Beech(Fagus) is agenusofdeciduoustreesin the familyFagaceae,native to subtropical (accessory forest element) and temperate (as dominant element ofmesophyticforests)EurasiaandNorth America.There are 14 accepted species in two distinct subgenera,EnglerianaeDenk & G.W.GrimmandFagus.[1]The subgenusEnglerianaeis 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 subgenusFagusare native toEurope,western and eastern Asia and eastern North America. They are high-branching trees with tall, stout trunks and smooth silver-grey bark.

Beech
European beech (Fagus sylvatica)
Scientific classificationEdit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Fagales
Family: Fagaceae
Subfamily: Fagoideae
K.Koch
Genus: Fagus
L.
Type species
Fagus sylvatica
Species

Seetext

The European beechFagus sylvaticais 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 excellentfirewood.Slats of washed beech wood are spread around the bottom of fermentation tanks forBudweiserbeer. Beech logs are burned to dry themaltused in some Germansmoked beers.Beech is also used to smokeWestphalian ham,andouillesausage, and some cheeses.

Description

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Leaf ofFagus sylvatica
Beechnuts in autumn

Beeches aremonoecious,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-pollinatingcatkins.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 smallburrsthat 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 asleafshape. The leaves of beech trees are entire or sparsely toothed, from 5–15 centimetres (2–6 inches) long and 4–10 cm (2–4 in) broad.

The bark is smooth and light gray. The fruit is a small, sharply three-anglednut10–15 mm (3858in) long, borne singly or in pairs in soft-spined husks1.5–2.5 cm (58–1 in) 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.[2]The nuts are called beechnuts[3]or beech mast and have a bitter taste (though not nearly as bitter asacorns) and a hightannincontent.

Taxonomy and systematics

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The most recent classification system of the genus recognizes 14 species in two distinct subgenera, subgenusEnglerianaeandFagus.[1]Beech species can be diagnosed byphenotypicaland/orgenotypicaltraits. Species of subgenusEnglerianaare found only in East Asia, and are notably distinct from species of subgenusFagusin that these beeches are low-branching trees, often made up of several major trunks with yellowish bark and a substantially differentnucleome (nuclear DNA),especially in noncoding, highly variable gene regions such as thespacersof the nuclear-encodedribosomal RNA genes (ribosomal DNA).[4][5]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,F. japonicaandF. okamotoi,and one Chinese species,F. engleriana.[2]While the status ofF. okamotoiremains uncertain, the most recent systematic treatment based on morphological and genetic data confirmed a third species,F. multinervis,endemic toUlleungdo,a South Korean island in theSea of Japan.[1]The beeches of Ulleungdo have been traditionally treated as a subspecies ofF. engleriana,to which they are phenotypically identical,[2][6]or as a variety ofF. japonica.[7]The differ from their siblings by their uniquenuclearandplastidgenotypes.[1][8][4]

The better known subgenusFagusbeeches are high-branching with tall, stout trunks and smooth silver-gray bark. This group includes five extant species in continental and insular East Asia (F. crenata,F. longipetiolata,F. lucida,and thecrypticsister speciesF. hayataeandF. pashanica),two pseudo-cryptic species ineastern North America(F. grandifolia,F. mexicana), and aspecies complexof at least four species (F. caspica,F. hohenackeriana,F. orientalis,F. sylvatica) inWestern Eurasia.Their genetics are highly complex and include both species-uniqueallelesas well as alleles and ribosomal DNA spacers that are shared between two or more species.[1]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 thePaleoceneofArcticNorth America (westernGreenland[9]) and quickly radiated across the high latitudes of theNorthern Hemisphere,with a firstdiversitypeak in theMioceneof northeasternAsia.[10][11]The contemporary species are the product of past, repeatedreticulate evolutionary processes(outbreeding,introgression,hybridization).[4]As far as studied, heterozygosity and intragenomic variation are common in beech species,[4][5][8]and their chloroplast genomes are nonspecific with the exception of the Western Eurasian and North American species.[1]

Fagusis the firstdiverginglineage in the evolution of theFagaceaefamily,[9][12]which also includesoaksandchestnuts.[13]The oldest fossils that can be assigned to the beech lineage are 81–82 million years oldpollenfrom theLate CretaceousofWyoming,United States.[9]Thesouthern beeches(genusNothofagus) historically thought closely related to beeches, are treated as members of a separate family, theNothofagaceae(which remains a member of the orderFagales). They are found throughout theSouthern HemisphereinAustralia,New Zealand,New Guinea,New Caledonia,as well asArgentinaandChile(principallyPatagoniaandTierra del Fuego).

Species

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Species treated in Denk et al. (2024) and listed inPlants of the World Online (POWO):[1]

Image Name Subgenus Status, systematic affinity Distribution Accepted as species in POWO as of April 2023[14]
Fagus caspicaDenk & G.W.Grimm– Caspian beech Fagus New species described in 2024;[1]first-diverging lineage within the Western Eurasian group TalyschandElburz Mountains,southeasternAzerbaijanand northernIran Populations included inF. sylvaticasubsp.orientalis
Fagus chieniiW.C.Cheng Fagus Possibly conspecific withF. lucida[6] Probably extinct, described from a single location in China (Sichuan). Individuals recently collected at the type locality were morphologically and genetically indistinguishable fromF. pashanica.[15] Yes
Fagus crenataBlume– Siebold's beech or Japanese beech Fagus Widespread species; complex history connecting it to both the Western Eurasian group and the other East Asian species of subgenusFagus[4] Japan; in the mountains ofKyushu,ShikokuandHonshu,down to sea-level in southernHokkaido. Yes
Fagus englerianaSeemenex Diels– Chinese beech Englerianae Widespread species; continental sister species ofF. japonica[5][8][4] China; south of theYellow River Yes
Fagus grandifoliaEhrh.– American beech Fagus Widespread species; sister species ofF. mexicana[8][4] Eastern North America; from E. Texas and N. Florida, United States, to theSt. Lawrence River,Canada at low to mid altitudes Yes, including Mexican beeches,F. mexicana
Fagus hayataePalib. ex Hayata Fagus Narrowendemicspecies; forming a cryptic sister species pair withF. pashanica[4][1] Taiwan;restricted to the mountains of northern Taiwan Yes
Fagus hohenackerianaPalib.Hohenacker's or Caucasian beech Fagus Dominant tree species of the Pontic andCaucasusMountains; intermediate betweenF. caspicaandF. orientalis.[16][17][18]Its genetic heterogeneity[1][19]may be indicative for ongoing speciation processes. Northeastern Anatolia (Pontic Mountains,Kaçkar Mountains) and Caucasus region (LesserandGreater Caucasus,Georgia,Armenia,Ciscaucasia;down to sea-level in southwestern Georgia) No, populations included inF. sylvaticasubsp.orientalis
Fagus japonicaMaxim. Englerianae Widespread species; insular sister species ofF. engleriana[4][5][8] Japan; Kyushu, Shikoku and Honshu from sea-level up to c. 1500 ma.s.l. Yes
Fagus longipetiolataSeemen Fagus Sym-toparapatricwithF. lucidaandF. pashanica,and sharing alleles with both species in addition to alleles indicating a sister relationship with the JapaneseF. crenata.[4][8] China, south of the Yellow River, into N.Vietnam;in montane areas up to 2400 m a.s.l.[20] Replaced byF. sinensis
Fagus lucidaRehder &E.H.Wilson Fagus Rare species; closest relatives areF. crenata[4][5][6]andF. longipetiolata[4][8] China; south of the Yellow River in montane areas between 800 and 2000 m a.s.l.[21] Yes
Fagus mexicanaMartínez Fagus Narrow endemic sister species ofF. grandifolia. F. mexicanadiffers fromF. grandifoliaby its slender leaves and less-evolved but more polymorphic set of alleles (higher level ofheterozygosity)[4][8] Hidalgo,Mexico; at 1400–2000 m a.s.l. as an element of the subtropical montane mesophilic forest "(bosque mesófilo de montaña) superimposing the tropical lowland rainforests. No, populations included inF. grandifolia
Fagus multinervisNakai Englerianae Narrow endemic species, first diverging lineage within subgenusEnglerianae[4][8] South Korea (Ulleungdo) Yes
Fagus orientalisLipsky– Oriental beech (in a narrow sense) Fagus Sister species ofF. sylvatica[17][18] Southeastern Europe (SEBulgaria,NEGreece,European Turkey) and adjacent northwestern Asia (NW and NAnatolia) No, treated as subspecies ofF. sylvatica
Fagus pashanicaC.C.Yang Fagus Continental sister species ofF. hayatae,with a set of alleles that puts it closer toF. longipetiolataandF. crenatathan its insular sister. China (Hubei, Hunan, Shaanxi, Sichuan, Zhe gian g), at 1300–2300 m a.s.l.(eFlora of China, asF. hayatae[22]) Yes
Fagus sinensisOliv. Fagus Invalid; the original material included material from two much different species:F. englerianaandF. longipetiolata[1][6] China (Hubei), Vietnam Yes, erroneously used as older synonym ofF. longipetiolata
Fagus sylvaticaL.– European beech Fagus Sister species of and closely related toF. orientalis[17][18] Europe Yes

Natural and potential hybrids

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Image Name Parentage Status Distribution
Fagus(×)moesiaca(K. Malý) Czeczott F. sylvatica×F. orientalis No evidence so far for hybrid origin. All individuals addressed asF. moesiacaincluded in genetic studies fell within the variation ofF. sylvatica.[5][23]They may represent a lowland ecotype ofF. sylvatica.[1][24]

Erroneously synonymized by some authors (e.g. POWO) with the CrimeanF. × taurica,from which it differs morphologically and genetically.

SoutheasternBalkans
Fagus okamotoiShen F. crenata×F. japonica? Unique phenotype, described from an area in whichF. crenataandF. japonicaare sympatric. So far, there is no genetic evidence for ongoing gene flow between the two Japanese species, which belong to different subgeneric lineages. Kanto,eastern Honshu
Fagus × tauricaPopl.– Crimean beech F. sylvatica×F. orientaliss.l. Hybrid status not yet tested by genetic data; according toisoenzymeprofiles a less-evolved, relict population ofF. sylvaticaor intermediate betweenF. sylvaticaand the species complex historically addressed as Oriental beech (F. orientalisin a broad sense)[16] Crimean peninsula

Fossil species

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Numerous species have been named globally from the fossil record spanning from theCretaceousto thePleistocene.[25]

Fossil species formerly placed inFagusinclude:[25]

Etymology

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The name of the tree in Latin,fagus(from whence thegeneric epithet), is cognate with English "beech" and ofIndo-Europeanorigin, and played an important role in early debates on the geographical origins of theIndo-European people,thebeech argument.Greekφηγός (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 southernGreece.[30]

Distribution and habitat

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European beech (Fagus sylvatica)
Beeches inEhrenbach,Germany
North American beech,seen in autumn
Chinese beech (Fagus engleriana)

Britain and Ireland

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Fagus sylvaticawas a late entrant toGreat Britainafter 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.[31]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.[32]Large areas of theChilternsare covered with beech woods, which are habitat to thecommon bluebelland other flora. TheCwm Clydach National Nature Reservein 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.[33]

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 650 m (2,100 ft).[34]The tallest and longest hedge in the world (according toGuinness World Records) is theMeikleour Beech HedgeinMeikleour,Perth and Kinross,Scotland.

Continental Europe

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Fagus sylvaticais one of the most common hardwood trees in north-central Europe, in France constituting alone about 15% of all nonconifers. 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 naturalspruceboundary, 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 ofBergenon the west coast of Norway. Near the city ofLarvikis 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.[35]

Theprimeval beech forests of the Carpathiansare 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 theUNESCO World Heritage Listin 2007.[36]

North America

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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). There are the only extant (surviving)Fagusspecies in the Western Hemisphere. Before thePleistoceneIce 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. grandifoliatolerates 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

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

Ecology

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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 withleaf litter.

In North America, they can formbeech-mapleclimaxforests by partnering with thesugar maple.

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

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.[37]

Diseases

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Beech bark diseaseis a fungal infection that attacks the American beech through damage caused by scale insects.[38]Infection can lead to the death of the tree.[39]

Beech leaf diseaseis 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.[40] 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.[41]

Cultivation

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

Uses

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Beech Treephotographed byEugène Atget,circa1910–1915

Wood

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Beech wood is an excellentfirewood,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 forBudweiserbeer. This provides a complex surface on which the yeast can settle, so that it does not pile up, preventing yeastautolysiswhich would contribute off-flavours to the beer.[citation needed]Beech logs are burned to dry themaltused in Germansmoked beers.[42]Beech is also used to smokeWestphalian ham,[43]traditionalandouille(an offal sausage) from Normandy,[44]and some cheeses.

Somedrumsare made from beech, which has a tone between those ofmapleandbirch,the two most popular drum woods.

The textilemodalis a kind ofrayonoften made wholly from reconstitutedcelluloseof pulped beech wood.[45][46][47]

The European speciesFagus sylvaticayields 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.[citation needed]

Beech wood is used for the stocks of military rifles when traditionally preferred woods such aswalnutare scarce or unavailable or as a lower-cost alternative.[48]

Food

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The edible fruit of the beech tree,[3]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 statesmanPliny the Elderin his workNatural History,beechnut was eaten by the people ofChioswhen 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".[49]They can also be roasted and pulverized into an adequatecoffee substitute.[50]The leaves can be steeped in liquor to give a light green/yellow liqueur.

Books

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Painting on beech wood - 1511

In antiquity, the bark of the beech tree was used byIndo-European peoplefor writing-related purposes, especially in a religious context.[51]Beech wood tablets were a commonwriting materialin Germanic societies before the development ofpaper.The Old Englishbōc[52]has the primary sense of "beech" but also a secondary sense of "book", and it is frombōcthat the modern word derives.[53]In modern German, the word for "book" isBuch,withBuchemeaning "beech tree". In modern Dutch, the word for "book" isboek,withbeukmeaning "beech tree". In Swedish, these words are the same,bokmeaning both "beech tree" and "book". There is a similar relationship in some Slavic languages. InRussianandBulgarian,the word for beech isбук(buk), while that for "letter" (as in a letter of the Alpha bet) is буква (bukva), whileSerbo-CroatianandSloveneuse "bukva"to refer to the tree.

Other

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The pigmentbistrewas made from beech woodsoot.Beechlitterraking as a replacement for straw inanimal husbandrywas an old non-timber practice in forest management that once occurred in parts ofSwitzerlandin the 17th century.[54][55][56][57]Beech has been listed as one of the 38 plants whose flowers are used to prepareBach flower remedies.[58]

See also

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References

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