulative effects. ...

'' (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 ''Ilex kaushue'' is a species of plant in the holly family, ''Aquifoliaceae'', which is used to make a very bitter tea called kuding Kuding or kuzding (; pronounced ) is a particularly bitter-tasting Chinese infusion, which due to their simil ...

'' (China) *'' Ilex khasiana'' (India) *'' Ilex latifolia'' – tarajo holly, ''tarayō'' (Japanese) (southern Japan and eastern and southern China ) *''

Ilex mitis ''Ilex mitis'' (commonly called Cape holly, African holly, waterboom or umDuma) is a tall, dense, evergreen tree that is indigenous to Sub-Saharan Africa and Madagascar. It makes an excellent fast-growing hedge for gardens - growing tall, stra ...

'' (southern Africa) *'' Ilex montana'' Torrey &

A.Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His '' Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually exc ...

– mountain winterberry ( Eastern United States) *''

Ilex mucronata ''Ilex mucronata'', the mountain holly or catberry, is a species of holly native to eastern North America, from Newfoundland west to Minnesota, and south to Maryland and West Virginia. Taxonomy It was formerly treated in its own monotypic genus ...

'' (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 ''Ilex pedunculosa'', called longstalk holly, is a species of flowering plant in 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 g ...

'' – longstalked holly *''

Ilex perado ''Ilex perado'', the Macaronesian holly, is a species of holly endemic to Macaronesia, distributed throughout the Azores, Madeira and Canary islands. It is an important component of the natural high-altitude Macaronesian rainforest, known as 'lau ...

'' – Macaronesian holly *'' Ilex quercetorum'' (Mexico and Guatemala) *'' Ilex rotunda'' *'' Ilex serrata'' – Japanese winterberry *''

Ilex verticillata ''Ilex verticillata'', the winterberry, is a species of holly native to eastern North America in the United States and southeast Canada, from Newfoundland west to Ontario and Minnesota, and south to Alabama. Other names that have been used incl ...

'' ( L.)

American winterberry (Eastern North America) *''

Ilex vomitoria ''Ilex vomitoria'', commonly known as yaupon () or yaupon holly, is a species of holly that is native to southeastern North America. The word ''yaupon'' was derived from the Catawban ''yą́pą'', from ''yą-'' tree + ''pą'' leaf. Another comm ...

'' – 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. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...

and subtropics, with a worldwide distribution in

temperate zone 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 ...

s. The greatest diversity of species is found in the Americas and in Southeast Asia. ''

'', formerly the type species of '' Nemopanthus'', is

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 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 ''

'', and in continental

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

by this species and ''

''. ''

'', 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 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 found elsew ...

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

evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...

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. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...

species are especially threatened by

habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...

and

overexploitation Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term app ...

. 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 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 Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...

s often take refuge in hollies, which provide shelter, protection from

predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...

s (by the spiny leaves), and

food Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is inge ...

. The flowers are sometimes eaten by the

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

of the

double-striped pug The double-striped pug (''Gymnoscelis rufifasciata'') is a moth of the family Geometridae. It is a widespread and common species, being found throughout the Palearctic region, including the Near East and North Africa. This is a variable species ...

moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...

(''Gymnoscelis rufifasciata''). Other

Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...

whose larvae feed on holly include ''

Bucculatrix ilecella ''Bucculatrix ilecella'' is a moth in the family Bucculatricidae. It is found in North America, where it has been recorded from Texas. It was first described by August Busck in 1915. The wingspan The wingspan (or just span) of a bird or an a ...

'', which feeds exclusively on hollies, and

the engrailed The engrailed and small engrailed (''Ectropis crepuscularia'') are moths of the family Geometridae found from the British Isles through central and eastern Europe to the Russian Far East and Kazakhstan. The western Mediterranean and Asia Minor ...

(''Ectropis crepuscularia'').

Toxicity

Holly can contain caffeic acid,

caffeoyl 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, on ...

derivatives, caffeoylshikimic acid, chlorogenic acid, feruloylquinic acid, quercetin,

quinic acid Quinic acid is a cyclitol, a cyclic polyol, and a cyclohexanecarboxylic acid. It is a colorless solid that can be extracted from plant sources. Quinic acid is implicated in the perceived acidity of coffee. Occurrence and preparation The compound ...

, kaempferol,

tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'', ...

s, rutin,

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 ...

, and

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 ...

. 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''; 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 ...

(''I. paraguariensis''), ''

'', Kuding (''

''), 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 plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...

s 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 type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...

s have been developed for garden use, among them the very popular "

Highclere Highclere (pronounced ) is a village and civil parish situated in the North Wessex Downs (an Area of Outstanding Natural Beauty) in the Basingstoke and Deane district of Hampshire, England. It lies in the northern part of the county, near the B ...

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 (Nort ...

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 ...

. Another hybrid is ''Ilex'' × ''koehneana'', with the cultivar 'Chestnut Leaf'.

Culture

Holly – more 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 a traditional Christmas decoration, used especially in wreaths and illustrations, for instance on

Christmas card A Christmas card is a greeting card sent as part of the traditional celebration of Christmas in order to convey between people a range of sentiments related to Christmastide and the holiday season. Christmas cards are usually exchanged during t ...

s. Since medieval times the plant has carried a

Christian symbolism Christian symbolism is the use of symbols, including archetypes, acts, artwork or events, by Christianity. It invests objects or actions with an inner meaning expressing Christian ideas. The symbolism of the early Church was characterized by bei ...

, 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; arc, ܡܪܝܡ, translit=Mariam; ar, مريم, translit=Maryam; grc, Μαρία, translit=María; la, Maria; cop, Ⲙⲁⲣⲓⲁ, translit=Maria was a first-century Jewish woman of Nazareth, the wife of Joseph and the mother o ...

. 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, rank and pedigree. Armory, the best-known branch ...

, 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 ''Harry Potter'' is a series of seven fantasy literature, fantasy novels written by British author J. K. Rowling. The novels chronicle the lives of a young Magician (fantasy), wizard, Harry Potter (character), Harry Potter, and his friends ...

'' novels, holly is used as the wood in Harry's wand. In some traditions of

Wicca Wicca () is a modern Pagan religion. Scholars of religion categorise it as both a new religious movement and as part of the occultist stream of Western esotericism. It was developed in England during the first half of the 20th century and was ...

, the Holly King is one of the faces of the Sun God. He is born at midsummer and rules from Mabon to Ostara.

Gallery

Image:Starr_020925-0058_Ilex_anomala.jpg, '' Ilex anomala'' Image:Contorted Hedgehog Holly.JPG, A contorted hedgehog holly ''

'' 'Ferox' Image:Ilex chinensis1.jpg, '' Ilex chinensis'' Image:Ilex pedunculosa5.jpg, ''

'' 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
i

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