An Oak is a tree or shrub in the genus Quercus (pronounced /ΛkwÉrkÉs/; Latin "oak tree"), of which about 600 species exist on earth. "Oak" may also appear in the names of species in related genera, notably Lithocarpus. The genus is native to the northern hemisphere, and includes deciduous and evergreen species extending from cold latitudes to tropical Asia and the Americas.
Oaks have spirally arranged leaves, with a lobed margin in many species; some have serrated leaves or entire leaves with a smooth margin. The flowers are catkins, produced in spring. The fruit is a nut called an acorn, borne in a cup-like structure known as a cupule; each acorn contains one seed (rarely two or three) and takes 6â18 months to mature, depending on species. The live oaks are distinguished for being evergreen, but are not actually a distinct group and instead are dispersed across the genus.
Oak trees are flowering plants. The genus is divided into two subgenera and a number of sections:
 Subgenus Quercus
The Subgenus Quercus is divided into the following sections:
- Sect. Quercus (synonyms Lepidobalanus and Leucobalanus), the white oaks of Europe, Asia and North America. Styles are short; acorns mature in 6 months and taste sweet or slightly bitter; the inside of acorn shell is hairless. The leaves mostly lack a bristle on their lobe tips, which are usually rounded.
- Sect. Mesobalanus, Hungarian oak and its relatives of Europe and Asia. Styles long; acorns mature in about 6 months and taste bitter; the inside of acorn shell is hairless. The section Mesobalanus is closely related to section Quercus and sometimes included in it.
- Sect. Cerris, the Turkey oak and its relatives of Europe and Asia. Styles long; acorns mature in 18 months and taste very bitter. The inside of the acorn shell is hairless. Its leaves typically have sharp lobe tips, with bristles at the lobe tip.
- Sect. Protobalanus, the Canyon live oak and its relatives, in southwest United States and northwest Mexico. Styles short, acorns mature in 18 months and taste very bitter. The inside of the acorn shell appears woolly. Leaves typically have sharp lobe tips, with bristles at the lobe tip.
- Sect. Lobatae (synonym Erythrobalanus), the red oaks of North America, Central America and northern South America. Styles long, acorns mature in 18 months and taste very bitter. The inside of the acorn shell appears woolly. The actual nut is encased in a thin, clinging, papery skin. Leaves typically have sharp lobe tips, with spiny bristles at the lobe.
 Subgenus Cyclobalanopsis
- The ring-cupped oaks of eastern and southeastern Asia. Evergreen trees growing 10â40 m tall. They are distinct from subgenus Quercus in that they have acorns with distinctive cups bearing concrescent rings of scales; they commonly also have densely clustered acorns, though this does not apply to all of the species. The Flora of China treats Cyclobalanopsis as a distinct genus, but most taxonomists consider it a subgenus of Quercus. It contains about 150 species.
Interspecific hybridisation is quite common among oaks, but usually only between species within the same section and most common in the white oak group (subgenus Quercus, section Quercus; see List of Quercus species). Inter-section hybrids, except between species of sections Quercus and Mesobalanus, are unknown. Recent systematic studies appear to confirm a high tendency of Quercus species to hybridize because of a combination of factors. White oaks are unable to discriminate against pollination by other species in the same section because they are wind pollinated and they have weak internal barriers to hybridisation, hybridization produces functional seeds and fertile hybrid offspring. Ecological stresses, especially near habitat margins, can also cause a breakdown of mate recognition as well as a reduction of male function (pollen quantity and quality) in one parent species.
Frequent hybridisation among oaks has consequences for oak populations around the world; most notably, hybridization has produced large populations of hybrids with copious amounts of introgression, and the evolution of new species. Frequent hybridisation and high levels of introgression have caused different species in the same populations to share up to 50% of their genetic information. The high rates of hybridisation and introgression, produces genetic data that often does not differentiate between two clearly morphologically distinct species, but instead differentiates populations. Numerous hypotheses have been proposed to explain how oak species are able to remain morphologically and ecologically distinct with such high levels of gene flow, but the problem is still largely a mystery to botanists.
The Fagaceae, or oak family, is a very slowly evolving clade compared to other angiosperms, and the hybridisation patterns in Quercus pose a great challenge to the concept of a species. A species is often defined as a group of âactually or potentially interbreeding populations which are reproductively isolated from other such groups.â By this definition, many species of Quercus would be lumped together according to their geographic and ecological habitat, despite clear distinctions in morphology and, to a large extent, genetic data.
Heart of oak beams
of the frame of Saint-Girons church in Monein
Oak wood has a density of about 0.75 g/cmâ³, great strength and hardness, and is very resistant to insect and fungal attack because of its high tannin content. It also has very attractive grain markings, particularly when quartersawn. Wide, quarter-sawn boards of oak have been prized since the Middle Ages for use in interior paneling of prestigious buildings such as the debating chamber of the British House of Commons in London, England, and in the construction of fine furniture. Oak wood, from Quercus robur and Quercus petraea, was used in Europe for the construction of ships, especially naval men of war, until the 19th century, and was the principal timber used in the construction of European timber-framed buildings. Today oak wood is still commonly used for furniture making and flooring, timber frame buildings, and for veneer production. Barrels in which red wines, sherry, brandy and spirits such as Scotch whisky and Bourbon whiskey are aged are made from European and American oak. The use of oak in wine can add many different dimensions to wine based on the type and style of the oak. Oak barrels, which may be charred before use, contribute to the colour, taste, and aroma, of the contents, imparting a desirable oaky vanillin flavour to these drinks. The great dilemma for wine producers is to choose between French and American oakwoods. French oaks (Quercus robur, Q. petraea) give the wine greater refinement and are chosen for best wines since they increase the price compared to those aged in American oak wood. American oak contributes greater texture and resistance to ageing, but produces more violent wine bouquets. Oak wood chips are used for smoking fish, meat, cheeses and other foods.
A cross section of the trunk of a cork oak
, Quercus suber
The bark of Quercus suber, or Cork oak, is used to produce wine stoppers (corks). This species grows in the Mediterranean Sea region, with Portugal, Spain, Algeria and Morocco producing most of the world's supply. Of the North American oaks, the Northern red oak Quercus rubra is the most prized of the red oak group for lumber, all of which is marketed as red oak regardless of the species of origin. It is not good for outdoor use due to the open capillaries. One can blow air through an end grain piece 10 inches long to make bubbles come out in a glass of water. These opening give fungus easy access when the finish deteriorates. The standard for the lumber of the white oak group, all of which is marketed as white oak, is the White Oak Quercus alba. White Oak is often used to make wine barrels. The wood of the deciduous Pedunculate Oak Quercus robur and Sessile Oak Quercus petraea account for most of the European oak production, but evergreen species, such as Holm oak Quercus ilex, and Cork oak Quercus suber also produce valuable timber.
The bark of the White Oak is dried and used in medical preparations. Oak bark is also rich in tannin, and is used by tanners for tanning leather. Acorns are used for making flour or roasted for acorn coffee. Oak galls were used for centuries as the main ingredient in manuscript ink, harvested at a specific time of year.
Japanese oak is used in the making of professional drums from manufacturer Yamaha Drums. The rough, hard surface of oak gives the drum a brighter and louder tone compared to traditional drum materials such as maple and birch.
A number of kinds of truffles, including the two well known varieties, the black PΓ©rigord truffle and the white Piedmont truffle, have symbiotic relationships with oak.
 Diseases and pests
Sudden Oak Death (Phytophthora ramorum) is a water mould that can kill oaks within just a few weeks. Oak Wilt, caused by the fungus Ceratocystis fagacearum (a fungus closely related to Dutch Elm Disease), is also a lethal disease of some oaks, particularly the red oaks (the white oaks can be infected but generally live longer). Other dangers include wood-boring beetles, as well as root rot in older trees which may not be apparent on the outside, often only being discovered when the trees come down in a strong gale. Oak apples are galls on oaks made by the gall wasp. The female kermes scale causes galls to grow on kermes oak. Oaks are used as food plants by the larvae of Lepidoptera (butterfly and moth) species such as the Gypsy Moth, Lymantria dispar, which can defoliate oak and other broadleaved tree species in North America.
A considerable number of galls are found on oak leaves, buds, flowers, roots, etc. Examples are Oak artichoke gall, Oak Marble gall, Oak apple gall, Knopper gall, and Spangle gall.
A number of species of fungus cause powdery mildew on oak species. In Europe the species, Erysiphe alphitoides is the most common cause.
A new, as yet little understood disease of mature oaks, Acute oak decline, has been reported in parts of the UK since 2009.
The leaves and acorns of the oak tree are poisonous to cattle, horses, sheep, and goats in large amounts due to the toxin tannic acid, and cause kidney damage and gastroenteritis. Additionally, once livestock have a taste for the leaves and acorns, they may seek them out. Symptoms of poisoning include lack of appetite, depression, constipation, diarrhea (which may contain blood), blood in urine, and colic. The exception to livestock and oak toxicity is the domestic pig, which may be fed entirely on acorns in the right conditions, and has traditionally been pastured in oak woodlands (such as the Spanish dehesa and the English system of pannage) for hundreds of years. Acorns are also edible to humans in processed form, and are a staple part of the forage consumed by wildlife.
 Cultural significance
 National symbol
The oak is a common symbol of strength and endurance and has been chosen as the national tree of many countries. Already an ancient Germanic symbol (in the form of the Donar Oak, for instance), certainly since the early nineteenth century it stands for the nation of Germany. In 2004 the Arbor Day Foundation, held a vote for the official National Tree of the United States of America. In November 2004, Congress passed legislation designating the oak as America's Nation Tree.
Other countries have also designated the oak as their national tree including England, Estonia, France, Germany, Moldova, Latvia, Lithuania, Poland, the United States, Basque Country, Wales, Bulgaria, and Serbia.
 Oaks as regional and state symbols
The oak is the emblem of County Londonderry in Northern Ireland, as a vast amount of the county was covered in forests of the tree until relatively recently. The name of the county comes from the city of Derry, which originally in Irish was known as Doire meaning oak.
Iowa has designated the oak as its official state tree in 1961, and the White Oak is the state tree of Connecticut, Illinois and Maryland. The Northern Red Oak is the provincial tree of Prince Edward Island, as well as the state tree of New Jersey. The Live Oak is the State Tree of Georgia.
 Military use
Oak leaves are traditionally an important part of German Army regalia. They also symbolize rank in the United States Armed Forces. A gold oak leaf indicates an O-4 (Major or Lt. Commander), whereas a silver oak leaf indicates an O-5 (Lt. Colonel or Commander). Arrangements of oak leaves, acorns and sprigs indicate different branches of the United States Navy Staff corps officers. Oak leaves are embroidered onto the covers worn by field grade officers and flag officers in the United States armed services.
 Political use
The oak tree is used as a symbol by a number of political parties. It is the symbol of the Conservative Party in the United Kingdom, and formerly of the Progressive Democrats in Ireland. In the cultural arena, the oakleaf is the symbol of the National Trust (UK) and The Royal Oak Foundation.
In Celtic mythology, it is the tree of doors, believed to be a gateway between worlds, or a place where portals could be erected.
In Norse mythology, the oak was sacred to the thunder god, Thor. Some scholars speculate that this is because the oak, as the largest tree in northern Europe, was the one most often struck by lightning. Thor's Oak was a sacred tree of the Germanic Chatti tribe. Its destruction marked the Christianisation of the heathen tribes by Saint Boniface.
In Classical mythology, the oak was a symbol of Zeus and his sacred tree. An example is the oracle of Dodona, which in prehistory consisted solely of a holy oak.
In the Bible, the oak tree at Shechem is the site where Jacob buries the foreign gods of his people (Gen. 35:4) . In addition, Joshua erects a stone under an oak tree as the first covenant of the Lord (Josh. 24.25-7). In Isaiah 61, the prophet refers to the Israelites as "Oaks of Righteousness".
In Slavonic mythology, oak was the most important tree of the god Perun.
Several individual oak trees, such as the Royal Oak in Britain and the Charter Oak in the United States, are of great historical or cultural importance; for a list of important oaks, see Individual oak trees.
"The Proscribed Royalist, 1651", a famous painting by John Everett Millais, depicted a Royalist fleeing from Cromwell's forces and hidden in an oak. Millais painted the picture in Hayes, Kent, from a local oak tree that became known as the Millais Oak.
The city of Raleigh, N.C. is known as "The City of Oaks."
The Jurupa Oak tree, a clonal colony of Quercus palmeria or Palmerâs Oak found in Riverside County, California, is believed to be the world's oldest organism at 13,000 years.
 Historical note on Linnaean species
Linnaeus described only five species of oak from eastern North America, based on general leaf form. These were White oak, Quercus alba; Chestnut oak, Q. Montana; Red oak, Q. rubra; Willow oak Q. phellos; and Water oak, Q. nigra. Because he was dealing with confusing leaf forms, the Q. prinus and Q. rubra specimens actually included mixed foliage of more than one species. For that reason, some taxonomists in the past proposed different names for these two species (Q. montana and Q. borealis, respectively), but the original Linnaean names have now been lectotypified by removing some of the specimens in Linnaeus' herbarium.
 See also
- ^ Sunset Western Garden Book, 1995:606â607
- ^ a b Williams, Joseph H.; Boecklen, William J.; Howard, Daniel J. (2001). "Reproductive processes in two oak (Quercus) contact zones with different levels of hybridisation". Heredity 87 (6): 680â690. doi:10.1046/j.1365-2540.2001.00968.x .
- ^ Arnold, M. L. (1997). Natural Hybridization and Evolution. New York: Oxford University Press. ISBN 0195099745 .
- ^ Conte, L.; Cotti, C.; Cristofolini, G. (2007). "Molecular evidence for hybrid origin of Quercus crenata Lam. (Fagaceae) from Q-cerris L. and Q-suber L.". Plant Biosystems 141 (2): 181â193. doi:10.1080/11263500701401463 .
- ^ Gomory, D.; Schmidtova, J. (2007). "Extent of nuclear genome sharing among white oak species (Quercus L. subgen. Lepidobalanus (Endl.) Oerst.) in Slovakia estimated by allozymes". Plant Systematics and Evolution 266 (3-4): 253â264. doi:10.1007/s00606-007-0535-0 .
- ^ Kelleher, C. T.; Hodkinson, T. R.; Douglas, G. C.; Kelly, D. L. (2005). "Species distinction in Irish populations of Quercus petraea and Q. robur: Morphological versus molecular analyses". Annals of Botany 96 (7): 1237â1246. doi:10.1093/aob/mci275. PMID 16199484 .
- ^ Frascaria, N.; Maggia, L.; Michaud, M.; Bousquet, J. (1993). "The RBCL Gene Sequence from Chestnut Indicates a Slow Rate of Evolution in the Fagaceae". Genome 36 (4): 668â671. doi:10.1139/g93-089. PMID 8405983 .
- ^ Manos, P. S.; Stanford, A. M. (2001b). "The historical biogeography of Fagaceae: Tracking the tertiary history of temperate and subtropical forests of the Northern Hemisphere". International Journal of Plant Sciences 162 (Suppl. 6): S77âS93. doi:10.1086/323280 .
- ^ Raven, Peter H.; Johnson, George B.; Losos, Jonathan B.; Singer, Susan R. (2005). Biology (Seventh ed.). New York: McGraw Hill. ISBN 0071111824 .
- ^ a b (Spanish) La crianza del vino La RazΓ³n 23 de Agosto de 2007
- ^ 200g Oak Smoked Wensleydale â Williams Deli â tearoom richmond north
- ^ 
- ^ "Truffle Glossary: Black Truffles". thenibble.com. =2010-07-01. http://www.thenibble.com/reviews/main/vegetables/truffle-glossary6.asp#glossary. Retrieved 2010-07-01.
- ^ "Truffle Glossary: White Truffles". thenibble.com. =2010-07-01. http://www.thenibble.com/reviews/main/vegetables/truffle-glossary8.asp#glossary. Retrieved 2010-07-01.
- ^ Tree News Autumn/Winter 2007 page 30
- ^ "Trees: Oak Insects and Diseases: Gypsy Moth". TreeHelp.com. http://www.treehelp.com/trees/oak/oak-insects-gypsymoth.asp. Retrieved 2010-04-27.
- ^ Mougou, A.; Dutech, C.; Desprez-Loustau, M. -L. (2008). "New insights into the identity and origin of the causal agent of oak powdery mildew in Europe". Forest Pathology 38: 275. doi:10.1111/j.1439-0329.2008.00544.x.
- ^ Kinver, Mark (28 April 2010). "Oak disease 'threatens landscape'". BBC News. http://news.bbc.co.uk/2/hi/science_and_environment/10089581.stm. Retrieved 29 April 2010.
- ^ Schierz, Kai Uwe (2004). "Von Bonifatius bis Beuys, oder: Vom Umgang mit heiligen Eichen". in Hardy Eidam, Marina Moritz, Gerd-Rainer Riedel, Kai-Uwe Schierz (in German). Bonifatius: Heidenopfer, Christuskreuz, Eichenkult. Stadtverwaltung Erfurt. pp. 139â45.
- ^ "Trees - Arbor Day Foundation". Arborday.org. http://www.arborday.org. Retrieved 2010-04-27.
- ^ "Oak Trees". arborday.org. http://www.arborday.org/Shopping/Trees/oak.cfm. Retrieved 2010-04-27.
- ^ Pickles, Eric. "The Conservative Party". Conservatives.com. http://www.conservatives.com. Retrieved 2010-04-27.
- ^ http://www.progressivedemocrats.ie/about_us/coalition_government_1989_to_1992/
- ^ Millais, J.G., Life and Letters of Sir John Everett Millais, vol. 1, p.166; See also Arborecology, containing a photograph of the Millais oak
- ^ Thaindian News: Jurupa Oak tree is worldâs oldest organism at 13,000 years . Posted: December 24, 2009
- Byfield, Liz (1990) An oak tree, Collins book bus, London : Collins Educational, ISBN 0-00-313526-8
- Philips, Roger. Trees of North America and Europe, Random House, Inc., New York ISBN 0-394-50259-0, 1979.
- Logan, William B. (2005) Oak : the frame of civilization, New York ; London : W.W. Norton, ISBN 0-393-04773-3
- Paterson, R.T. (1993) Use of trees by livestock, 5: Quercus, Chatham : Natural Resources Institute, ISBN 0-85954-365-X
- Royston, Angela (2000) Life cycle of an oak tree, Heinemann first library, Oxford : Heinemann Library, ISBN 0-431-08391-6
- Savage, Stephen (1994) Oak tree, Observing nature series, Hove : Wayland, ISBN 0-7502-1196-2
- Tansley, Arthur G., Sir (1952) Oaks and oak woods, Field study books, London : Methuen.
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