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Nothofagus

{{Short description|Genus of plants}}

{{Automatic taxobox

| name = Nothofagus

| image = Nothofagus cunninghamii kz02.jpg

| image_caption = Nothofagus cunninghamii, Eastern Australia.

| fossil_range =
Late Cretaceous to recent {{Fossil range|83.6|0}}

| parent_authority = Kuprian.{{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 }}

| display_parents = 2

| taxon = Nothofagus

| authority = Blume

| synonyms = * Calucechinus Hombr. & Jacquinot ex Decne. in J.S.C.Dumont d'Urville

  • Calusparassus Hombr. & Jacquinot ex Decne. in J.S.C.Dumont d'Urville
  • Cliffortioides Dryand. ex Hook.
  • Fagaster Spach
  • Fuscospora (R.S.Hill & J.Read) Heenan & Smissen
  • Lophozonia Turcz.
  • Myrtilloides Banks & Sol. ex Hook.
  • Trisyngyne Baill.

| synonyms_ref = {{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}}

| subdivision_ranks =

| subdivision =

| range_map = Nothofagus Distribution.svg

| range_map_caption = The range of Nothofagus.

}}

Image:Nothofagus obliqua Shoot LeavesCupules.jpg of N. obliqua]]

Image:Beech trees southern North Island New Zealand.JPG

File:Nothofagus range including New Caledonia.jpg, 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 trees and shrubs native to the Southern Hemisphere, found across southern South America (Chile, Argentina) and east and southeast Australia, New Zealand, New Guinea, and New Caledonia.{{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 }} The species are ecological dominants in many temperate forests in these regions.{{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}} Some species are reportedly naturalised in Germany and Great Britain.[http://apps.kew.org/wcsp/namedetail.do?name_id=135736 Kew World Checklist of Selected Plant Families] The genus has a rich fossil record of leaves, cupules, and pollen, with fossils extending into the late Cretaceous period and occurring in Australia, New Zealand, Antarctica, and South America.{{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 }}

Description

The leaves are toothed or entire, evergreen or deciduous. The fruit is a small, flattened or triangular 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 suckering (clonal reproduction), being remnant forest from a cooler time. Sexual reproduction has since been shown to be possible.{{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}}

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.{{cite web |title=Nothofagus |url=https://biodiversity.org.au/nsl/services/apc-format/display/95806|website=Australian Plant Census |access-date=21 April 2020}}{{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}} Nothofagus means "false beech", which Blume chose to indicate that Nothofagus species were different from beeches in the Northern Hemisphere.{{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}}

In the past, they were included in the family Fagaceae, but genetic tests revealed them to be genetically distinct, and they are now included in their own family, Nothofagaceae.{{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}}

=Species list=

The following is a list of species, hybrids and varieties accepted by the Plants of the World Online as of April 2023:

=Subgenera=

Four subgenera are recognized, based on morphology and DNA analysis:

  • 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.{{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 }} The proposed new genera are not accepted at the World Checklist of Selected Plant Families.{{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 }}

=Extinct species=

The following additional species are listed as extinct:{{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}}{{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 }}{{Cite web |title=Fossilworks: Nothofagus |url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=250689 |access-date=2022-12-04 |website=Paleobiology Database}}

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.{{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 }} More recent studies suggest that the Antarctic land bridge likely played a major role in the dispersal of the genus between these continents.{{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}} However, genetic evidence using 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.{{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 }} 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.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 }}

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.{{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}} 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 shrubs, similar to Salix arctica (Arctic willow).{{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}}

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.{{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}}

Although the genus now mostly occurs in cool, isolated, high-altitude environments at temperate and tropical latitudes, the fossil record shows that it survived in climates that appear to be much warmer than those that Nothofagus now occupies.{{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 }}

Ecology

Nothofagus species are used as food plants by the larvae of hepialid moths 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. 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 and Chubut in Western Argentina.''{{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 }}

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 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.{{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}}

The New Caledonian species are endemic to the main island (Grand Terre), most commonly on soils derived from ultramafic rocks 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.

Beech mast

Every four to six years or so, Nothofagus produces a heavier crop of seeds and is known as the beech mast. In New Zealand, the beech mast causes an increase in the population of introduced mammals such as mice, rats, and stoats. When the rodent population collapses, the stoats begin to prey on native bird species, many of which are threatened with extinction.{{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}} 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