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An oak is a hardwood tree or shrub in the genus Quercus of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisphere; it includes some 500 species, both deciduous and evergreen. Fossil oaks date back to the Middle Eocene. Molecular phylogeny shows that the genus is divided into Old World and New World clades, but many oak species hybridise freely, making the genus's history difficult to resolve. Ecologically, oaks are keystone species in habitats from Mediterranean semi-desert to subtropical rainforest. They live in association with many kinds of fungi including truffles. Oaks support more than 950 species of caterpillar, many kinds of gall wasp which form distinctive galls, roundish woody lumps such as the oak apple, and a large number of pests and diseases. Oak leaves and acorns contain enough tannin to be toxic to cattle, but pigs are able to digest them safely. Oak timber is strong and hard, and has found many uses in construction and furniture-making. The bark was traditionally used for tanning leather. Wine barrels are made of oak; these are used for aging alcoholic beverages such as sherry and whisky, giving them a range of flavours, colours, and aromas. The spongy bark of the cork oak is used to make traditional wine bottle corks. Almost a third of oak species are threatened with extinction due to climate change, invasive pests, and habitat loss. In culture, the oak tree is a symbol of strength and serves as the national tree of many countries. In Indo-European and related religions, the oak is associated with thunder gods. Individual oak trees of cultural significance include the Royal Oak in Britain, the Charter Oak in the United States, and the Guernica Oak in the Basque Country.
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