An oak is a tree or shrub in the genus Quercus ( ; Latin "oak tree"), of which about 400 species exist. "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 6–18 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.

Classification

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 10–40 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.

Hybridisation

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 “actually or potentially interbreeding populations which are reproductively isolated from other such groups.” 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. Thus, although it may be difficult to place a definition on a species within a genus like Quercus, it is trivial and uninformative to apply the biological species concept to all forms of life.

Uses

Oak wood has a density of about 0.75 g/cm³, 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.

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.

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, ect. Examples are Oak artichoke gall, Oak Marble gall, Oak apple gall, Knopper gall, and Spangle gall.

Toxicity

The leaves and acorns of the oak tree are poisonous to horses in large amounts, due to the toxin tannic acid, and cause kidney damage and gastroenteritis. Additionally, once horses have a taste for the leaves and acorns, they may seek them out. Therefore, horse owners are encouraged to fence out oak trees from their pasture, especially if forage is scarce. Symptoms of poisoning include lack of appetite, depression, constipation, diarrhea (which may contain blood), blood in urine, and colic.

Cultural significance

Political or symbolic

The oak is a common symbol of strength and endurance and has been chosen as the national tree of England, Estonia, France, Germany, Moldova, Latvia, Lithuania, Poland, the United States, Basque Country, Wales, Bulgaria and Serbia. 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.

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.

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.

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.

Mythological

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 the Franks .

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.

The Oak tree is traditionally sacred to Serbs and is widely used throughout Serbia on national and regional symbols both old and new.

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".

Historical

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 a Thousand Oaks."

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.

Gallery

Image:Raunkiaer.jpg|A Pedunculate oak in DenmarkImage:Oak_at_night.JPG|An oak tree at nightImage:Oakbark.jpg|Bark of Quercus roburImage:Spanish-moss-tree.jpg|Southern live oak with spanish mossImage:Quercus mongolica mongolian oak MN 2007.JPG|Quercus mongolica Mongolian Oak in Minnesota Landscape ArboretumImage:Quercus robur JPG (a).jpg|Old oak in Liernu, Belgium.Image:Quercus robur JPG (d2).jpg|The Duke Prosper Oak in Enghien (Belgium).Image:Sivry-Rance AR1aJPG.jpg|Quercus petraea Sessile Oak in Sivry-Rance (Belgium).Image:Kindred Spirit® Hybrid Oak.jpg|Form of Kindred Spirit Hybrid Oak.Image:Angel Oak Johns Island SC horz.jpg|Angel Oak on Johns Island, South Carolina.Image:Robin Hood Major Oak.jpg| The Major Oak is an 800–1000 year old oak in Sherwood Forest, famed as the alleged principal hideout of Robin Hood.File:Mustard Powder Lichen.JPG|The Mustard Powder lichen or Gold Dust lichen is common on old oak bark.

See also

• List of plants poisonous to equines

Notes

1. Sunset Western Garden Book, 1995:606–607
2. Williams, Joseph H., William J. Boecklen, and Daniel J. Howard. 2001 Reproductive processes in two oak (Quercus) contact zones with different levels of hybridisation. Heredity 87: 680–690.
3. Arnold, M. L. 1997. Natural Hybridization and Evolution. Oxford University Press, New York.
4. Conte, L., Cotti, C., and 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): 181–193.
5. Gomory, D. and 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): 253–264.
6. Kelleher, CT., TR Hodkinson, GC Doublas, and DL Kelly. 2005 Species distinction in Irish populations of Quercus petraea and Q. robur: Morphological versus molecular analyses. Annals of Botany 96 (7): 1237–1246.
7. Frascaria, N., L. Maggia, M. Michaud, and J. Bousquet. 1993 The RBCL Gene Sequence from Chestnut Indicates a Slow Rate of Evolution in the Fagaceae. Genome 36 (4): 668–671.
8. Manos, PS., AM Stanford. 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: S77-S93 Suppl. 6.
9. Raven, Peter H., George B. Johnson, Jonathan B. Losos, Susan R. Singer. Biology: Seventh Edition. McGraw Hill, New York, NY 2005.
10. La crianza del vino La Razón 23 de Agosto de 2007
11. 200g Oak Smoked Wensleydale – Williams Deli – tearoom richmond north
12. http://www.swaledalecheese.co.uk/cheese.htm
13. http://www.treehelp.com/trees/oak/oak-insects-gypsymoth.asp
14. http://www.conservatives.com
15. http://www.progressivedemocrats.ie/about_us/coalition_government_1989_to_1992/
16. 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

References

• Byfield, Liz (1990) An oak tree, Collins book bus, London : Collins Educational, ISBN 0-00-313526-8
• 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, 50 p.

External links

• Flora of North America: Quercus
• Flora of China: Quercus
• Flora of China – Cyclobalanopsis
• Flora Europaea: Quercus
• Oak Trees of the Lowcountry (South Carolina) – Beaufort County Library
• Oaks from Bialowieza Forest
• Common Oaks of Florida
• Oak leaves and their use in aquariums
• Oaks of the world