About 600, see text
ILEX /ˈaɪlɛks/ , or HOLLY, is a genus of 400 to 600 species of
flowering plants in the family AQUIFOLIACEAE, and the only living
genus in that family. The species are evergreen or deciduous trees,
shrubs, and climbers from tropics to temperate zones worldwide.
* 1 Description
* 2 Etymology
* 3 History
* 4 Range
* 5 Ecology
* 6 Toxicity
* 7 Usage
* 8 Ornamental use
* 9 Culture
* 10 Selected species
* 11 Gallery
* 12 References
* 13 External links
The genus Ilex is widespread throughout the temperate 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 and many
species are adapted to laurel forest habitat. It occurs from sea level
to more than 2,000 metres (6,600 ft) with high mountain species. It is
a genus of small, evergreen trees with smooth, glabrous, or pubescent
branchlets. The plants are generally slow-growing with some species
growing to 25 m (82 ft) tall. The type species 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 fruits of Ilex, although often referred to as berries , are
technically drupes . They range in color from red to brown to black,
and rarely green or yellow. The "bones" contain up to ten seeds each.
Some species produce fruits parthenogenetically , 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 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 to British Columbia, English
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
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
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 hole(ġ)n, Middle English Holin, later Hollen. The French
word for holly, houx, derives from the
Old Low Franconian
Old Low Franconian *hulis
(Middle Dutch huls). Both are related to
Old High German hulis, huls,
Low German /
Low Franconian terms like Hülse or hulst. These
Germanic words appear to be related to words for holly in Celtic
languages , such as Welsh celyn, Breton kelen(n) and Irish cuileann.
Several romance languages use the Latin word acrifolium (turned into
aquifolium in modern time), so Italian agrifoglio,
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 stamen and
reduced, sterile stamens with no pollen.
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
Laurasia about 82 million 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
around five million years ago, mountain formation diversified the
landscape and provided new opportunities for speciation within the
The fossil record indicates that the Ilex lineage was already
widespread prior to the end of the
Cretaceous period. Based on the
molecular clock the common ancestor of most of the extant species
probably appeared during the
Eocene , about 50 million years ago,
suggesting that older representatives of the genus belong to now
extinct branches. The laurel forest covered great areas of the Earth
Paleogene , when the genus was more prosperous. This type
of forest extended during the
Neogene , more than 20 million years
ago. Most of the last remaining temperate 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 in coastal enclaves, archipelagos, and coastal
mountains sufficiently far from the extreme cold and aridity and
protected by the oceanic influence.
The genus includes about 400 to 600 species, divided into three
* Ilex subg. Byronia, with the type species Ilex polypyrena
* Ilex subg. Prinos, with 12 species
* Ilex subg. Ilex, with the rest of the over 400 species
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
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; 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 by
this species and (
Ilex mitis ).
Ilex canariensis , from
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 . A species which stands out for its economic importance in
Spanish-speaking countries is
Ilex paraguariensis or Yerba mate.
Having evolved numerous species that are endemic to islands and small
mountain ranges, and being highly useful plants, many hollies are now
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.
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 , 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).
Holly can contain caffeic acid , caffeoyl derivatives,
caffeoylshikimic acid , chlorogenic acid , feruloylquinic acid ,
quercetin , quinic acid , kaempferol , tannins , rutin , caffeine ,
and theobromine .
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.
Leaves of some holly species are used by some cultures to make daily
tea. These species are
Yerba mate (I. paraguariensis),
Ilex guayusa ,
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.
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)
Moreover, many hundreds of hybrids and cultivars 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). Hollies are often used for
hedges ; the spiny leaves make them difficult to penetrate, and they
take well to pruning and shaping.
Christmas card with holly and mistletoe . Circa
Holly – more specifically the European holly,
Ilex aquifolium –
is commonly referenced at
Christmas 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 cards . Since medieval
times the plant has carried a
Christian symbolism , as expressed in
the well-known Christian
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:
Christians have identified a wealth of symbolism in its form. The
sharpness of the leaves help to recall the crown of thorns worn by
Jesus; the red berries serve as a reminder of the drops of blood that
were shed for salvation ; 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
Christmas season , it naturally came to be associated with the
In heraldry , holly is used to symbolize truth. The Norwegian
Stord has a yellow twig of holly in its Coat-of-arms.
Druids held that "leaves of holly offered protection against evil
spirits" and thus "wore holly in their hair".
Harry Potter novels, holly is used as the wood in Harry 's
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 Mabon to Ostara.
* Ilex aculeolata
* Ilex affinis
* Ilex aggregata
* Ilex × altaclarensis
* Ilex amazonensis –
Ilex ambigua – Sand holly
* Ilex amboroica
* Ilex angulata
* Ilex anodonta
Arn. – Kāwaʻu (Hawaiʻi )
* Ilex apicidens
Ilex aquifolium – European holly, English holly, Christ's thorn
* Ilex archeri
* Ilex ardisiifrons
* Ilex argentina
* Ilex arimensis
* Ilex atabapoensis
* Ilex atrata
* Ilex auriculata
* Ilex austrosinensis
* Ilex belizensis
* Ilex berteroi
* Ilex bidens
* Ilex bioritsensis
* Ilex blanchetii
* Ilex boliviana
* Ilex brandegeeana
* Ilex brevicuspis
* Ilex buergeri
* Ilex buxoides
* Ilex casiquiarensis
Ilex cassine – Dahoon holly, cassena
* Ilex cauliflora
* Ilex centrochinensis
* Ilex chamaebuxus
* Ilex chamaedryfolia
* Ilex championii
* Ilex chartaceifolia
* Ilex chengbuensis
* Ilex cheniana
* Ilex chinensis
* Ilex chingiana
* Ilex chiriquensis
* Ilex ciliospinosa
* Ilex cinerea
* Ilex clementis
* Ilex cochinchinensis
* Ilex colombiana
* Ilex condensata
* Ilex confertiflora
* Ilex corallina
Ilex coriacea – Gallberry
Ilex cornuta –
Chinese holly , horned holly
Ilex crenata – Japanese holly, box-leaved holly, inutsuge
* Ilex cubana
* Ilex culmenicola
* Ilex cupreonitens
* Ilex curtissii
* Ilex cuzcoana
* Ilex cyrtura
* Ilex danielis
* Ilex daphnogenea
* Ilex dasyclada
* Ilex dasyphylla
Ilex decidua Walter – Possumhaw (Eastern United States,
* Ilex dehongensis
* Ilex delavayi
* Ilex denticulata
* Ilex dianguiensis
* Ilex dicarpa
* Ilex dioica
* Ilex dipyrena – Himalayan holly
* Ilex dicolor
* Ilex diuretica
* Ilex divaricata
* Ilex dolichopoda
* Ilex dubia
* Ilex dugesii
* Ilex duidae
* Ilex dumosa
* Ilex dunniana
* Ilex editicostata
* Ilex elliptica
* Ilex elmerrilliana
* Ilex embelioides
* Ilex eoa
* Ilex estriata
* Ilex excelsa
* Ilex fargesii
* Ilex ficifolia
* Ilex ficoidea
* Ilex formosana
* Ilex forrestii
* Ilex fragilis
* Ilex franchetiana
* Ilex fukeinensis
* Ilex gabinetensis
* Ilex gabrielleana
Ilex gardneriana (extinct : 20th century?)
* Ilex georgei
* Ilex gintungensis
Ilex glabra L. A.Gray –
Evergreen winterberry, bitter gallberry,
inkberry (Eastern North America)
* Ilex glaucophylla
* Ilex godajam
* Ilex goshiensis
* Ilex gracilis
* Ilex gransabanensis
* Ilex guangnanensis
Ilex guayusa – Guayusa
* Ilex guianensis
* Ilex guizhouensis
* Ilex gundlachiana
* Ilex haberi
* Ilex hainanensis
* Ilex hanceana
* Ilex hayatana
* Ilex hippocrateoides
* Ilex hirsuta
* Ilex hookeri
* Ilex huana
* Ilex hylonoma
* Ilex hypaneura
* Ilex hyreana
Ilex integra – Mochi tree, Nepal holly
* Ilex intricata
Ilex jamaicana Proctor (
* Ilex kingiana
* Ilex kusanoi
Ilex laevigata – Smooth winterberry
Ilex latifolia – Tarajo holly, tarayō (Japanese)
* Ilex leucoclada
* Ilex longipes
* Ilex macoucoua
* Ilex macrocarpa
* Ilex macropoda
* Ilex margratesavage
* Ilex × meserveae
* Ilex microdonta
Ilex montana Torrey & A.Gray – Mountain winterberry (Eastern
United States )
Ilex mucronata (L.) M.Powell, Savol., ">
A contorted hedgehog holly
Ilex aquifolium 'Ferox'
Ilex × meserveae
Ilex pernyi from west China.
* ^ Sunset Western Garden Book 1995:606–607
* ^ "Index Nominum Genericorum". Smithsonian Institution. 2013.
Retrieved 19 February 2017.
* ^ "Kew Plants and Fungi". 2012-04-11.
* ^ "Noxious Weed Control Board (NWCB) - WA State - Weed Detail
* ^ "English Holly". Invasive
Species Week. Archived from the
original on 6 September 2015.
* ^ "Cal-IPC: Ilex aquifolium". cal-ipc.org.
* ^ https://www.portlandoregon.gov/auditor/34460?a=322280
* ^ Shorter Oxford English dictionary, 6th ed. United Kingdom:
Oxford University Press. 2007. p. 3804. ISBN 0199206872 .
* ^ Abbe, Elfriede Martha (1965). The plants of Virgil\'s Georgics.
Cornell University Press. p. 88.
* ^ Middle English Holin, later Hollen. Skeat, Walter William
(1887). Principles of English etymology, Volume 1. London, U.K.:
Clarendon Press. p. 371.
* ^ T. F. Hoad, English Etymology, Oxford University Press, 1993
(ISBN 0-19-283098-8 ), p. 218b.
* ^ Skeat, Walter William (2005). A Concise Etymological Dictionary
of the English Language. Cosimo, Inc. p. 244. ISBN 978-1-59605-092-1 .
* ^ "HOUX : Etymologie de HOUX". cnrtl.fr.
* ^ Pick, Edward (1869). An etymological dictionary of the French
language. Murray. p. 106.
* ^ Skeat, Walter William (1882). An etymological dictionary of the
English language. Clarendon Press. p. 269.
* ^ Nettleship, Henry (1889). Contributions to Latin lexicography.
Clarendon Press. p. 27.
* ^ Loizeau, P.-A.; Barriera G.; Manen J.-F.; Broennimann O.
(2005). "Towards an understanding of Ilex L. (Aquifoliaceae) on a
World-wide scale". In Friis I.; Balslev H.
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. 55. Kongelige Danske videnskabernes
selskab. pp. 507–517. ISBN 978-87-7304-304-2 . Retrieved 9 April
* ^ Germplasm Resources Information Network: Ilex mucronata
Archived 5 June 2011 at the
Wayback Machine .
* ^ Powell, M., Savolainen, V., Cuénod, P., Manen, J. F., Gustavo
C. Giberti; Lidia Poggio (2005). "Molecular analyses of the genus Ilex
(Aquifoliaceae) in southern South America, evidence from AFLP and ITS
American Journal of Botany . 92 (2): 352–369. doi
JSTOR 4123880 . PMID 21652411 .
* ^ International Union for Conservation of Nature and Natural
Resources (IUCN) (2007): 2007IUCN Red List of Threatened Species:Ilex]
* ^ Kellie P. Burris; Federico M. Harte; P. Michael Davidson; C.
Neal Stewart Jr; Svetlana Zivanovic (2012). "Composition and bioactive
properties of yerba mate (
Ilex paraguariensis A. St.-Hil.): A review".
Chilean Journal of Agricultural Research. 72 (2): 268–274.
* ^ A B "Toxicity of Holly". Retrieved 3 March 2014.
* ^ A B C "Holly". webmd.com. Retrieved 3 March 2014.
* ^ "American Holly". aspca. Retrieved 3 March 2014.
* ^ Austin, Daniel F. (2004). Florida Ethnobotany. CRC Press. p.
* ^ RHS A-Z encyclopedia of garden plants. United Kingdom: Dorling
Kindersley. 2008. p. 1136. ISBN 1405332964 .
* ^ Huxley, A., ed. (1992). New RHS Dictionary of Gardening.
Macmillan ISBN 0-333-47494-5 .
* ^ Northumbria Police: Security starts at the Garden Gate
* ^ 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.
* ^ A B C D E Mosteller, Angie (2008). Christmas. Itasca Books. pp.
86–87, 89. ISBN 9781607910084 .
* ^ Ciesla, William M. (2002). Non-wood Forest Products from
Temperate Broad-leaved Trees.
Food and Agriculture Organization of the
United Nations. p. 13. ISBN 9789251048559 .
Holly is still a popular
Christmas decoration among Christian cultures.
* ^ Struthers, Jane (4 October 2010). The Book of Christmas. Ebury
Publishing. p. 218. ISBN 9781448148936 . The British native holly
(Ilex aquifolim) has tremendous religious significance at Christmas.
Its prickly leave 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.
* ^ Little Jr., Elbert L.; Roger G. Skolmen (1989). "Kāwaʻu,
Hawaiian holly" (
United States Forest Service .
* ^ Ulloa Ulloa padding:0.75em; background:#f9f9f9;"> Find more
aboutHOLLYat's sister projects
* Definitions from Wiktionary
* Media from Commons
* Quotations from Wikiquote
* Taxonomy from Wikispecies
* Eichhorn, Markus (March 2011). "The
Holly Tree". Test Tube. Brady
Haran for the
University of Nottingham
University of Nottingham .
Aquifoliaceae in BoDD – Botanical Dermatology Database
* Wd : Q117085
* EoL : 40226
FloraBase : 22454
* FOC : 116366
Fossilworks : 319957
* GRIN : 6018
* IPNI : 60437220-2
* ITIS : 27980
* NCBI : 4295
* PLANTS : ILEX