Editing Nothofagus

{{Short description|Genus of plants}}
{{Automatic taxobox
| name = ''Nothofagus''
| image = Nothofagus cunninghamii kz02.jpg
| image_caption = ''[[Nothofagus cunninghamii]]'',  Eastern Australia.
| fossil_range = <br/>[[Late Cretaceous]] to recent {{Fossil range|83.6|0}}
| parent_authority = [[Ludmila Andreevna Kuprianova|Kuprian.]]<ref name="APGIII-2009">{{Cite journal |last=Angiosperm Phylogeny Group |year=2009 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=105–121|doi=10.1111/j.1095-8339.2009.00996.x |doi-access=free |hdl=10654/18083 |hdl-access=free }}</ref>
| display_parents = 2
| taxon = Nothofagus
| authority = [[Carl Ludwig Blume|Blume]]
| synonyms = * ''Calucechinus'' <small>Hombr. & Jacquinot ex Decne. in J.S.C.Dumont d'Urville</small>
* ''Calusparassus'' <small>Hombr. & Jacquinot ex Decne. in J.S.C.Dumont d'Urville</small>
* ''Cliffortioides'' <small>Dryand. ex Hook.</small>
* ''Fagaster'' <small>Spach</small>
* ''Fuscospora'' <small>(R.S.Hill & J.Read) [[Peter Brian Heenan|Heenan]] & Smissen</small>
* ''Lophozonia'' <small>Turcz.</small>
* ''Myrtilloides'' <small>Banks & Sol. ex Hook.</small>
* ''Trisyngyne'' <small>Baill.</small>
| synonyms_ref = <ref name="POWO">{{cite web |title=''Nothofagus'' |url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:330346-2 |publisher=Plants of the World Online - Kew Science |access-date=19 April 2023}}</ref>
| subdivision_ranks = 
| subdivision = 
| range_map = Nothofagus Distribution.svg
| range_map_caption = The range of ''Nothofagus''.
}}

[[Image:Nothofagus obliqua Shoot LeavesCupules.jpg|thumb|Shoots, leaves, and [[calybium and cupule|cupules]] of ''N. obliqua'']]
[[Image:Beech trees southern North Island New Zealand.JPG|thumb|Southern beech trees in New Zealand]]
[[File:Nothofagus range including New Caledonia.jpg|thumb|The ''Nothofagus'' plant genus illustrates the distribution on fragments of the old supercontinent Gondwana: [[Australia]], [[New Guinea]], [[New Zealand]], [[New Caledonia]], [[Argentina]], and [[Chile]]. Fossils show that the genus originated on [[Gondwana]].]]

'''''Nothofagus''''', also known as the '''southern beeches''', is a [[genus]] of 43 [[species]] of [[tree]]s and [[shrub]]s native to the Southern Hemisphere, found across southern South America (Chile, Argentina) and east and southeast Australia, New Zealand, [[New Guinea]], and [[New Caledonia]].<ref name="Christenhusz-2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | doi-access = free |bibcode=2016Phytx.261..201C }}</ref> The species are ecological dominants in many temperate forests in these regions.<ref>{{Cite book|title=Ecology and Biogeography of Nothofagus Forests|last1=Veblen|first1=Thomas|last2=Hill|first2=Robert|last3=Read|first3=Jennifer|publisher=Yale University Press|year=1996|isbn=978-0-300-06423-0|location=New Haven, CT}}</ref> Some species are reportedly naturalised in Germany and Great Britain.<ref name="KewChecklist">[http://apps.kew.org/wcsp/namedetail.do?name_id=135736 Kew World Checklist of Selected Plant Families]</ref> The genus has a rich fossil record of leaves, [[Calybium and cupule|cupule]]s, and pollen, with fossils extending into the late Cretaceous period and occurring in Australia, New Zealand, Antarctica, and South America.<ref name="Hill-2001">{{Cite journal|title = Biogeography, evolution and palaeoecology of Nothofagus (Nothofagaceae): The contribution of the fossil record|last = Hill|first = Robert|authorlink1=Robert Hill (botanist)|date = 2001|journal = Australian Journal of Botany|doi = 10.1071/BT00026|volume=49|issue = 3|pages=321| bibcode=2001AuJB...49..321H }}</ref>

==Description==
The [[leaf|leaves]] are toothed or entire, [[evergreen]] or [[deciduous]]. The [[fruit]] is a small, flattened or triangular [[nut (fruit)|nut]], borne in cupules containing one to seven nuts.

==Reproduction==
Many individual trees are extremely old, and at one time, some populations were thought to be unable to reproduce in present-day conditions where they were growing, except by [[basal shoot|sucker]]ing ([[asexual reproduction|clonal reproduction]]), being remnant forest from a cooler time. [[Sexual reproduction]] has since been shown to be possible.<ref>{{cite web |url=http://cgi.cse.unsw.edu.au/~lambert/cgi-bin/clim/2005/07/ |title=Abstracts on Global Climate Change |website=cgi.cse.unsw.edu.au |url-status=dead |archive-url=https://web.archive.org/web/20080107011616/http://cgi.cse.unsw.edu.au/~lambert/cgi-bin/clim/2005/07/ |archive-date=2008-01-07}}</ref>

==Taxonomy==
The genus ''Nothofagus'' was first formally described in 1850 by [[Carl Ludwig Blume]] who published the description in his book ''Museum botanicum Lugduno-Batavum, sive, Stirpium exoticarum novarum vel minus cognitarum ex vivis aut siccis brevis expositio et descriptio''.<ref name="AusPlantCensus">{{cite web |title=''Nothofagus'' |url=https://biodiversity.org.au/nsl/services/apc-format/display/95806|website=Australian Plant Census |access-date=21 April 2020}}</ref><ref name="Blume-1850">{{cite book |last1=Blume |first1=Carl Ludwig |title=Museum botanicum Lugduno-Batavum, sive, Stirpium exoticarum novarum vel minus cognitarum ex vivis aut siccis brevis expositio et descriptio |date=1850 |pages=306–307 |url=https://www.biodiversitylibrary.org/item/200651#page/316/mode/1up |access-date=22 April 2020}}</ref> ''Nothofagus'' means "false beech", which Blume chose to indicate that ''Nothofagus'' species were different from beeches in the [[Northern Hemisphere]].<ref>{{Cite journal |last=Ryan |first=John Charles |date=2021-07-25 |title="Solitary in Your Rainy Kingdom:" Postcolonial Poetic Narratives of the Southern Beech |url=https://asle-brasil.com/journal/index.php/aslebr/article/view/158 |journal=Revista Interdisciplinar de Literatura e Ecocrítica |language=en |volume=7 |issue=1 |pages=5–28}}</ref>

In the past, they were included in the family [[Fagaceae]], but genetic tests revealed them to be genetically distinct,<ref name="Manos-1997"/> and they are now included in their own family, '''Nothofagaceae'''.<ref name="Manos-1997">{{Cite journal|title = Phylogenetic analyses of 'higher' Hamamelididae based on plastid sequence data|last = Manos|first = Paul|date = 1997|journal = American Journal of Botany|doi = 10.2307/2446139|pmid = 21708548|jstor=2446139|volume=84|issue = 10|pages=1407–1419|doi-access = free}}</ref>

===Species list===
The following is a list of species, hybrids and varieties accepted by the [[Plants of the World Online]] as of April 2023:<ref name="POWO"/>
*''[[Nothofagus aequilateralis]]'' <small>([[Marcel Gustav Baumann-Bodenheim|Baum.-Bod.]]) [[Steenis]]</small> (New Caledonia)
*''[[Nothofagus alessandrii]]'' <small>[[Marcial Ramón Espinosa Bustos|Espinosa]]</small> (Central Chile)
*''[[Nothofagus alpina]]'' <small>([[Poepp.]] & [[Endl.]]) [[Oerst.]]</small> (Argentina South, Chile Central, Chile South)
*''[[Nothofagus antarctica]]'' <small>([[G.Forst.]]) Oerst.</small> (Argentina South, Chile Central, Chile South)
*''[[Nothofagus balansae]]'' <small>([[Baill.]]) Steenis</small> (New Caledonia)
*''[[Nothofagus baumanniae]]'' <small>(Baum.-Bod.) Steenis</small> (New Caledonia)
*''[[Nothofagus betuloides]]'' <small>([[Mirb.]]) Oerst.</small> (Argentina South, Chile South)
*''[[Nothofagus brassii]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus carrii]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus cliffortioides]]'' <small>([[Hook.f.]]) Oerst.</small> (New Zealand North, New Zealand South)
*''[[Nothofagus codonandra]]'' <small>(Hook.f.) Oerst.</small> (New Caledonia)
*''[[Nothofagus crenata]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus cunninghamii]]'' <small>(Hook.f.) Oerst.</small> (Tasmania, Victoria)
*''[[Nothofagus discoidea]]'' <small>(Baum.-Bod.) Steenis</small> (New Caledonia)
*''[[Nothofagus dombeyi]]'' <small>(Mirb.) Oerst.</small> (Argentina South, Chile Central, Chile South)
*''[[Nothofagus flaviramea]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus fusca]]'' <small>(Hook.f.) Oerst.</small> (New Zealand North, New Zealand South)
*''[[Nothofagus glauca]]'' <small>([[Phil.]]) [[Johan Carl Krauss|Krasser]]</small> (Chile Central)
*''[[Nothofagus grandis]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus gunnii]]'' <small>(Hook.f.) Oerst.</small> (Tasmania)
*''[[Nothofagus macrocarpa]]'' <small>([[A.DC.]]) [[Francisco María Vázquez|F.M.Vázquez]] & [[Roberto A. Rodríguez|R.A.Rodr.]]</small> (Chile Central)
*''[[Nothofagus menziesii]]'' <small>(Hook.f.) Oerst.</small> (New Zealand North, New Zealand South)
*''[[Nothofagus moorei]]'' <small>([[F.Muell.]]) Krasser</small> (New South Wales, Queensland)
*''[[Nothofagus nitida]]'' <small>(Phil.) Krasser</small> (Chile South)
*''[[Nothofagus nuda]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus obliqua]]'' <small>(Mirb.) Oerst.</small> (Argentina South, Chile Central, Chile South)
*''[[Nothofagus perryi]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus pseudoresinosa]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus pullei]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus pumilio]]'' <small>(Poepp. & Endl.) Krasser</small> (Argentina South, Chile Central, Chile South)
*''[[Nothofagus resinosa]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus rubra]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus rutila]]'' <small>[[Pierfelice Ravenna|Ravenna]]</small> (Chile Central)
*''[[Nothofagus solandri]]'' <small>(Hook.f.) Oerst.</small> (New Zealand North, New Zealand South)
*''[[Nothofagus starkenborghiorum]]'' <small>Steenis</small> (Bismarck Archipelago, New Guinea)
*''[[Nothofagus stylosa]]'' <small>Steenis</small> (New Guinea)
*''[[Nothofagus truncata]]'' <small>([[William Colenso|Colenso]]) [[Leonard C. Cockayne|Cockayne]]</small> (New Zealand North, New Zealand South)
*''[[Nothofagus womersleyi]]'' <small>Steenis</small> (New Guinea)
*''Nothofagus × apiculata'' <small>(Colenso) Cockayne</small> (New Zealand North, New Zealand South)
*''Nothofagus × blairii'' <small>[[Thomas Kirk (botanist)|Kirk]]</small> (New Zealand North, New Zealand South)
*''Nothofagus × dodecaphleps'' <small>[[Michael Livingstone Grant|Mike L.Grant]] & [[E.J. Clement|E.J.Clement]]</small> (artificial hybrid)
*''Nothofagus × eugenananus'' <small>[[Ken Gillanders|Gilland.]]</small> (artificial hybrid)
*''[[Nothofagus × leoni]]'' <small>Espinosa</small> (Chile Central)
*''Nothofagus × solfusca'' <small>[[Harry Howard Barton Allan|Allan]]</small> (New Zealand North)

===Subgenera===
Four subgenera are recognized, based on morphology and DNA analysis:<ref name="Heenan-2013"/>
* Subgenus ''Fuscospora'', six species (''N. alessandri, N. cliffortioides, N. fusca, N. gunnii, N. solandri'', and ''N. truncata'') in New Zealand, Tasmania, and southern South America.
* Subgenus ''Lophozonia'', seven species (''N. alpina, N. cunninghamii, N. glauca, N. macrocarpa, N. menziesii, N. moorei'', and ''N. obliqua'') in New Zealand, Australia, and southern South America.
* Subgenus ''Nothofagus'', five species (''N. antarctica, N. betuloides, N. dombeyi, N. nitida'', and ''N. pumilio'') in southern South America. 
* Subgenus ''Brassospora'' (or ''Trisyngyne''), 20 accepted species (''N. aequilateralis, N. balansae, N. baumanniae, N. brassii, N. carrii, N. codonandra, N. crenata, N. discoidea,  N. flaviramea, N. grandis, N. nuda, N. perryi, N. pseudoresinosa, N, pullei, N. recurva, N. resinosa, N. rubra, N. starkenborghiorum, N. stylosa'', and ''N. womersleyi'') in New Guinea and New Caledonia.

In 2013, [[Peter Brian Heenan]] and [[Rob D. Smissen]] proposed splitting the genus into four, turning the four recognized subgenera into the new genera ''Fuscospora'', ''Lophozonia'' and ''Trisyngyne'', with the five South American species of subgenus ''Nothofagus'' remaining in genus ''Nothofagus''.<ref name="Heenan-2013">{{cite journal | last1 = Heenan | first1 = P.B. | last2 = Smissen | first2 = R.D. | year = 2013 | title = Revised circumscription of ''Nothofagus'' and recognition of the segregate genera ''Fuscospora'', ''Lophozonia'', and ''Trisyngyne'' (Nothofagaceae) | journal = Phytotaxa | volume = 146 | issue = 1| pages = 1–31 | doi = 10.11646/phytotaxa.146.1.1 | doi-access = free | bibcode = 2013Phytx.146....1H }}</ref> The proposed new genera are not accepted at the [[World Checklist of Selected Plant Families]].<ref name="KewChecklist" /><ref>{{cite journal | last1 = Hill | first1 = RS |authorlink1=Robert Hill (botanist)| last2 = Jordan | first2 = GJ | last3 = Macphail | first3 = MK | year = 2015 | title = Why we should retain ''Nothofagus sensu lato'' | journal = Australian Systematic Botany | volume = 28 | issue = 3| pages = 190–193 | doi=10.1071/sb15026| bibcode = 2015AuSyB..28..190H | s2cid = 83733526 }}</ref>

===Extinct species===
The following additional species are listed as extinct:<ref name="Hill-2001" /><ref>{{cite journal | last1 = Carpenter | first1 = RJ | last2 = Bannister | first2 = JM | last3 = Lee | first3 = DE | last4 = Jordan | first4 = GJ | year = 2014 | title = ''Nothofagus'' subgenus ''Brassospora'' (Nothofagaceae) leaf fossils from New Zealand: A link to Australia and New Guinea? | journal = Botanical Journal of the Linnean Society | volume = 174 | issue = 4| pages = 503–515 | doi=10.1111/boj.12143}}</ref><ref>{{cite journal | last1 = Jordan | first1 = GJ | year = 1999 | title = A new Early Pleistocene species of Nothofagus and the climatic implications of co-occurring Nothofagus fossils | url = https://eprints.utas.edu.au/1800/1/pachyphylla_paper_preprint.pdf| journal = Australian Systematic Botany | volume = 12 | issue = 6| pages = 757–765 | doi=10.1071/sb98025| bibcode = 1999AuSyB..12..757J }}</ref><ref>{{Cite web |title=Fossilworks: Nothofagus |url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=250689 |access-date=2022-12-04 |website=[[Paleobiology Database]]}}</ref>
*†''[[Nothofagus australis]]'' (Argentina, Early Oligocene-Early Miocene)
*†''[[Nothofagus balfourensis]]'' (Tasmania, Late Oligocene-Early Miocene)
*†''[[Nothofagus beardmorensis]]'' (Antarctica, Late Pliocene)<ref>{{cite journal| doi=10.1016/S0034-6667(96)00003-6 | volume=94 | issue=1–2 | title=Nothofagus beardmorensis (Nothofagaceae), a new species based on leaves from the Pliocene Sirius Group, Transantarctic Mountains, Antarctica | year=1996 | journal=Review of Palaeobotany and Palynology | pages=11–24 | last1 = Hill | first1 = R.S. |authorlink1=Robert Hill (botanist)| last2 = Harwood | first2 = D.M. | last3 = Webb | first3 = P.-N.| bibcode=1996RPaPa..94...11H }}</ref>
*†''[[Nothofagus bulbosa]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus cethanica]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus cooksoniae]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus crenulata]]'' (Argentina, Mid Oligocene-Early Miocene)
*†''[[Nothofagus cretacea]]'' (Antarctica, Late Cretaceous)
*†''[[Nothofagus densinervosa]]'' (Argentina, Mid Oligocene-Early Miocene)
*†''[[Nothofagus elongata]]'' (Argentina, Early Oligocene-Early Miocene)
*†''[[Nothofagus glandularis]]'' (Tasmania, Mid Oligocene-Early Miocene)
*†''[[Nothofagus glaucifolia]]'' (Antarctica, Late Cretaceous)
*†''[[Nothofagus lanceolata]]'' (Argentina, Late Oligocene-Early Miocene)
*†''[[Nothofagus lobata]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus magelhaenica]]'' (Argentina, Early Oligocene-Early Miocene)
*†''[[Nothofagus magellanica]]'' (Argentina, Late Oligocene-Mid Miocene)
*†''[[Nothofagus maideni]]'' (Tasmania, Early Oligocene-Mid Miocene)
*†''[[Nothofagus microphylla]]'' (Tasmania, Late Oligocene-Mid Miocene)
*†''[[Nothofagus mucronata]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus muelleri]]'' (New South Wales, Late Eocene)
*†''[[Nothofagus novae-zealandiae]]'' (New Zealand, Mid-Late Miocene)
*†''[[Nothofagus pachyphylla]]'' (Tasmania, Early Pleistocene)
*†''[[Nothofagus palustris]]'' (New Zealand, Late Oligocene-Early Miocene)
*†''[[Nothofagus peduncularis]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus robusta]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus serrata]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus serrulata]]'' (Argentina, Mid Oligocene-Early Miocene)
*†''[[Nothofagus simplicidens]]'' (Argentina, Mid Oligocene-Early Miocene)
*†''[[Nothofagus smithtonensis]]'' (Tasmania, Early Oligocene)
*†''[[Nothofagus tasmanica]]'' (Tasmania, Eocene-Early Oligocene)
*†''[[Nothofagus ulmifolia]]'' (Antarctica, Late Cretaceous)
*†''[[Nothofagus variabilis]]'' (Argentina, Oligocene)
*†''[[Nothofagus zastawniakiae]]'' (Antarctica, Late Cretaceous)

==Distribution==
The pattern of distribution around the southern [[Pacific Rim]] suggests the dissemination of the genus dates to the time when Antarctica, Australia, and South America were connected in a common land-mass or [[supercontinent]] referred to as [[Gondwana]].<ref>{{Cite web |url=http://www.nativeforest.org/campaigns/gondwana/index.htm |title=Native Forest Network (2003) ''Gondwana Forest Sanctuary'' |access-date=2007-11-06 |archive-url=https://web.archive.org/web/20080516052545/http://www.nativeforest.org/campaigns/gondwana/index.htm |archive-date=2008-05-16 |url-status=dead }}</ref> More recent studies suggest that the [[Antarctic land bridge]] likely played a major role in the dispersal of the genus between these continents.<ref name=":1">{{Cite journal |last1=van den Ende |first1=Conrad |last2=White |first2=Lloyd T. |last3=van Welzen |first3=Peter C. |date=2017-04-01 |title=The existence and break-up of the Antarctic land bridge as indicated by both amphi-Pacific distributions and tectonics |url=https://linkinghub.elsevier.com/retrieve/pii/S1342937X16302829 |journal=Gondwana Research |volume=44 |pages=219–227 |bibcode=2017GondR..44..219V |doi=10.1016/j.gr.2016.12.006 |issn=1342-937X}}</ref> However, genetic evidence using [[Molecular clock|molecular dating]] methods has been used to argue that the species in New Zealand and New Caledonia evolved from species that arrived in these landmasses by dispersal across oceans.<ref>{{cite journal | last1 = Knapp | first1 = M | last2 = Stockler | first2 = K | last3 = Havell | first3 = D | last4 = Delsuc | first4 = F | last5 = Sebastiani | first5 = F | last6 = Lockhart | first6 = PJ | year = 2005 | title = Relaxed molecular clock provides evidence for long-distance dispersal of ''Nothofagus'' (Southern Beech) | journal = PLOS Biology | volume = 3 | issue = 1| pages = 38–43 | doi = 10.1371/journal.pbio.0030014 | pmid=15660155 | pmc=539330 | doi-access = free }}</ref> Uncertainty exists in molecular dates and controversy rages as to whether the distribution of ''Nothofagus'' derives from the break-up of Gondwana (i.e. [[vicariance]]), or if long-distance dispersal has occurred across oceans. In South America, the northern limit of the genus can be construed as [[La Campana National Park]] and the [[Vizcachas Mountains]] in the central part of Chile.<ref>C. Michael Hogan (2008) [http://globaltwitcher.auderis.se/artspec_information.asp?thingid=82831 ''Chilean Wine Palm: Jubaea chilensis'', GlobalTwitcher.com, ed. Nicklas Stromberg] {{webarchive|url=https://web.archive.org/web/20121017013207/http://globaltwitcher.auderis.se/artspec_information.asp?thingid=82831 |date=2012-10-17 }}
</ref>

== Evolutionary history ==
''Nothofagus'' first appeared in Antarctica during the early [[Campanian]] stage (83.6 to 72.1 million years ago) of the [[Late Cretaceous]]. During the Campanian ''Nothofagus'' diversified and became dominant within Antarctic ecosystems, with the appearance of all four modern subgenera by the end of the stage. ''Nothofagus'' shows a progressive decline in the Antarctic pollen record through the [[Maastrichtian]], before substantially recovering after the [[Cretaceous–Paleogene boundary|Cretaceous-Paleogene boundary]].<ref>{{Citation|last=Cantrill|first=David J.|title=Cretaceous to Paleogene Vegetation Transition in Antarctica|date=2018|url=https://linkinghub.elsevier.com/retrieve/pii/B9780128130124000279|work=Transformative Paleobotany|pages=645–659|publisher=Elsevier|language=en|doi=10.1016/b978-0-12-813012-4.00027-9|isbn=978-0-12-813012-4|access-date=2021-05-19|url-access=subscription}}</ref> ''Nothofagus'' persisted in Antarctica deep into the Cenozoic, despite the increasingly inhospitable conditions, with the final records from the late [[Neogene]], around 15-5 million years old, which were small tundra-adapted [[prostrate shrub]]s, similar to ''[[Salix arctica]]'' (Arctic willow).<ref>{{Cite journal|last1=Rees-Owen|first1=Rhian L.|last2=Newton|first2=Robert J.|last3=Ivanovic|first3=Ruza F.|last4=Francis|first4=Jane E.|last5=Riding|first5=James B.|last6=Marca|first6=Alina D.|date=February 2021|title=A calibration of cellulose isotopes in modern prostrate Nothofagus and its application to fossil material from Antarctica|journal=Science of the Total Environment|language=en|volume=754|pages=142247|doi=10.1016/j.scitotenv.2020.142247|pmid=33254952|bibcode=2021ScTEn.75442247R|doi-access=free}}</ref>

''Nothofagus'' first appeared in southern South America during the late Campanian. During the Paleocene and Eocene they were mostly restricted to southern Patagonia, before reaching a peak abundance during the Miocene. Their distribution contracted westwards during the late Miocene due to the aridification of Patagonia.<ref>{{Cite journal|last1=Pujana|first1=Roberto R|last2=Fernández|first2=Damián A|last3=Panti|first3=Carolina|last4=Caviglia|first4=Nicolás|date=2020-12-31|title=The micro- and megafossil record of Nothofagaceae from South America|url=http://dx.doi.org/10.1093/botlinnean/boaa097|journal=Botanical Journal of the Linnean Society|volume=196|issue=1|pages=1–20|doi=10.1093/botlinnean/boaa097|issn=0024-4074|url-access=subscription}}</ref>

Although the genus now mostly occurs in cool, isolated, high-altitude environments at [[temperate]] and [[tropical]] [[latitude]]s, the fossil record shows that it survived in climates that appear to be much warmer than those that ''Nothofagus'' now occupies.<ref>{{cite journal | last1 = Carpenter | first1 = RJ | last2 = Jordan | first2 = GJ | last3 = Macphail | first3 = MK | last4 = Hill | first4 = RS |author-link4=Robert Hill (botanist)| year = 2012 | title = Near-tropical early eocene terrestrial temperatures at the Australo-Antarctic margin, western Tasmania | journal = Geology | volume = 40 | issue = 3| pages = 267–270 | doi = 10.1130/G32584.1 | bibcode = 2012Geo....40..267C }}</ref>

==Ecology==
''Nothofagus'' species are used as food plants by the [[larva]]e of [[Hepialidae|hepialid]] [[moth]]s of the genus ''[[Aenetus]]'', including ''A. eximia'' and ''A. virescens''. ''[[Zelopsis nothofagi]]'' is a leaf hopper, endemic to New Zealand, which is found on ''Nothofagus''.

''[[Cyttaria]]'' is genus of [[ascomycete]] [[fungi]] found on or associated with ''Nothofagus'' in Australia and South America. ''[[Misodendrum]]'' are specialist parasitic plants found on various species of ''Nothofagus'' in South America.<ref name="Read-1996" /> Additionally, the beetle, [[Brachysternus prasinus]], has been known to live in ''Nothofagus'' in Chile and in parts of Argentina. The geographic range of B. prasinus is highly dependent on the availability and distribution of Nothofagus on which B. prasinus is believed to feed. B. prasinus have been observed in the Nothofagus forests near the cities of Coquimbo and Llanquihue in Chile as well as the areas of [[Neuquén Province|Neuquén]] and Chubut in Western Argentina.''<ref>{{Cite journal |last1=Jameson |first1=Mary Liz |last2=Smith |first2=Andrew B. T. |date=September 1, 2002 |title=Revision of the South American Genus BrachysternusGuérin-Méneville (Coleoptera: Scarabaeidae: Rutelinae: Anoplognathini: Brachysternina) |url=https://bioone.org/journals/the-coleopterists-bulletin/volume-56/issue-3/0010-065X_2002_056_0321_ROTSAG_2.0.CO_2/Revision-of-the-South-American-Genus-BrachysternusGu%c3%a9rin-M%c3%a9neville-Coleoptera/10.1649/0010-065X(2002)056[0321:ROTSAG]2.0.CO;2.full |journal=The Coleopterists Bulletin |volume=56 |issue=3 |pages=321–366 |doi=10.1649/0010-065X(2002)056[0321:ROTSAG]2.0.CO;2 |issn=0010-065X|hdl=10057/3386 |hdl-access=free }}</ref>

The species of subgenus ''Brassospora'' are evergreen, and distributed in the tropics of New Guinea, New Britain, and New Caledonia. In New Guinea and New Britain ''Nothofagus'' is characteristic of lower montane rain forests between 1000 and 2500 meters elevation, occurring infrequently at elevations as low as 600 meters, and in upper montane forests between 2500 and 3150 meters elevation. ''Nothofagus'' is most commonly found above the ''[[Castanopsis]]-[[Lithocarpus]]'' zone in the lower montane forests, and below the [[conifer]]-dominated upper montane forests. ''Nothofagus'' grows in mixed stands with trees of other species or in pure stands, particularly on ridge crests and upper slopes. The [[New Guinea Highlands|Central Range]] has the greatest diversity of species, with fewer species distributed among the mountains of western and northern New Guinea, New Britain, and Goodenough and Normanby islands.<ref name="Read-1996">{{cite book |last1=Read |first1=Jennifer |first2=Geoffrey S. |last2=Hope |year=1996 |chapter=Ecology of Nothofagus forests of New Guinea and New Caledonia |chapter-url={{GBurl|SKSBF-0ormAC|p=200}} |title=The Ecology and Biogeography of Nothofagus Forests |editor1-last=Veblen |editor1-first=Thomas T |editor2-first=Robert S. |editor2-last=Hill |editor-link2=Robert Hill (botanist)|editor3-first=Jennifer |editor3-last=Read |publisher=Yale University Press |isbn=978-0-300-06423-0 |pages=200–256}}</ref>

The New Caledonian species are endemic to the main island (Grand Terre), most commonly on soils derived from [[ultramafic rock]]s between 150 and 1350 meters elevation. They occur in isolated stands, forming a low or stunted and irregular and fairly open canopy. The conifers ''[[Agathis]]'' and ''[[Araucaria]]'' are sometimes present as emergents, rising 10 to 20 meters above the ''Nothofagus'' canopy.<ref name="Read-1996"/>

==Beech mast==
Every four to six years or so, ''Nothofagus'' produces a heavier crop of seeds and is known as the beech [[Mast (botany)|mast]]. In New Zealand, the beech mast causes an increase in the population of introduced mammals such as mice, rats, and [[stoat]]s. When the rodent population collapses, the stoats begin to prey on native bird species, many of which are threatened with extinction.<ref>{{cite web|url=http://www.doc.govt.nz/conservation/native-plants/beech-forest/|title=Beech forest: Native plants|publisher=Department of Conservation|access-date=26 August 2012}}</ref>  This phenomenon is covered in more detail in the article on [[stoats in New Zealand]].

==References==
{{Reflist|2}}

==External links==
{{Commons category|Nothofagus}}
{{Wikispecies|Nothofagus}}
*{{cite web |title=''Nothofagus'' Blume |publisher=Atlas of Living Australia |url=https://bie.ala.org.au/species/https://id.biodiversity.org.au/node/apni/7911560}}

{{Angiosperm families}}
{{Taxonbar|from1=Q218274|from2=Q13634230}}
{{Authority control}}

[[Category:Nothofagaceae]]
[[Category:Trees of New Zealand]]
[[Category:Fagales genera]]
[[Category:Extant Campanian first appearances]]
[[Category:Taxa named by Carl Ludwig Blume]]