Fagus Leptoneuron

Beech (genus ''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to subtropical (accessory forest element) and temperate (as dominant element of mesophytic forests) Eurasia and North America. There are 14 accepted species in two distinct subgenera, ''Englerianae'' and ''Fagus''. The subgenus ''Englerianae'' is found only in East Asia, distinctive for its low branches, often made up of several major trunks with yellowish bark. The better known species of subgenus ''Fagus'' are native to Europe, western and eastern Asia and eastern North America. They are high-branching trees with tall, stout trunks and smooth silver-grey bark.

The European beech ''Fagus sylvatica'' is the most commonly cultivated species, yielding a utility timber used for furniture construction, flooring and engineering purposes, in plywood, and household items. The timber can be used to build homes. Beechwood makes excellent firewood. Slats of washed beech wood are spread around the bottom of fermentation tanks for Budweiser beer. Beech logs are burned to dry the malt used in some German smoked beers. Beech is also used to smoke Westphalian ham, andouille sausage, and some cheeses.

Description

Beeches are monoecious, bearing both male and female flowers on the same plant. The small flowers are unisexual, the female flowers borne in pairs, the male flowers wind-pollinating catkins. They are produced in spring shortly after the new leaves appear. The fruit of the beech tree, known as beechnuts or mast, is found in small burrs that drop from the tree in autumn. They are small, roughly triangular, and edible, with a bitter, astringent, or mild and nut-like taste.

The European beech (''Fagus sylvatica'') is the most commonly cultivated, although few important differences are seen between species aside from detail elements such as leaf shape. The leaves of beech trees are entire or sparsely toothed, from long and broad.

The bark is smooth and light gray. The fruit is a small, sharply three-angled nut (fruit), nut long, borne singly or in pairs in soft-spined husks long, known as cupules. The husk can have a variety of spine- to scale-like appendages, the character of which is, in addition to leaf shape, one of the primary ways beeches are differentiated. The nuts are called beechnuts or beech mast and have a bitter taste (though not nearly as bitter as acorns) and a high tannin content.

Taxonomy and systematics

The most recent classification system of the genus recognizes 14 species in two distinct subgenera, subgenus ''Englerianae'' and ''Fagus''. Beech species can be diagnosed by Phenotype, phenotypical and/or Genotype, genotypical traits. Species of subgenus ''Engleriana'' are found only in East Asia, and are notably distinct from species of subgenus ''Fagus'' in that these beeches are low-branching trees, often made up of several major trunks with yellowish bark and a substantially different Nuclear DNA, nucleome (nuclear DNA), especially in noncoding, highly variable gene regions such as the Spacer DNA, spacers of the nuclear-encoded Ribosomal DNA, ribosomal RNA genes (ribosomal DNA). Further differentiating characteristics include the whitish bloom on the underside of the leaves, the visible tertiary leaf veins, and a long, smooth cupule-peduncle. Originally proposed but not formalized by botanist Chung-Fu Shen in 1992, this group comprised two Japanese species, ''Fagus japonica, F. japonica'' and ''Fagus okamotoi, F. okamotoi,'' and one Chinese species, ''Fagus engleriana, F. engleriana''. While the status of ''F. okamotoi'' remains uncertain, the most recent systematic treatment based on morphological and genetic data confirmed a third species, ''F. multinervis,'' endemic to Ulleungdo, a South Korean island in the Sea of Japan. The beeches of Ulleungdo have been traditionally treated as a subspecies of ''F. engleriana,'' to which they are phenotypically identical, or as a variety of ''F. japonica.'' The differ from their siblings by their unique Nuclear DNA, nuclear and Chloroplast DNA, plastid genotypes.

The better known subgenus ''Fagus'' beeches are high-branching with tall, stout trunks and smooth silver-gray bark. This group includes five extant species in continental and insular East Asia (''Fagus crenata, F. crenata, Fagus longipetiolata, F. longipetiolata, Fagus lucida, F. lucida'', and the Cryptic speciation, cryptic sister species ''Fagus hayatae, F. hayatae'' and ''Fagus pashanica, F. pashanica'')'','' two pseudo-cryptic species in eastern North America (Fagus grandifolia, ''F. grandifolia'''', Fagus mexicana, F. mexicana''), and a species complex of at least four species (''Fagus caspica, F. caspica, Fagus hohenackeriana, F. hohenackeriana, Fagus orientalis, F. orientalis, Fagus sylvatica, F. sylvatica'') in Western Eurasia. Their genetics are highly complex and include both species-unique alleles as well as alleles and ribosomal DNA spacers that are shared between two or more species. The western Eurasian species are characterized by morphological and genetical gradients.

Research suggests that the first representatives of the modern-day genus were already present in the Paleocene of Arctic North America (western Greenland) and quickly radiated across the high latitudes of the Northern Hemisphere, with a first Biodiversity, diversity peak in the Miocene of northeastern Asia. The contemporary species are the product of past, repeated Reticulate evolution, reticulate evolutionary processes (Outcrossing, outbreeding, introgression, Hybrid (biology), hybridization). As far as studied, heterozygosity and intragenomic variation are common in beech species, and their chloroplast genomes are nonspecific with the exception of the Western Eurasian and North American species.

''Fagus'' is the first Divergent evolution, diverging lineage in the evolution of the Fagaceae family, which also includes oaks and chestnuts. The oldest fossils that can be assigned to the beech lineage are 81–82 million years old pollen from the Late Cretaceous of Wyoming, United States. The southern beeches (genus ''Nothofagus'') historically thought closely related to beeches, are treated as members of a separate family, the Nothofagaceae (which remains a member of the order Fagales). They are found throughout the Southern Hemisphere in Australia, New Zealand, New Guinea, New Caledonia, as well as Argentina and Chile (principally Patagonia and Tierra del Fuego).

Species

Species treated in Denk et al. (2024) and listed in Plants of the World Online, Plants of the World Online (POWO):

Natural and potential hybrids

Phylogeny

A cladogram of 11 beech species is shown below.

Fossil species

Numerous species have been named globally from the fossil record spanning from the Cretaceous to the Pleistocene.

*†''Fagus aburatoensis''
*†''Fagus alnitifolia''
*†''Fagus altaensis''
*†''Fagus ambigua''
*†''Fagus angusta''
*†''Fagus antipofii''
*†''Fagus aperta''
*†''Fagus arduinorum''
*†''Fagus aspera (Berry) Brown, Fagus aspera''
*†''Fagus aspera (Chelebaeva), Fagus aspera'' (jr homonym)
*†''Fagus atlantica''
*†''Fagus attenuata''
*†''Fagus aurelianii''
*†''Fagus australis''
*†''Fagus betulifolia''
*†''Fagus bonnevillensis''
*†''Fagus castaneifolia''
*†''Fagus celastrifolia''
*†''Fagus ceretana''
*†''Fagus chamaephegos''
*†''Fagus chankaica''
*†''Fagus chiericii''
*†''Fagus chinensis''
*†''Fagus coalita''
*†''Fagus cordifolia''
*†''Fagus cretacea''
*†''Fagus decurrens''
*†''Fagus dentata''
*†''Fagus deucalionis''
*†''Fagus dubia (Mirb), Fagus dubia''
*†''Fagus dubia (Watelet), Fagus dubia'' (jr homonym)
*†''Fagus echinata''
*†''Fagus eocenica''
*†''Fagus etheridgei''
*†''Fagus ettingshausenii''
*†''Fagus europaea''
*†''Fagus evenensis''
*†''Fagus faujasii''
*†''Fagus feroniae''
*†''Fagus florinii''
*†''Fagus forumlivii''
*†''Fagus friedrichii''
*†''Fagus gortanii''
*†''Fagus grandifoliiformis''
*†''Fagus gussonii''
*†''Fagus haidingeri''
*†''Fagus herthae''
*†''Fagus hitchcockii''
*†''Fagus hondoensis''
*†''Fagus hookeri''
*†''Fagus horrida''
*†''Fagus humata''
*†''Fagus idahoensis''
*†''Fagus inaequalis''
*†''Fagus incerta''
*†''Fagus integrifolia''
*†''Fagus intermedia''
*†''Fagus irvajamensis''
*†''Fagus japoniciformis''
*†''Fagus japonicoides''
*†''Fagus jobanensis''
*†''Fagus jonesii''
*†''Fagus juliae''
*†''Fagus kitamiensis''
*†''Fagus koraica''
*†''Fagus kraeuselii''
*†''Fagus kuprianoviae''
*†''Fagus lancifolia'' (nomen nudum)
*†''Fagus langevinii''
*†''Fagus laptoneura''
*†''Fagus latissima''
*†''Fagus leptoneuron''
*†''Fagus macrophylla''
*†''Fagus maorica''
*†''Fagus marsillii''
*†''Fagus menzelii''
*†''Fagus microcarpa''
*†''Fagus miocenica''
*†''Fagus napanensis''
*†''Fagus nelsonica''
*†''Fagus oblonga (Suzuki), Fagus oblonga''
*†''Fagus oblonga (Andreánszky), Fagus oblonga''
*†''Fagus obscura''
*†''Fagus olejnikovii''
*†''Fagus orbiculatum''
*†''Fagus orientaliformis''
*†Fagus orientalis, ''Fagus orientalis'' var ''fossilis''
*†Fagus orientalis, ''Fagus orientalis'' var ''palibinii''
*†''Fagus pacifica''
*†''Fagus palaeococcus''
*†''Fagus palaeocrenata''
*†''Fagus palaeograndifolia''
*†''Fagus palaeojaponica''
*†''Fagus pittmanii''
*†''Fagus pliocaenica'' (jr homonym)
*†''Fagus pliocenica''
*†''Fagus polycladus''
*†''Fagus praelucida''
*†''Fagus praeninnisiana''
*†''Fagus praeulmifolia''
*†''Fagus prisca''
*†''Fagus pristina''
*†''Fagus producta''
*†''Fagus protojaponica''
*†''Fagus protolongipetiolata''
*†''Fagus protonucifera''
*†''Fagus pseudoferruginea''
*†''Fagus pygmaea''
*†''Fagus pyrrhae''
*†''Fagus salnikovii''
*†''Fagus sanctieugeniensis''
*†''Fagus saxonica''
*†''Fagus schofieldii''
*†''Fagus septembris''
*†''Fagus shagiana''
*†''Fagus stuxbergii''
*†''Fagus subferruginea''
*†''Fagus succinea''
*†Fagus sylvatica, ''Fagus sylvatica'' var ''diluviana''
*†Fagus sylvatica, ''Fagus sylvatica'' var ''pliocenica''
*†''Fagus tenella''
*†''Fagus uemurae''
*†''Fagus uotanii''
*†''Fagus vivianii''
*†''Fagus washoensis''

Fossil species formerly placed in ''Fagus'' include:

*†''Alnus paucinervis''
*†''Castanea abnormalis''
*†''Fagopsis longifolia''
*†''Fagopsis undulata''
*†''Fagoxylon grandiporosum''
*†''Fagus-pollenites parvifossilis''
*†''Juglans ginannii'' (new name for ''F. ginannii'')
*†''Nothofagaphyllites novae-zealandiae''
*†''Nothofagus benthamii''
*†''Nothofagus dicksonii''
*†''Nothofagus lendenfeldii''
*†''Nothofagus luehmannii''
*†''Nothofagus magelhaenica''
*†''Nothofagus maidenii''
*†''Nothofagus muelleri''
*†''Nothofagus ninnisiana''
*†''Nothofagus risdoniana''
*†''Nothofagus ulmifolia''
*†''Nothofagus wilkinsonii''
*†''Trigonobalanus minima''

Etymology

The name of the tree in Latin, ''fagus'' (from whence the generic epithet), is cognate with English "beech" and of Indo-European language, Indo-European origin, and played an important role in early debates on the geographical origins of the Proto-Indo-Europeans, Indo-European people, the beech argument. Greek language, Greek φηγός (figós) is from the same root, but the word was transferred to the oak tree (e.g. Iliad 16.767) as a result of the absence of beech trees in southern Greece.

Distribution and habitat

Britain and Ireland

''Fagus sylvatica'' was a late entrant to Great Britain after the last glaciation, and may have been restricted to basic soils in the south of England. Some suggest that it was introduced by Neolithic tribes who planted the trees for their edible nuts. The beech is classified as a native in the south of England and as a non-native in the north where it is often removed from 'native' woods. Large areas of the Chilterns are covered with beech woods, which are habitat to the common bluebell and other flora. The Cwm Clydach National Nature Reserve in southeast Wales was designated for its beech woodlands, which are believed to be on the western edge of their natural range in this steep limestone gorge.

Beech is not native to Ireland; however, it was widely planted in the 18th century and can become a problem shading out the native woodland understory.

Beech is widely planted for hedging and in deciduous woodlands, and mature, regenerating stands occur throughout mainland Britain at elevations below about . The tallest and longest hedge in the world (according to ''Guinness World Records'') is the Meikleour Beech Hedges, Meikleour Beech Hedge in Meikleour, Perth and Kinross, Scotland.

Continental Europe

''Fagus sylvatica'' is one of the most common hardwood trees in north-central Europe, in France constituting alone about 15% of all nonconifers. Balkans, The Balkans are also home to the lesser-known oriental beech (''F. orientalis'') and Crimean beech (''F. taurica'').

As a naturally growing forest tree, beech marks the important border between the European deciduous forest zone and the northern pine forest zone. This border is important for wildlife and fauna.

In Denmark and Scania at the southernmost peak of the Scandinavian peninsula, southwest of the natural spruce boundary, it is the most common forest tree. It grows naturally in Denmark and southern Norway and Sweden up to about 57–59°N. The most northern known naturally growing (not planted) beech trees are found in a small grove north of Bergen on the west coast of Norway. Near the city of Larvik is the largest naturally occurring beech forest in Norway, Bøkeskogen.

Some research suggests that early agriculture patterns supported the spread of beech in continental Europe. Research has linked the establishment of beech stands in Scandinavia and Germany with cultivation and fire disturbance, i.e. early agricultural practices. Other areas which have a long history of cultivation, Bulgaria for example, do not exhibit this pattern, so how much human activity has influenced the spread of beech trees is as yet unclear.

The Ancient and Primeval Beech Forests of the Carpathians and Other Regions of Europe, primeval beech forests of the Carpathians are also an example of a singular, complete, and comprehensive forest dominated by a single tree species - the beech tree. Forest dynamics here were allowed to proceed without interruption or interference since the last ice age. Nowadays, they are amongst the last pure beech forests in Europe to document the undisturbed postglacial repopulation of the species, which also includes the unbroken existence of typical animals and plants. These virgin beech forests and similar forests across 12 countries in continental Europe were inscribed on the World Heritage Site, UNESCO World Heritage List in 2007.

North America

The American beech (''Fagus grandifolia'') occurs across much of the eastern United States and southeastern Canada, with a disjunct sister species in Mexico (''F. mexicana''). It is the only extant (surviving) ''Fagus'' species in the Western Hemisphere. Before the Pleistocene Ice Age, it is believed to have spanned the entire width of the continent from the Atlantic Ocean to the Pacific but now is confined to the east of the Great Plains. ''F. grandifolia'' tolerates hotter climates than European species but is not planted much as an ornamental due to slower growth and less resistance to urban pollution. It most commonly occurs as an overstory component in the northern part of its range with sugar maple, transitioning to other forest types further south such as beech-magnolia. American beech is rarely encountered in developed areas except as a remnant of a forest that was cut down for land development.

The dead brown leaves of the American beech remain on the branches until well into the following spring, when the new buds finally push them off.

Asia

East Asia is home to eight species of ''Fagus'', only one of which (''F. crenata'') is occasionally planted in Western countries. Smaller than ''F. sylvatica'' and ''F. grandifolia'', this beech is one of the most common hardwoods in its native range.

Ecology

Beech grows on a wide range of soil types, acidic or basic, provided they are not waterlogged. The tree canopy casts dense shade and thickens the ground with leaf litter.

In North America, they can form Beech-maple forest, beech-maple climax community, climax forests by partnering with the sugar maple.

The beech blight aphid (''Grylloprociphilus imbricator'') is a common pest of American beech trees. Beeches are also used as food plants by some species of Lepidoptera.

Beech bark is extremely thin and scars easily. Since the beech tree has such delicate bark, carvings, such as lovers' initials and other forms of graffiti, remain because the tree is unable to heal itself.

Diseases

Beech bark disease is a fungal infection that attacks the American beech through damage caused by scale insects. Infection can lead to the death of the tree.

Beech leaf disease is a disease that affects American beeches spread by the newly discovered nematode, ''Litylenchus crenatae mccannii''. This disease was first discovered in Lake County, Ohio, in 2012 and has now spread to over 41 counties in Ohio, Pennsylvania, New York, and Ontario, Canada.
As of 2024, the disease has become widespread in Connecticut, Massachusetts and Rhode Island, and in portions of coastal New Hampshire and coastal and central Maine.

Cultivation

The beech most commonly grown as an ornamental tree is the European beech (''Fagus sylvatica''), widely cultivated in North America as well as its native Europe. Many varieties are in cultivation, notably the weeping beech ''F. sylvatica'' 'Pendula', several varieties of copper or purple beech, the fern-leaved beech ''F. sylvatica'' 'Asplenifolia', and the tricolour beech ''F. sylvatica'' 'Roseomarginata'. The columnar Dawyck beech (''F. sylvatica'' 'Dawyck') occurs in green, gold, and purple forms, named after Dawyck Botanic Garden in the Scottish Borders, one of the four garden sites of the Royal Botanic Garden Edinburgh.

Uses

Wood

Beech wood is an excellent firewood, easily split and burning for many hours with bright but calm flames. Slats of beech wood are washed in caustic soda to leach out any flavour or aroma characteristics and are spread around the bottom of fermentation tanks for Budweiser beer. This provides a complex surface on which the yeast can settle, so that it does not pile up, preventing yeast autolysis (biology), autolysis which would contribute off-flavours to the beer. Beech logs are burned to dry the malt used in German smoked beers. Beech is also used to smoke Westphalian ham, traditional andouille (an offal sausage) from Normandy, and some cheeses.

Some drums are made from beech, which has a tone between those of maple and birch, the two most popular drum woods.

The textile Modal (textile), modal is a kind of rayon often made wholly from reconstituted cellulose of pulped beech wood.

The European species ''Fagus sylvatica'' yields a tough, utility timber. It weighs about 720 kg per cubic metre and is widely used for furniture construction, flooring, and engineering purposes, in plywood and household items, but rarely as a decorative wood. The timber can be used to build chalets, houses, and log cabins.

Beech wood is used for the stocks of military rifles when traditionally preferred woods such as Juglans#Wood, walnut are scarce or unavailable or as a lower-cost alternative.

Food

The edible fruit of the beech tree, known as beechnuts or mast, is found in small burrs that drop from the tree in autumn. They are small, roughly triangular, and edible, with a bitter, astringent, or in some cases, mild and nut-like taste. According to the Roman statesman Pliny the Elder in his work Natural History (Pliny), ''Natural History'', beechnut was eaten by the people of Chios when the town was besieged, writing of the fruit: "that of the beech is the sweetest of all; so much so, that, according to Cornelius Alexander, the people of the city of Chios, when besieged, supported themselves wholly on mast". They can also be roasted and pulverized into an adequate coffee substitute. The leaves can be steeped in liquor to give a light green/yellow liqueur.

Books

In antiquity, the bark of the beech tree was used by Proto-Indo-Europeans, Indo-European people for writing-related purposes, especially in a religious context. Beech wood tablets were a common writing material in Germanic societies before the development of paper. The Old English ''bōc'' has the primary sense of "beech" but also a secondary sense of "book", and it is from ''bōc'' that the modern word derives. In modern German, the word for "book" is ''Buch,'' with ''Buche'' meaning "beech tree". In modern Dutch, the word for "book" is ''boek,'' with ''beuk'' meaning "beech tree". In Swedish, these words are the same, ''bok'' meaning both "beech tree" and "book". There is a similar relationship in some Slavic languages. In Russian and Bulgarian language, Bulgarian, the word for beech is :wikt:бук, бук (''buk''), while that for "letter" (as in a letter of the alphabet) is буква (''bukva''), while Serbo-Croatian language, Serbo-Croatian and Slovene language, Slovene use ":wikt:bukva, bukva" to refer to the tree.

Other

The pigment bistre was made from beech wood soot. Beech Plant litter, litter raking as a replacement for straw in animal husbandry was an old non-timber practice in forest management that once occurred in parts of Switzerland in the 17th century. Beech has been listed as one of the 38 plants whose flowers are used to prepare Bach flower remedies.

See also

* Ancient and Primeval Beech Forests of the Carpathians and Other Regions of Europe
* English Lowlands beech forests
* Weeping Beech (Queens)

References

External links

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Traditional and Modern Use of Beech

{{Authority control

Edible nuts and seeds
Fagus,
Ornamental trees