Beech

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Template:Short description Template:Good article Script error: No such module "about". Script error: No such module "Distinguish". Template:Use British English Template:Use dmy dates Template:Automatic taxobox 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 Denk & G.W.GrimmScript error: No such module "Check for unknown parameters". and Fagus.[1] 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.

The European beech Fagus sylvatica is the most commonly cultivated species, with several ornamental varieties, and forest trees yielding a timber used for furniture, flooring and construction, 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 some beers. 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. The fruit is a three-angled nut, with two in a spiny dehiscent cupule. The bark is smooth. The leaves have a central vein with side-veins parallel to each other and ending in a tooth on the thin leaf-blade. The tree is deciduous, dropping its leaves in autumn.[2]

Evolution

Evolutionary history

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File:Fagus sylvatica pliocenica MHNT.PAL.VEG.2002.31 (cropped).jpg
Fagus sylvatica pliocenica, Piacenzian, 3.6 to 2.6 mya

Numerous species have been named globally from the fossil record spanning from the Cretaceous to the Pleistocene.[3] Some fossil species formerly placed in Fagus have been moved to other genera, namely Alnus, Castanea, Fagopsis, Fagoxylon, Fagus-pollenites, Juglans, Nothofagaphyllites, Nothofagus, and Trigonobalanus.[3]

Fagus is the first diverging lineage in the evolution of the Fagaceae family,[4][5] which includes oaks and chestnuts.[6] 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.[4]

The first representatives of the modern-day genus were likely already present in the Paleocene of Arctic North America (western Greenland[4]) and quickly radiated across the high latitudes of the Northern Hemisphere, with a first diversity peak in the Miocene of northeastern Asia.[7][8] The contemporary species are the product of past, repeated reticulate evolutionary processes (outbreeding, introgression, hybridization).[9] As far as studied, heterozygosity and intragenomic variation are common in beech species,[9][10][11] and their chloroplast genomes are nonspecific with the exception of the Western Eurasian and North American species.[1]

Phylogeny

A cladogram of 11 extant beech species is shown below. The subgenera Engleriana and Fagus diverged from each other in the Early Oligocene era, 32.1 to 33.4 million years ago.[12]

Template:Clade

Taxonomy

The most recent classification system of the genus recognizes 14 species in two distinct subgenera, subgenus Englerianae and Fagus.[1] Beech species can be diagnosed by phenotypical and/or 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 nucleome (nuclear DNA), especially in noncoding, highly variable gene regions such as the spacers of the nuclear-encoded ribosomal RNA genes (ribosomal DNA).[9][10] 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. japonica and F. okamotoi, and one Chinese species, F. engleriana.[13] 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.[1] The beeches of Ulleungdo have been traditionally treated as a subspecies of F. engleriana, to which they are phenotypically identical,[13][14] or as a variety of F. japonica.[15] The differ from their siblings by their unique nuclear and plastid genotypes.[1][11][9]

The better known subgenus Fagus beeches are high-branching with tall, stout trunks and smooth silver-grey bark. This group includes five extant species in continental and insular East Asia (F. crenata, F. longipetiolata, F. lucida, and the cryptic sister species F. hayatae and F. pashanica), two pseudo-cryptic species in eastern North America (F. grandifolia, F. mexicana), and a species complex of at least four species (F. caspica, F. hohenackeriana, F. orientalis, 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.[1] The western Eurasian species are characterised by morphological and genetical gradients.[1]

Species

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

Image Name Subgenus Status, systematic affinity Distribution Accepted in POWO, Sept. 2025[16]
Fagus caspica Denk & G.W.GrimmScript error: No such module "Check for unknown parameters". – Caspian beech Fagus New species described in 2024;[1] first-diverging lineage within the Western Eurasian group Talysch and Elburz Mountains, southeastern Azerbaijan and northern Iran No mention
Fagus chienii W.C.ChengScript error: No such module "Check for unknown parameters". Fagus Possibly conspecific with F. lucida[14] Probably extinct, described from a single location in China (Sichuan). Individuals collected there were morphologically and genetically indistinguishable from F. pashanica.[17] Yes
File:Fagus crenata in Ogasayama 2010-10-17.jpg Fagus crenata BlumeScript error: No such module "Check for unknown parameters". – 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 subgenus Fagus[9] Japan; in the mountains of Kyushu, Shikoku and Honshu, down to sea-level in southern Hokkaido. Yes
File:Fagus engleriana - Morris Arboretum - DSC00475.JPG Fagus engleriana Seemen ex DielsScript error: No such module "Check for unknown parameters". – Chinese beech Englerianae Widespread species; continental sister species of F. japonica[10][11][9] China; south of the Yellow River Yes
File:Fagus grandifolia JPG1Ms.jpg Fagus grandifolia Ehrh.Script error: No such module "Check for unknown parameters". – American beech Fagus Widespread species; sister species of F. mexicana[11][9] Eastern North America; from E. Texas and N. Florida, United States, to the St. Lawrence River, Canada at low to mid altitudes Yes
File:Fagus hayatae 98412.jpg Fagus hayatae Palib. ex HayataScript error: No such module "Check for unknown parameters". Fagus Narrow endemic species; forming a cryptic sister species pair with F. pashanica[9][1] Taiwan; restricted to the mountains of northern Taiwan Yes
File:Fagus hohenackeriana near Fioletovo village, Armenia S-N 04.jpg Fagus hohenackeriana Palib.Script error: No such module "Check for unknown parameters". – Caucasian or Hohenacker's beech Fagus Dominant tree species of the Pontic and Caucasus Mountains; intermediate between F. caspica and F. orientalis.[18][19][20] Its genetic heterogeneity[1][21] may be indicative for ongoing speciation processes. Northeastern Anatolia (Pontic Mountains, Kaçkar Mountains) and Caucasus region (Lesser and Greater Caucasus, Georgia, Armenia, Ciscaucasia; down to sea-level in southwestern Georgia) Yes
File:Forest in Tanzawa 08.jpg Fagus japonica Maxim.Script error: No such module "Check for unknown parameters".File:Fagus mexicana, Zacualtipán de Ángeles, Hidalgo, Mexico 5737290.jpg Englerianae Widespread species; insular sister species of F. engleriana[9][10][11] Japan; Kyushu, Shikoku and Honshu from sea-level up to c. 1500 m a.s.l. Yes
Fagus longipetiolata SeemenScript error: No such module "Check for unknown parameters". Fagus Sym- to parapatric with F. lucida and F. pashanica, and sharing alleles with both species in addition to alleles indicating a sister relationship with the Japanese F. crenata.[9][11] China, south of the Yellow River, into N. Vietnam; in montane areas up to 2400 m a.s.l.[22] Replaced by F. sinensis
File:Fagus lucida Buk 2020-07-18 01.jpg Fagus lucida Rehder & E.H.WilsonScript error: No such module "Check for unknown parameters". Fagus Rare species; closest relatives are F. crenata[9][10][14] and F. longipetiolata[9][11] China; south of the Yellow River in montane areas between 800 and 2000 m a.s.l.[23] Yes
File:Fagus mexicana, Zacualtipán de Ángeles, Hidalgo, Mexico 5737290.jpg Fagus mexicana MartínezScript error: No such module "Check for unknown parameters". Fagus Narrow endemic sister species of F. grandifolia. F. mexicana differs from F. grandifolia by its slender leaves and less-evolved but more polymorphic set of alleles (higher level of heterozygosity)[9][11] 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. Yes
Fagus multinervis NakaiScript error: No such module "Check for unknown parameters". Englerianae Narrow endemic species, first diverging lineage within subgenus Englerianae[9][11] South Korea (Ulleungdo) Yes
File:TR Yedigöller asv2021-10 img13.jpg Fagus orientalis LipskyScript error: No such module "Check for unknown parameters". – Oriental beech (in a narrow sense) Fagus Sister species of F. sylvatica[19][20] Southeastern Europe (SE Bulgaria, NE Greece, East Thrace (European Turkey) and adjacent northwestern Asia (NW and N Anatolia) Yes
Fagus pashanica C.C.YangScript error: No such module "Check for unknown parameters". Fagus Continental sister species of F. hayatae, with a set of alleles that puts it closer to F. longipetiolata and F. crenata than its insular sister. China (Hubei, Hunan, Shaanxi, Sichuan, Zhejiang), at 1300–2300 m a.s.l.(eFlora of China, as F. hayatae[24]) Yes
Fagus sinensis Oliv.Script error: No such module "Check for unknown parameters". Fagus Invalid; the original material included material from two much different species: F. engleriana and F. longipetiolata[1][14] China (Hubei), Vietnam Yes, erroneously used as older synonym of F. longipetiolata
File:Fagus sylvatica TK 2023-05-06 5.jpg Fagus sylvatica L.Script error: No such module "Check for unknown parameters". – European beech Fagus Sister species of and closely related to F. orientalis[19][20] Europe Yes

Natural and potential hybrids

Name Parentage Status Distribution
Fagus (×) moesiaca (K. Malý) CzeczottScript error: No such module "Check for unknown parameters". F. sylvatica × F. orientalis No evidence so far for hybrid origin. All individuals addressed as F. moesiaca included in genetic studies fell within the variation of F. sylvatica.[10][25] They may represent a lowland ecotype of F. sylvatica.[1][26] Southeastern Balkans
Fagus × taurica Popl.Script error: No such module "Check for unknown parameters". – Crimean beech F. sylvatica × F. orientalis s.l. Hybrid status not yet tested by genetic data; according to isoenzyme profiles a less-evolved, relict population of F. sylvatica or intermediate between F. sylvatica and the species complex historically addressed as Oriental beech (F. orientalis in a broad sense)[18] Crimean peninsula

Etymology

The name of the tree in Latin, fagus (whence the generic epithet), is cognate with English "beech" and of Indo-European origin. It played a role in early debates on the geographical origins of the Indo-European people, the beech argument. 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.[27]

The common name of "beech" is from the Anglo-Saxon boc, bece or beoce, the German buche, the Swedish box - all meaning "book" as well as beech and derived from the Sanskrit boko or letter and bokos or writings. This connection to "beech" seems to have derived from the fact that the old Runic tablets were of beech wood.[28]

Ecology

Habitat and distribution

Beech requires a deep soil with good drainage and a neutral or slightly acidic soil, pH 6 to 7.5. It is vulnerable to drought as its root system is relatively shallow. It does not live in waterlogged areas, but it can grow in windy places, shade from other trees, and cold. In northern Europe it is a lowland species, while further south it is montane, growing at an altitude of up to Template:Convert.[29]

The English Lowlands beech forests is an ecoregion of high-canopy forest dominated by European beech in southeastern England, surviving as remnants such as the Template:Convert New Forest.[30] The species arrived in Britain after the last glaciation, and may have been restricted to basic soils in the south of England. It could have been introduced by Neolithic tribes who planted the trees for their edible nuts.[31] In southeast Wales, the Cwm Clydach National Nature Reserve holds beech woodlands on the western edge of their natural range in a steep limestone gorge.[32] The primeval beech forests of the Carpathians have been dominated since the last ice age by the beech.[33] In North America, beech can form Beech-maple forest, seen by some ecologists as a climax community, by partnering with the sugar maple.[34]

Pests and diseases

The beech blight aphid, Grylloprociphilus imbricator, is a common pest of American beech trees.[35]

Beech bark disease is a fungal infection of trees in the Eastern US, Canada, and Europe. Following damage caused by the scale insects Xylococculus betulae and Cryptococcus fagisuga, the fungi Neonectria faginata and Neonectria ditissima produce cankers each year; these may eventually girdle and kill the tree.[36]

Beech leaf disease is a disease that affects beeches spread by the nematode Litylenchus crenatae mccannii. The disease was discovered in Ohio in 2012.[37] It has spread through the Eastern United States and Canada.[38]

Uses

Furniture and construction

The European beech Fagus sylvatica yields a tough timber. It weighs about 720 kg per cubic metre and is widely used for furniture construction, flooring, plywood, and household items. The timber can be used to build chalets, houses, and log cabins.[39]

Ornamental tree

The European beech, Fagus sylvatica, is widely cultivated in most regions that have a suitable climate, including North and South America, Europe, South Africa, Australia, and New Zealand.[40] Many varieties are in cultivation, including the weeping beech F. sylvatica 'Pendula', several varieties of copper or purple beech, the fern-leaved beech F. sylvatica 'Asplenifolia',[41] and the tricolour beech F. sylvatica 'Roseomarginata'.[42] The columnar Dawyck beech (F. sylvatica 'Dawyck') is named after Dawyck Botanic Garden in the Scottish Borders.[43]

The tallest beech hedge in the world, and the longest hedge in Britain, is the Meikleour Beech Hedge in Perth and Kinross, Scotland.[44]

Food and food preparation

The fruit of the beech tree is an edible nut, known as beech mast.[45][46] According to the Roman statesman Pliny the Elder in his Natural History, the fruit: "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".[47] They can also be roasted and pulverized into a coffee substitute.[48]

Slats of beech wood, its flavour removed with caustic soda, are spread inside fermentation tanks for beers such as Budweiser to prevent the yeast from piling up on the bottom and possibly spoiling the beer's flavour.[49] Beech logs are burned to dry the malt used in German smoked beers.[50] Beech is also used to smoke Westphalian ham,[51] traditional andouille (an offal sausage) from Normandy,[52] and some smoked cheeses.[53]

Other

File:Venus with a Mirror (study) (cropped).jpg
Painting Venus with a Mirror. Oil and plaster on beech wood. Titian, 1511

In antiquity, Indo-European people used beech bark as a writing material, especially in a religious context.[54] Beech wood tablets were a common writing material in Germanic societies before the development of paper. The Old English bōc[55] has the primary sense of "beech" with a secondary sense of "book".[56]

The pigment bistre was made from beech wood soot. Beech litter raking was used as a replacement for straw in animal husbandry in Switzerland in the 17th century.[57][58][59][60] Beech is one of the 38 plants whose flowers are used to prepare Bach flower remedies.[61] Beech makes an excellent firewood.[62] Some drum shells are made from beech.[63] The textile modal is a kind of rayon often made wholly from cellulose extracted from pulped beech wood.[64][65][66]

In Gallo-Roman religion, Fagus (Latin for "beech") was a god known from four inscriptions found in the Hautes-Pyrénées.[67]

See also

References

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External links

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  64. ↑ holistic-interior-designs.com, Modal Fabric Template:Webarchive, retrieved 9 October 2011
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  67. ↑ Nicole Jufer & Thierry Luginbühl (2001). Les dieux gaulois : répertoire des noms de divinités celtiques connus par l'épigraphie, les textes antiques et la toponymie. Paris: Editions Errance. Template:ISBN.
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