Holly

taxobox
name = Holly


image_width = 240px
image_caption = European Holly ("Ilex aquifolium") leaves and fruit
regnum = Plantae
divisio = Magnoliophyta
classis = Magnoliopsida
subclassis = Asteridae
unranked_ordo = Euasterids II
ordo = Aquifoliales
familia = Aquifoliaceae
familia_authority = DC. ex A.Rich.
genus = "Ilex"
genus_authority = L.
subdivision_ranks = Species
subdivision = About 600, see text

Holly ("Ilex") is a genus of about 600 species of flowering plants in the family Aquifoliaceae, and the only living genus in that family. One other genus, the monotypic "Nemopanthus" (Mountain Holly), was formerly separated from "Ilex" on the basis that its flowers have a reduced calyx and narrow petals, and also in cytology, being tetraploid, whereas "Ilex" is diploid. However, following analysis of molecular data, Mountain Holly has now been merged into "Ilex", as "I. mucronata"; it is closely related to "I. amelanchier". [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.] [Gottlieb, A. M., Giberti, G. C., & Poggio, L. (2005). Molecular analyses of the genus Ilex (Aquifoliaceae) in southern South America, evidence from AFLP and ITS sequence data. "Amer. J. Bot". 92: 352-369. Available [http://www.amjbot.org/cgi/content/full/92/2/352 online] .]

Description and ecology

Hollies are shrubs and trees from 2–25 m tall, with a wide distribution in Asia, Europe, Africa, and North and South America. Most species are found in the tropics and subtropics of America and Asia - China has 204 native species [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=10055] - but this account is based largely on the well-studied species of Europe and North America. The leaves are simple, and can be either deciduous or evergreen depending on the species, and may be entire, finely toothed, or with widely-spaced, spine-tipped serrations. They are mostly dioecious, with male and female flowers on different plants, with some exceptions. Pollination is mainly by bees and other insects. The fruits are small drupes, usually with four to six pits, and range in color from red to purple-black. [Judd, W. S., C. S. Campbell, E. A. Kellogg, P. F. Stevens, and M. J. Donoghue. (2008). "Plant Systematics: A Phylogenetic Approach, Third Edition". Sinauer Associates, Inc., Sunderland, MA. p. 494.] One species in China, "Ilex chapaensis", has large green fruits that are eaten by civets [Tsang, A.C.W. and R.T. Corlett (2005) Reproductive biology of the Ilex species (Aquifoliaceae) in Hong Kong, China. Canadian Journal of Botany 83: 1645-1654] .

Holly berries are mildly toxic and will cause vomiting and/or diarrhea when ingested by people. However they are extremely important food for numerous species of birds, and also are eaten by other wild animals. In the fall and early winter the berries are hard and apparently unpalatable. After being frozen or frosted several times, the berries 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"). The Japanese Beetle ("Popillia japonica") is another well-known animal feeding on holly leaves. Holly is commonly referenced at Christmas time.

Having evolved numerous species that are endemic to islands and small mountain ranges, and being highly useful plants, many hollies are now becoming rare. Tropical species are especially often threatened by habitat destruction and overexploitation, and at least two have become extinct, with numerous others barely surviving.International Union for the Conservation of Nature and Natural Resources (IUCN) (2007): "2007 IUCN Red List of Threatened Species": [http://www.iucnredlist.org/search/search.php?freetext=ilex&modifier=phrase&criteria=wholedb&taxa_species=1&redlistCategory%5B%5D=all&country%5B%5D=all&cty_default=1&aquatic%5B%5D=all&aqu_default=1&regions%5B%5D=all&reg_default=1&habitats%5B%5D=all&threats%5B%5D=all&redlistAssessyear%5B%5D=all&growths%5B%5D=all "Ilex"] ] :)

elected species [Ulloa Ulloa & Jørgensen (1993), eFloras.org (2007a, b), IUCN (2007), RBGE (2007), USDA (2007a, b)]

* "Ilex abscondita"
* "Ilex acutidenticulata"
* "Ilex affinis"
* "Ilex × altaclarensis"
* "Ilex altiplana"
* "Ilex amara" – Caachira
* "Ilex ambigua" – Sand Holly
* "Ilex amelanchier" – Swamp Holly
* "Ilex anomala"
* "Ilex anonoides"
* "Ilex aquifolium" – European Holly, English Holly, Christ's Thorn
* "Ilex aracamuniana"
* "Ilex argentina"
* "Ilex arisanensis"
* "Ilex bioritsensis"
* "Ilex brachyphylla"
* "Ilex brasiliensis"
* "Ilex brevicuspis"
* "Ilex brevipedicellata"
* "Ilex buergeri"
* "Ilex canariensis" – Small-leaved Holly, Acebino
* "Ilex caniensis"
* "Ilex cassine" – Dahoon Holly, Cassena
* "Ilex centrochinensis"
* "Ilex cerasifolia"
* "Ilex chamædryfolia"
* "Ilex chengkouensis"
* "Ilex chinensis"
* "Ilex chuniana"
* "Ilex ciliolata"
* "Ilex ciliospinosa"
* "Ilex cognata"
* "Ilex colchica"
* "Ilex collina"
* "Ilex conocarpa"
* "Ilex cookii" – Cook's Holly
* "Ilex corallina"
* "Ilex coriacea" – Gallberry
* "Ilex cornuta" – Chinese Holly, Horned Holly
* "Ilex costaricensis"
* "Ilex cowanii"
* "Ilex crenata" – Japanese Holly, Box-leaved Holly, "inutsuge" (Japanese)
* "Ilex crepitans"
* "Ilex cyrtura"
* "Ilex dabieshanensis"
* "Ilex davidsei"
* "Ilex decidua" – Deciduous Holly, Meadow Holly, Swamp Holly, Possum Haw
* "Ilex dehongensis"
* "Ilex dimorphophylla"
* "Ilex diospyroides"
* "Ilex dipyrena" – Himalayan Holly
* "Ilex dumosa"
* "Ilex ericoides"
* "Ilex euryoides"
* "Ilex fargesii"
* "Ilex fengqingensis"
* "Ilex fertilis"
* "Ilex florifera"
* "Ilex gardneriana" (extinct: 20th century?)
* "Ilex geniculata"
* "Ilex georgei"
* "Ilex gigantea"
* "Ilex glabella"
* "Ilex glabra" – Evergreen Winterberry, Appalachian Tea, Dye-leaves, Gallberry, Bitter Gallberry, Inkberry
* "Ilex gleasoniana"
* "Ilex goshiensis"
* "Ilex graciliflora"
* "Ilex grandiflora"
* "Ilex guaiquinimae"
* "Ilex guayusa" – Guayusa
* "Ilex guianensis"
* "Ilex harrisii"
* "Ilex holstii"
* "Ilex huachamacariana"
* "Ilex Humboldtiana"
* "Ilex ignicola"
* "Ilex illustris"
* "Ilex integerrima"
* "Ilex integra" – Mochi Tree, Nepal Holly
* "Ilex intricata"
* "Ilex jamaicana"
* "Ilex jauaensis"
* "Ilex jelskii"
* "Ilex karuaiana"
* "Ilex khasiana"
* "Ilex kingiana"
* "Ilex kudingcha"
* "Ilex kusanoi"
* "Ilex lævigata" – Smooth Winterberry
* "Ilex lasseri"
* "Ilex latifolia" – Tarajo Holly, "tarayō" (Japanese)
* "Ilex lechleri"
* "Ilex leucoclada"
* "Ilex longipes"
* "Ilex longzhouensis"
* "Ilex machilifolia"
* "Ilex maclurei"
* "Ilex macoucoua"
* "Ilex macrocarpa"
* "Ilex macropoda"
* "Ilex magnifructa"
* "Ilex maingayi"
* "Ilex marahuacae"
* "Ilex marginata"
* "Ilex margratesavage"
* "Ilex mathewsii"
* "Ilex × meserveae"
* "Ilex microdonta"
* "Ilex mitis"
* "Ilex montana" – Mountain Winterberry
* "Ilex mucronata" – Mountain Holly, Catberry
* "Ilex myrtifolia" – Myrtle Holly, Myrtle Dahoon
* "Ilex neblinensis"
* "Ilex nothofagifolia"
* "Ilex oblonga"
* "Ilex occulta"
* "Ilex opaca" – American Holly, White Holly, Holly Berry
* "Ilex ovalifolia"
* "Ilex palawanica"
* "Ilex pallida"
* "Ilex paraguariensis" – Maté, yerba maté, "erva-mate" (Portuguese)
* "Ilex parvifructa"
* "Ilex patens"
* "Ilex pauciflora"
* "Ilex paujiensis"
* "Ilex pedunculosa"
* "Ilex peiradena"
* "Ilex perado" – Madeiran Holly
* "Ilex perlata"
* "Ilex pernyi" – Perny's Holly
* "Ilex polita"
* "Ilex praetermissa"
* "Ilex pringlei"
* "Ilex pseudobuxus"
* "Ilex puberula"
* "Ilex pubescens"
* "Ilex pubiflora"
* "Ilex purpurea"
* "Ilex qianlingshanensis"
* "Ilex quercetorum"
* "Ilex quercifolia"
* "Ilex rarasanensis"
* "Ilex reticulata"
* "Ilex rotunda"
* "Ilex rugosa"
* "Ilex sclerophylla"
* "Ilex serrata" – Japanese Winterberry
* "Ilex sessilifructa"
* "Ilex shimeica"
* "Ilex sikkimensis"
* "Ilex sintenisii" – Sintenis' Holly
* "Ilex sipapoana"
* "Ilex socorroensis"
* "Ilex spinigera"
* "Ilex spruceana"
* "Ilex steyermarkii"
* "Ilex subrotundifolia"
* "Ilex subtriflora"
* "Ilex sugerokii"
* "Ilex sulcata"
* "Ilex syzygiophylla"
* "Ilex tahanensis"
* "Ilex tateana"
* "Ilex taubertiana"
* "Ilex ternatiflora" (extinct: 20th century?)
* "Ilex theezans"
* "Ilex tiricae"
* "Ilex tolucana"
* "Ilex trachyphylla"
* "Ilex trichocarpa"
* "Ilex tugitakayamensis"
* "Ilex uraiensis"
* "Ilex vaccinoides"
* "Ilex venezuelensis"
* "Ilex venulosa"
* "Ilex verticillata" – Deciduous Holly, Michigan Holly, Swamp Holly, Black Alder, Brook Alder, False Alder, Striped Alder, White Alder, Coralberry, Inkberry, Possum Haw, Fever Bush, Winterberry Holly, American Winterberry, Black Alder Winterberry, Deciduous Winterberry, Virginian Winterberry
* "Ilex vomitoria" – Yaupon Holly, "casseena" (Spanish)
* "Ilex vulcanicola"
* "Ilex wenchowensis"
* "Ilex williamsii"
* "Ilex wilsonii"
* "Ilex yunnanensis"
* "Ilex wugonshanensis"
* "Ilex yuiana"

Etymology

The origin of the word "holly" is Old English _an. "holegn", which is related to Old High German _go. "hulis". The French word for holly, _fr. "houx", derives from the Old High German word, as do 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 _cy. "celyn" and Irish _ga. "cuilleann".

The botanical name "ilex" was the original Latin name for the Holm Oak ("Quercus ilex"), which has similar foliage to common holly, and is occasionally confused with it.

Uses

In many western cultures, holly is a traditional Christmas decoration, used especially in wreaths. The wood is heavy, hard and whitish; one traditional use is for chess pieces, with holly for the white pieces, and ebony for the black. Other uses include turnery, inlay work and as firewood. Looms in the 1800s used holly for the spinning rod. Because holly is dense and can be sanded very smooth, the rod was less likely than other woods to snag threads being used to make cloth.

Many of the hollies are widely used as ornamental plants in gardens and parks. Several hybrids and numerous cultivars have been developed for garden use, among them the very popular "Ilex × altaclerensis" ("I. aquifolium" × "I. perado") and "Ilex × meserveae" ("I. aquifolium" × "I. rugosa").Huxley, A., ed. (1992). "New RHS Dictionary of Gardening". Macmillan ISBN 0-333-47494-5.] Hollies are often used for hedges; the spiny leaves make them difficult to penetrate, and they take well to pruning and shaping. [Northumbria Police: [http://ww2.northumbria.police.uk/ePolicing/Web/WMS.nsf/CrimeReductionContentDocs/CRD000301 Security starts at the Garden Gate] ] In Heraldry, holly is used to symbolise truth.

Between the thirteenth and eighteenth century, before the introduction of turnips, holly was cultivated for use as winter fodder for cattle and sheep. [Spray, M. (1981). Holly as a Fodder in England. "Agricultural History Review" 29 (2): 97. Available [http://www.bahs.org.uk/29n2a3.pdf online] (pdf file). British Agricultural History Society.] Less spiny varieties of holly were preferred, and in practice the leaves growing near the top of the tree have far fewer spines making them more suitable for fodder.

Several holly species are used to make caffeine-rich herbal teas. The South American Yerba Mate ("I. paraguariensis") is boiled for the popular revigorating drinks Mate, and "Chimarrão", and steeped in water for the cold "Tereré". Guayusa ("I. guayusa") is used both as a stimulant and as an admixture to the entheogenic tea ayahuasca; its leaves have the highest known caffeine content of any plant. In North and Central America, Yaupon ("I. vomitoria"), was used by southeastern Native Americans as a ceremonial stimulant and emetic known as "the black drink" [Cherokee: "Gvnega adatasti" (ᎬᏁᎦ ᎠᏓᏔᏍᏘ), "Asi" (ᎠᏏ).] . As the name suggests, the tea's purgative properties were one of its main uses, most often ritually. Evergreen Winterberry (Appalachian Tea, "I. glabra") is a milder substitute for Yaupon. In China, the young leaf buds of "I. kudingcha" are processed in a method similar to green tea to make a tisane called kǔdīng chá (苦丁茶, roughly "bitter spikeleaf tea").

References