beech

{{Short description|Genus of flowering plants in the family Fagaceae}}

{{About|the genus of trees|other uses|Beech (disambiguation)|and|Beechwood (disambiguation)}}

{{Distinguish|Beach|Birch|Fagus (disambiguation)}}

{{Use dmy dates|date=December 2024}}

{{Automatic taxobox

| name = Beech

| image = Fagus sylvatica Purpurea JPG4a.jpg

| image_caption = European beech (Fagus sylvatica)

| parent_authority = K.Koch

| taxon = Fagus

| authority = L.

| type_species = Fagus sylvatica

| type_species_authority = L.

| subdivision_ranks = Species

| subdivision = See text

}}

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 {{Small|Denk & G.W.Grimm}} and Fagus.{{Cite journal |last1=Denk |first1=Thomas |last2=Grimm |first2=Guido W. |last3=Cardoni |first3=Simone |last4=Csilléry |first4=Katalin |last5=Kurz |first5=Mirjam |last6=Schulze |first6=Ernst-Detlef |last7=Simeone |first7=Marco Cosimo |last8=Worth |first8=James R. P. |date=2024 |title=A subgeneric classification of Fagus (Fagaceae) and revised taxonomy of western Eurasian beeches |journal=Willdenowia |volume=54 |issue=2–3 |doi=10.3372/wi.54.54301 |issn=0511-9618|doi-access=free }} 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

File:Fagus sylvatica leaf 001.jpg]]

File:Beechnuts during autumn.jpg

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 {{convert|5|–|15|cm|in|0|abbr=off}} long and {{convert|4|–|10|cm|in|0|abbr=on}} broad.

The bark is smooth and light gray. The fruit is a small, sharply three-angled nut {{convert|10|–|15|mm|in|frac=8|abbr=on}} long, borne singly or in pairs in soft-spined husks {{convert|1.5|–|2.5|cm|in|frac=8|abbr=on}} 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.{{cite thesis |last=Shen |first=Chung-Fu |title=A Monograph of the Genus Fagus Tourn. Ex L. (Fagaceae) |date=1992 |type=PhD |publisher=City University of New York |oclc=28329966}} The nuts are called beechnuts{{Cite book |last=Lyle |first=Katie Letcher |title=The Complete Guide to Edible Wild Plants, Mushrooms, Fruits, and Nuts: How to Find, Identify, and Cook Them |publisher=FalconGuides |year=2010 |isbn=978-1-59921-887-8 |edition=2nd |location=Guilford, CN |pages=138 |oclc=560560606 |orig-year=2004}} 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 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).{{Citation |last1=Cardoni |first1=Simone |title=5S-IGS rDNA in wind-pollinated trees (Fagus L.) encapsulates 55 million years of reticulate evolution and hybrid origins of modern species |date=2021-10-19 |url=https://www.biorxiv.org/content/10.1101/2021.02.26.433057v2 |access-date=2024-10-24 |language=en |doi=10.1111/tpj.15601|biorxiv=10.1101/2021.02.26.433057 |last2=Piredda |first2=Roberta |last3=Denk |first3=Thomas |last4=Grimm |first4=Guido W. |last5=Papageorgiou |first5=Aristotelis C. |last6=Schulze |first6=Ernst-Detlef |last7=Scoppola |first7=Anna |last8=Shanjani |first8=Parvin Salehi |last9=Suyama |first9=Yoshihisa|journal=The Plant Journal |volume=109 |issue=4 |pages=909–926 |pmid=34808015 |pmc=9299691 }}{{Cite journal |last1=Denk |first1=Thomas |last2=Grimm |first2=Guido W. |last3=Hemleben |first3=Vera |date=June 2005 |title=Patterns of molecular and morphological differentiation in Fagus (Fagaceae): phylogenetic implications |url=https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.92.6.1006 |journal=American Journal of Botany |language=en |volume=92 |issue=6 |pages=1006–1016 |doi=10.3732/ajb.92.6.1006 |pmid=21652485 |issn=0002-9122}} 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. 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,{{Cite journal |last=Denk |first=T. |date=2003-09-01 |title=Phylogeny of Fagus L. (Fagaceae) based on morphological data |url=https://link.springer.com/article/10.1007/s00606-003-0018-x |journal=Plant Systematics and Evolution |language=en |volume=240 |issue=1 |pages=55–81 |doi=10.1007/s00606-003-0018-x |bibcode=2003PSyEv.240...55D |issn=1615-6110|url-access=subscription }} or as a variety of F. japonica.{{Cite journal |last1=Oh |first1=Sang-Hun |last2=Youm |first2=Jung-Won |last3=Kim |first3=Yong-In |last4=Kim |first4=Young-Dong |date=2016-09-01 |title=Phylogeny and Evolution of Endemic Species on Ulleungdo Island, Korea: The Case of Fagus multinervis (Fagaceae) |url=http://openurl.ingenta.com/content/xref?genre=article&issn=0363-6445&volume=41&issue=3&spage=617 |journal=Systematic Botany |language=en |volume=41 |issue=3 |pages=617–625 |doi=10.1600/036364416X692271|url-access=subscription }} The differ from their siblings by their unique nuclear and plastid genotypes.{{Cite journal |last1=Jiang |first1=Lu |last2=Bao |first2=Qin |last3=He |first3=Wei |last4=Fan |first4=Deng-Mei |last5=Cheng |first5=Shan-Mei |last6=López-Pujol |first6=Jordi |last7=Chung |first7=Myong Gi |last8=Sakaguchi |first8=Shota |last9=Sánchez-González |first9=Arturo |last10=Gedik |first10=Aysun |last11=Li |first11=De-Zhu |last12=Kou |first12=Yi-Xuan |last13=Zhang |first13=Zhi-Yong |date=July 2022 |title=Phylogeny and biogeography of Fagus (Fagaceae) based on 28 nuclear single/low-copy loci |url=https://onlinelibrary.wiley.com/doi/10.1111/jse.12695 |journal=Journal of Systematics and Evolution |language=en |volume=60 |issue=4 |pages=759–772 |doi=10.1111/jse.12695 |issn=1674-4918|url-access=subscription }}

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 (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. 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{{Cite journal |last1=Grímsson |first1=Friðgeir |last2=Grimm |first2=Guido W. |last3=Zetter |first3=Reinhard |last4=Denk |first4=Thomas |date=2016-12-01 |title=Cretaceous and Paleogene Fagaceae from North America and Greenland: evidence for a Late Cretaceous split between Fagus and the remaining Fagaceae |url=https://acpa.botany.pl/Cretaceous-and-Paleogene-Fagaceae-nfrom-North-America-and-Greenland-evidence-nfor,118915,0,2.html |journal=Acta Palaeobotanica |volume=56 |issue=2 |pages=247–305 |doi=10.1515/acpa-2016-0016 |issn=2082-0259|doi-access=free }}) and quickly radiated across the high latitudes of the Northern Hemisphere, with a first diversity peak in the Miocene of northeastern Asia.{{Cite journal |last1=Denk |first1=Thomas |last2=Grimm |first2=Guido W. |date=2009 |title=The biogeographic history of beech trees |url=https://linkinghub.elsevier.com/retrieve/pii/S0034666709001353 |journal=Review of Palaeobotany and Palynology |language=en |volume=158 |issue=1–2 |pages=83–100 |doi=10.1016/j.revpalbo.2009.08.007|bibcode=2009RPaPa.158...83D |url-access=subscription }}{{Cite journal |last1=Renner |first1=S. S. |last2=Grimm |first2=Guido W. |last3=Kapli |first3=Paschalia |last4=Denk |first4=Thomas |date=2016-07-19 |title=Species relationships and divergence times in beeches: new insights from the inclusion of 53 young and old fossils in a birth–death clock model |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |language=en |volume=371 |issue=1699 |pages=20150135 |doi=10.1098/rstb.2015.0135 |issn=0962-8436 |pmc=4920336 |pmid=27325832}} The contemporary species are the product of past, repeated reticulate evolutionary processes (outbreeding, introgression, 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 diverging lineage in the evolution of the Fagaceae family,{{Cite journal |last1=Zhou |first1=Biao-Feng |last2=Yuan |first2=Shuai |last3=Crowl |first3=Andrew A. |last4=Liang |first4=Yi-Ye |last5=Shi |first5=Yong |last6=Chen |first6=Xue-Yan |last7=An |first7=Qing-Qing |last8=Kang |first8=Ming |last9=Manos |first9=Paul S. |last10=Wang |first10=Baosheng |date=2022-03-14 |title=Phylogenomic analyses highlight innovation and introgression in the continental radiations of Fagaceae across the Northern Hemisphere |journal=Nature Communications |language=en |volume=13 |issue=1 |pages=1320 |doi=10.1038/s41467-022-28917-1 |issn=2041-1723 |pmc=8921187 |pmid=35288565|bibcode=2022NatCo..13.1320Z }} which also includes oaks and chestnuts.{{cite journal |last1=Manos |first1=Paul S. |first2=Kelly P. |last2=Steele |title=Phylogenetic analysis of "Higher" Hamamelididae based on Plasid Sequence Data |journal=American Journal of Botany |volume=84 |issue=10 |pages=1407–19 |date=1997 |doi=10.2307/2446139 |jstor=2446139 |pmid=21708548 |url=|doi-access=free }} 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 (POWO):

class="wikitable collapsible"
ImageName

!Subgenus

!Status, systematic affinity

Distribution

!Accepted as species in POWO as of April 2023{{cite web |date=2022-05-07 |title=Fagus L. - Plants of the World Online |url=http://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30048723-2 |access-date=2023-04-24 |website=Plants of the World Online}}

|Fagus caspica {{Small|Denk & G.W.Grimm}} – Caspian beech

|Fagus

|New species described in 2024; first-diverging lineage within the Western Eurasian group

|Talysch and Elburz Mountains, southeastern Azerbaijan and northern Iran

|Populations included in F. sylvatica subsp. orientalis

Fagus chienii {{small|W.C.Cheng}}

|Fagus

|Possibly conspecific with F. lucida

Probably extinct, described from a single location in China (Sichuan). Individuals recently collected at the type locality were morphologically and genetically indistinguishable from F. pashanica.{{Cite journal |last1=Li |first1=Dan-Qi |last2=Jiang |first2=Lu |last3=Liang |first3=Hua |last4=Zhu |first4=Da-Hai |last5=Fan |first5=Deng-Mei |last6=Kou |first6=Yi-Xuan |last7=Yang |first7=Yi |last8=Zhang |first8=Zhi-Yong |date=2023-09-01 |title=Resolving a nearly 90-year-old enigma: The rare Fagus chienii is conspecific with F. hayatae based on molecular and morphological evidence |journal=Plant Diversity |volume=45 |issue=5 |pages=544–551 |doi=10.1016/j.pld.2023.01.003 |issn=2468-2659 |pmc=10625896 |pmid=37936819|bibcode=2023PlDiv..45..544L }}

|Yes

120pxFagus crenata {{small|Blume}} – 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

Japan; in the mountains of Kyushu, Shikoku and Honshu, down to sea-level in southern Hokkaido.

|Yes

120pxFagus engleriana {{small|Seemen ex Diels}} – Chinese beech

|Englerianae

|Widespread species; continental sister species of F. japonica

China; south of the Yellow River

|Yes

120pxFagus grandifolia {{small|Ehrh.}} – American beech

|Fagus

|Widespread species; sister species of F. mexicana

Eastern North America; from E. Texas and N. Florida, United States, to the St. Lawrence River, Canada at low to mid altitudes

|Yes, including Mexican beeches, F. mexicana

120pxFagus hayatae {{small|Palib. ex Hayata}}

|Fagus

|Narrow endemic species; forming a cryptic sister species pair with F. pashanica

Taiwan; restricted to the mountains of northern Taiwan

|Yes

|Fagus hohenackeriana {{Small|Palib.}} – Hohenacker's or Caucasian beech

|Fagus

|Dominant tree species of the Pontic and Caucasus Mountains; intermediate between F. caspica and F. orientalis.{{Cite journal |last1=Gömöry |first1=Dušan |last2=Paule |first2=Ladislav |date=2010-07-01 |title=Reticulate evolution patterns in western-Eurasian beeches |url=https://link.springer.com/article/10.1007/s00035-010-0068-y |journal=Botanica Helvetica |language=en |volume=120 |issue=1 |pages=63–74 |doi=10.1007/s00035-010-0068-y |bibcode=2010BotHe.120...63G |issn=1420-9063|url-access=subscription }}{{Cite journal |last1=Gömöry |first1=Dušan |last2=Paule |first2=Ladislav |last3=Mačejovský |first3=Vladimír |date=2018-06-29 |title=Phylogeny of beech in western Eurasia as inferred by approximate Bayesian computation |url=https://pbsociety.org.pl/journals/index.php/asbp/article/view/asbp.3582 |journal=Acta Societatis Botanicorum Poloniae |language=en |volume=87 |issue=2 |page=3582 |doi=10.5586/asbp.3582 |bibcode=2018AcSBP..87.3582G |issn=2083-9480}}{{Cite journal |last1=Kurz |first1=Mirjam |last2=Kölz |first2=Adrian |last3=Gorges |first3=Jonas |last4=Pablo Carmona |first4=Beatriz |last5=Brang |first5=Peter |last6=Vitasse |first6=Yann |last7=Kohler |first7=Martin |last8=Rezzonico |first8=Fabio |last9=Smits |first9=Theo H. M. |last10=Bauhus |first10=Jürgen |last11=Rudow |first11=Andreas |last12=Kim Hansen |first12=Ole |last13=Vatanparast |first13=Mohammad |last14=Sevik |first14=Hakan |last15=Zhelev |first15=Petar |date=2023-03-01 |title=Tracing the origin of Oriental beech stands across Western Europe and reporting hybridization with European beech – Implications for assisted gene flow |url=https://linkinghub.elsevier.com/retrieve/pii/S0378112723000348 |journal=Forest Ecology and Management |volume=531 |pages=120801 |doi=10.1016/j.foreco.2023.120801 |bibcode=2023ForEM.53120801K |issn=0378-1127|hdl=20.500.11850/597076 |hdl-access=free }} Its genetic heterogeneity{{Cite journal |last1=Sękiewicz |first1=Katarzyna |last2=Danelia |first2=Irina |last3=Farzaliyev |first3=Vahid |last4=Gholizadeh |first4=Hamid |last5=Iszkuło |first5=Grzegorz |last6=Naqinezhad |first6=Alireza |last7=Ramezani |first7=Elias |last8=Thomas |first8=Peter A. |last9=Tomaszewski |first9=Dominik |last10=Walas |first10=Łukasz |last11=Dering |first11=Monika |date=2022 |title=Past climatic refugia and landscape resistance explain spatial genetic structure in Oriental beech in the South Caucasus |journal=Ecology and Evolution |language=en |volume=12 |issue=9 |pages=e9320 |doi=10.1002/ece3.9320 |issn=2045-7758 |pmc=9490144 |pmid=36188519|bibcode=2022EcoEv..12E9320S }} 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)

|No, populations included in F. sylvatica subsp. orientalis

120pxFagus japonica {{small|Maxim.}}

|Englerianae

|Widespread species; insular sister species of F. engleriana

Japan; Kyushu, Shikoku and Honshu from sea-level up to c. 1500 m a.s.l.

|Yes

|Fagus longipetiolata {{Small|Seemen}}

|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.

|China, south of the Yellow River, into N. Vietnam; in montane areas up to 2400 m a.s.l.{{Cite web |title=Fagus longipetiolata in Flora of China @ efloras.org |url=http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=200006255 |access-date=2024-10-24 |website=www.efloras.org}}

|Replaced by F. sinensis

120pxFagus lucida {{small|Rehder & E.H.Wilson}}

|Fagus

|Rare species; closest relatives are F. crenata and F. longipetiolata

China; south of the Yellow River in montane areas between 800 and 2000 m a.s.l.{{Cite web |title=Fagus lucida in Flora of China @ efloras.org |url=http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=200006256 |access-date=2024-10-24 |website=www.efloras.org}}

|Yes

|Fagus mexicana {{Small|Martínez}}

|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)

|Hidalgo, Mexico; at 1400–2000 m a.s.l. as an element of the subtropical montane mesophilic forest"([https://www.gob.mx/conabio/prensa/bosques-mesofilos-de-montana-de-mexico?idiom=es bosque mesófilo de montaña]) superimposing the tropical lowland rainforests.

|No, populations included in F. grandifolia

Fagus multinervis {{small|Nakai}}

|Englerianae

|Narrow endemic species, first diverging lineage within subgenus Englerianae

South Korea (Ulleungdo)

|Yes

120pxFagus orientalis {{small|Lipsky}} – Oriental beech (in a narrow sense)

|Fagus

|Sister species of F. sylvatica

Southeastern Europe (SE Bulgaria, NE Greece, European Turkey) and adjacent northwestern Asia (NW and N Anatolia)

|No, treated as subspecies of F. sylvatica

Fagus pashanica {{small|C.C.Yang}}

|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{{Cite web |title=Fagus hayatae in Flora of China @ efloras.org |url=http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=200006254 |access-date=2024-10-24 |website=www.efloras.org}})

|Yes

Fagus sinensis {{small|Oliv.}}

|Fagus

|Invalid; the original material included material from two much different species: F. engleriana and F. longipetiolata

China (Hubei), Vietnam

|Yes, erroneously used as older synonym of F. longipetiolata

120pxFagus sylvatica {{small|L.}} – European beech

|Fagus

|Sister species of and closely related to F. orientalis

Europe

|Yes

=Natural and potential hybrids=

class="wikitable collapsible"
ImageNameParentage

!Status

Distribution
|Fagus (×) moesiaca {{Small|(K. Malý) Czeczott}}

|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.{{Cite journal |last1=Ulaszewski |first1=Bartosz |last2=Meger |first2=Joanna |last3=Mishra |first3=Bagdevi |last4=Thines |first4=Marco |last5=Burczyk |first5=Jarosław |date=2021 |title=Complete Chloroplast Genomes of Fagus sylvatica L. Reveal Sequence Conservation in the Inverted Repeat and the Presence of Allelic Variation in NUPTs |journal=Genes |language=en |volume=12 |issue=9 |pages=1357 |doi=10.3390/genes12091357 |doi-access=free |issn=2073-4425 |pmc=8468245 |pmid=34573338}} They may represent a lowland ecotype of F. sylvatica.{{Cite journal |last=Denk |first=Th. |date=January 1999 |title=The taxonomy of Fagus in western Eurasia. 2: Fagus sylvatica subsp. sylvatica |url=https://onlinelibrary.wiley.com/doi/10.1002/fedr.19991100510 |journal=Feddes Repertorium |language=en |volume=110 |issue=5–6 |pages=381–412 |doi=10.1002/fedr.19991100510 |issn=0014-8962|url-access=subscription }}

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

|Southeastern Balkans

|Fagus okamotoi {{Small|Shen}}

|F. crenata × F. japonica ?

|Unique phenotype, described from an area in which F. crenata and F. japonica are 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

120pxFagus × taurica {{small|Popl.}} – Crimean beechF. 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)

Crimean peninsula

=Phylogeny=

A cladogram of 11 beech species is shown below.{{cite journal |author1=Jiang, Lu |display-authors=etal |title=Phylogeny and biogeography of Fagus (Fagaceae) based on 28 nuclear single/low-copy loci |journal=Journal of Systematics and Evolution |date=10 October 2020 |volume=60 |issue=4 |pages=759–772 |doi=10.1111/jse.12695 |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/jse.12695|url-access=subscription }}

{{clade

|label1=Fagus

|1={{clade

|label1=subg. Engleriana

|1={{clade

|1=F. multinervis

|2={{clade

|1=F. engleriana

|2=F. japonica

}}

}}

|label2=subg. Fagus

|2={{clade

|label1=sect. Grandifolia

|1={{clade

|1=F. grandifolia

|2=F. mexicana

}}

|2={{clade

|label1=sect. Fagus

|1={{clade

|1=F. orientalis

|2=F. sylvatica

}}

|label2=sect. Lucida

|2={{clade

|1=F. hayatae

|2={{clade

|1=F. crenata

|2={{clade

|1=F. longipetiolata

|2=F. lucida

}}

}}

}}

}}

}}

}}

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=Fossil species=

Numerous species have been named globally from the fossil record spanning from the Cretaceous to the Pleistocene.{{cite web |url=http://ifpni.org/genus.htm?id=3EE9EA96-C4DC-4621-B6EB-697227CF797B |website=The International Fossil Plant Names Index |title=Fagus |accessdate=6 Feb 2023}}

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Fossil species formerly placed in Fagus include:

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= Etymology =

The name of the tree in Latin, fagus (from whence the generic epithet), is cognate with English "beech" and of Indo-European origin, and played an important 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.Robert Beekes, Etymological Dictionary of Greek, Leiden and Boston 2010, pp. 1565–6

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.{{cite web |url=http://linnaeus.nrm.se/flora/di/faga/fagus/fagusylv.jpg |title= Map|website=linnaeus.nrm.se |format=JPG|access-date=2019-08-07}} 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.{{cite web |url=http://www.forestry.gov.uk/newsrele.nsf/WebPressReleases/1A301105A92950FE80257012002508A0 |title=International Foresters Study Lake District's greener, friendlier forests |publisher=Forestry Commission |access-date=4 August 2010 |archive-date=28 January 2010 |archive-url=https://web.archive.org/web/20100128215333/http://www.forestry.gov.uk/newsrele.nsf/WebPressReleases/1A301105A92950FE80257012002508A0 |url-status=dead }} 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.{{cite web |url=http://www.ccw.gov.uk/landscape--wildlife/protecting-our-landscape/special-landscapes--sites/protected-landscapes/national-nature-reserves/cwm-clydach.aspx |title=Cwm Clydach |publisher=Countryside Council for Wales Landscape & wildlife |access-date=4 August 2010 |url-status = dead|archive-url=https://web.archive.org/web/20100925090500/http://www.ccw.gov.uk/landscape--wildlife/protecting-our-landscape/special-landscapes--sites/protected-landscapes/national-nature-reserves/cwm-clydach.aspx |archive-date=25 September 2010 }}

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 {{convert|650|m|ft|-2|abbr=on}}.{{cite book |first1=C.D. |last1=Preston |first2=D. |last2=Pearman |first3=T.D. |last3=Dines |title=New Atlas of the British Flora |publisher=Oxford University Press |location= |date=2002 |isbn=978-0-19-851067-3 |pages= |url=}} The tallest and longest hedge in the world (according to Guinness World Records) is the 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. 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.{{cite journal |last1=Bradshaw |first1=R.H.W. |first2=N. |last2=Kito and |first3=T. |last3=Giesecke |title=Factors influencing the Holocene history of Fagus |journal=Forest Ecology and Management |volume=259 |issue=11 |pages=2204–12 |date=2010 |doi=10.1016/j.foreco.2009.11.035 |bibcode=2010ForEM.259.2204B |url=}}

The 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 UNESCO World Heritage List in 2007.{{cite web |url = https://whc.unesco.org/en/list/1133 |title = Ancient and Primeval Beech Forests of the Carpathians and Other Regions of Europe |website = UNESCO World Heritage Centre |publisher = United Nations Educational, Scientific, and Cultural Organization |access-date = 13 November 2021}}

= 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 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.{{cite book |last1=Lawrence |first1=Gale |first2=Adelaide |last2=Tyrol |title=A Field Guide to the Familiar: Learning to Observe the Natural World |publisher=Prentice-Hall |location= |date=1984 |isbn=978-0-13-314071-2 |pages=75–76 |url=}}

= Diseases =

Beech bark disease is a fungal infection that attacks the American beech through damage caused by scale insects."beech." The Columbia Encyclopedia. New York: Columbia University Press, 2008. Credo Reference. Web. 17 September 2012. Infection can lead to the death of the tree.{{cite book |first= |last= |chapter=beech bark disease |chapter-url= |editor= |title=Dictionary of Microbiology & Molecular Biology |publisher=Wiley |location= |date=2006 |isbn=978-0-470-03545-0 |pages= |url=}} Credo Reference. Web. 27 September 2012.

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.{{cite news |last=Crowley |first=Brendan |date=2020-09-28 |title=Deadly 'Beech Leaf Disease' Identified Across Connecticut and Rhode Island |work=The Connecticut Examiner |url=https://ctexaminer.com/2020/09/28/deadly-beech-leaf-disease-identified-across-connecticut-and-rhode-island/ |access-date=2020-11-15}}

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.University of New Hampshire

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 which would contribute off-flavours to the beer.{{Citation needed|date= March 2018}} Beech logs are burned to dry the malt used in German smoked beers.{{cite web |url=https://www.schlenkerla.de/rauchbier/prozess/prozess.html |title=Der Brauprozeß von Schlenkerla Rauchbier |author= |date=2011 |website=Schlenkerla - die historische Rauchbierbrauerei |publisher=Schlenkerla |language=de |access-date=11 December 2020}} Beech is also used to smoke Westphalian ham,{{cite web |url=http://www.germanfoods.org/consumer/facts/guidetoham.cfm |title=GermanFoods.org - Guide to German Sausages and German Hams |access-date=2012-05-17 |url-status = dead|archive-url=https://web.archive.org/web/20121123232931/http://www.germanfoods.org/consumer/facts/guidetoham.cfm |archive-date=2012-11-23 }} traditional andouille (an offal sausage) from Normandy,{{Cite web |url=http://www.cookthink.com/reference/823/What_is_andouille |title=What is andouille? | Cookthink |access-date=2012-11-22 |archive-url=https://web.archive.org/web/20120512015109/http://www.cookthink.com/reference/823/What_is_andouille |archive-date=2012-05-12 |url-status=dead }} 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 is a kind of rayon often made wholly from reconstituted cellulose of pulped beech wood.holistic-interior-designs.com, [http://www.holistic-interior-designs.com/modal-fabric.html Modal Fabric] {{Webarchive|url=https://web.archive.org/web/20111009010337/http://www.holistic-interior-designs.com/modal-fabric.html |date=2011-10-09 }}, retrieved 9 October 2011uniformreuse.co.uk, [http://www.uniformreuse.co.uk/fabric_modal.html?KeepThis=true&TB_iframe=true&height=460&width=800 Modal data sheet] {{Webarchive|url=https://web.archive.org/web/20111024015844/http://www.uniformreuse.co.uk/fabric_modal.html?KeepThis=true&TB_iframe=true&height=460&width=800 |date=2011-10-24 }}, retrieved 9 October 2011fabricstockexchange.com, [http://www.fabricstockexchange.com/blog/resources/fiber-dictionary/ Modal] {{Webarchive|url=https://web.archive.org/web/20110925102655/http://fabricstockexchange.com/blog/resources/fiber-dictionary/ |date=2011-09-25 }} (dictionary entry), retrieved 9 October 2011

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.{{cite journal | last1=Skarvelis | first1=Michalis | last2=Mantanis | first2=George I. | title=Physical and mechanical properties of beech wood harvested in the Greek public forests | journal=Wood Research | publisher=Pulp and Paper Research Institute | volume=58 | issue=1 | date=2012-12-29 | issn=1336-4561 | url=https://www.researchgate.net/publication/237840835 | access-date=2024-12-24 | pages=123–130}}

Beech wood is used for the stocks of military rifles when traditionally preferred woods such as walnut are scarce or unavailable or as a lower-cost alternative.{{cite book |first=J. |last=Walter |title=Rifles of the World |publisher=Krause Publications |edition=3rd |date=2006 |isbn=978-0-89689-241-5 |pages= |url=}}

=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, 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".{{Cite web|url=https://interestingearth.com/how_did_beech_mast_save_the_people_of_chios.html|title=How did beech mast save the people of Chios? - Interesting Earth|website=interestingearth.com|access-date=2019-10-07}} They can also be roasted and pulverized into an adequate coffee substitute.{{Cite book |url=https://www.worldcat.org/oclc/277203364 |title=The Complete Guide to Edible Wild Plants |publisher=Skyhorse Publishing |author=United States Department of the Army |year=2009 |isbn=978-1-60239-692-0 |location=New York |pages=29 |language=en-US |oclc=277203364}} The leaves can be steeped in liquor to give a light green/yellow liqueur.

=Books=

File:Venus with a Mirror (study).jpg

In antiquity, the bark of the beech tree was used by Indo-European people for writing-related purposes, especially in a religious context.{{cite book |first= Saskia |last=Pronk-Tiethoff |title= The Germanic loanwords in Proto-Slavic |url= https://books.google.com/books?id=0iWLAgAAQBAJ&pg=PA81 |date= 25 October 2013 |publisher= Rodopi |isbn= 978-94-012-0984-7 |pages= 81}} Beech wood tablets were a common writing material in Germanic societies before the development of paper. The Old English bōcA Concise Anglo-Saxon Dictionary, Second Edition (1916), [http://lexicon.ff.cuni.cz/html/oe_clarkhall/b0047.html Blōtan-Boldwela], John Richard Clark Hall has the primary sense of "beech" but also a secondary sense of "book", and it is from bōc that the modern word derives.{{cite web|url= http://www.etymonline.com/index.php?term=book |title= Book |author= Douglas Harper |work= Online Etymological Dictionary |access-date= 2011-11-18}} 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, the word for beech is бук (buk), while that for "letter" (as in a letter of the alphabet) is буква (bukva), while Serbo-Croatian and Slovene use "bukva" to refer to the tree.

=Other=

The pigment bistre was made from beech wood soot. Beech 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.{{cite journal |title= Three objectives of historical ecology: the case of litter collecting in Central European forests |author= Bürgi, M. |author2=Gimmi, U. |year= 2007 |doi= 10.1007/s10980-007-9128-0 |journal= Landscape Ecology |volume= 22 |issue= S1 |pages= 77–87|bibcode= 2007LaEco..22S..77B |hdl= 20.500.11850/58945 |s2cid= 21130814 |url= http://doc.rero.ch/record/316425/files/10980_2007_Article_9128.pdf |hdl-access= free }}{{cite journal |title= Soil carbon pools in Swiss forests show legacy effects from historic forest litter raking |author= Gimmi, U. |author2= Poulter, B. |author3= Wolf, A. |author4= Portner, H. |author5= Weber, P. |author6= Bürgi, M. |year= 2013 |doi= 10.1007/s10980-012-9778-4 |journal= Landscape Ecology |volume= 28 |issue= 5 |pages= 385–846|bibcode= 2013LaEco..28..835G |hdl= 20.500.11850/66782 |s2cid= 16930894 |url= http://doc.rero.ch/record/315772/files/10980_2012_Article_9778.pdf |hdl-access= free }}{{cite journal |title= Reconstructing European forest management from 1600 to 2010 |author= McGrath, M.J. |year= 2015 |doi= 10.5194/bg-12-4291-2015 |journal= Biogeosciences |volume= 12 |issue= 14 |pages= 4291–4316|display-authors=etal|bibcode= 2015BGeo...12.4291M |doi-access= free }}{{cite journal |title= Consequence of litter removal on pedogenesis: A case study in Bachs and Irchel (Switzerland) |author= Scalenghe, R. |author2= Minoja, A.P. |author3= Zimmermann, S. |author4= Bertini, S. |year= 2016 |doi= 10.1016/j.geoderma.2016.02.024 |journal= Geoderma |volume= 271 |pages= 191–201|bibcode= 2016Geode.271..191S |url= https://zenodo.org/record/889561 }} Beech has been listed as one of the 38 plants whose flowers are used to prepare Bach flower remedies.{{cite book|author=D. S. Vohra|title=Bach Flower Remedies: A Comprehensive Study|url=https://books.google.com/books?id=icG8onA0ys8C&pg=PR3|access-date=2 September 2013|date=1 June 2004|publisher=B. Jain Publishers|isbn=978-81-7021-271-3|page=3}}

See also

References

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