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[[File:Gunnera insignis.jpg|thumb|right|200px|''Gunnera insignis'', [[Costa Rica]]]] | [[File:Gunnera insignis.jpg|thumb|right|200px|''Gunnera insignis'', [[Costa Rica]]]] | ||
'''''Gunnera''''' is the sole genus of [[herbaceous]] [[flowering plant]]s in the family '''Gunneraceae''', which contains 63 species. Some species in this genus, namely those in the subgenus ''Panke'', have extremely large leaves. Species in the genus are variously native to [[Latin America]], [[Australia]], [[New Zealand]], [[Papuasia]], [[Hawaii]], [[insular Southeast Asia]], [[Africa]], and [[Madagascar]].<ref name=l>[http://apps.kew.org/wcsp/namedetail.do?name_id=370359 Kew World Checklist of Selected Plant Families]</ref> The stalks of some species are edible.<ref name="TasteAtlas" /> | '''''Gunnera''''' is the sole genus of [[herbaceous]] [[flowering plant]]s in the family '''Gunneraceae''', which contains 63 species. Some species in this genus, namely those in the subgenus ''Panke'', have extremely large leaves. Species in the genus are variously native to [[Latin America]], [[Australia]], [[New Zealand]], [[Papuasia]], [[Hawaii]], [[insular Southeast Asia]], eastern [[Africa]], and [[Madagascar]].<ref name=l>[http://apps.kew.org/wcsp/namedetail.do?name_id=370359 Kew World Checklist of Selected Plant Families]</ref> The stalks of some species are edible.<ref name="TasteAtlas" /> | ||
== Taxonomy == | == Taxonomy == | ||
''Gunnera'' is the only genus in the family Gunneraceae.<ref name="Christenhusz-Byng2016">{{cite journal | vauthors = Christenhusz MJ, Byng JW |year=2016 |title=The number of known plants species in the world and its annual increase |url=http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |journal=Phytotaxa |volume=261 |issue=3 |pages=201–217 |doi=10.11646/phytotaxa.261.3.1 |doi-access=free}}</ref> The [[APG II system]] | ''Gunnera'' is the only genus in the family Gunneraceae.<ref name="Christenhusz-Byng2016">{{cite journal | vauthors = Christenhusz MJ, Byng JW |year=2016 |title=The number of known plants species in the world and its annual increase |url=http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |journal=Phytotaxa |volume=261 |issue=3 |pages=201–217 |doi=10.11646/phytotaxa.261.3.1 |doi-access=free}}</ref> The [[APG II system]] of 2003 accepted this family and assigned it to the order Gunnerales in the clade [[core eudicots]]. The family then consisted of one or two genera, ''Gunnera'' and, optionally, ''[[Myrothamnus]]'', the latter optionally segregated as a separate family, [[Myrothamnaceae]]. This represents a change from the [[APG system]] of 1998, which accepted two separate families, unplaced as to order. The [[APG III system]] and [[APG IV system]] accept the family Gunneraceae, and places ''Myrothamnus'' in Myrothamnaceae; both families are placed in the order Gunnerales in the core eudicots.<ref name="APGIV2016">{{Cite journal |author =Angiosperm Phylogeny Group |year=2016 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |journal=[[Botanical Journal of the Linnean Society]] |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385 |issn=0024-4074 |doi-access=free |author-link=Angiosperm Phylogeny Group}}</ref> | ||
The genus ''Gunnera'' was named after the Norwegian botanist [[Johann Ernst Gunnerus]]. At first it was assigned to the family [[Haloragaceae]], though that presented difficulties that led to the general recognition of the family Gunneraceae, as had been proposed about the beginning of the 20th century. In the meantime, in many publications it had been referred to as being in the Haloragaceae, variously misspelt (as for example "Halorrhagidaceae".<ref name="WattPP">{{cite book | vauthors = Watt JM, Breyer-Brandwijk MG, Gerdina M | title = The Medicinal and Poisonous Plants of Southern and Eastern Africa | publisher = E & S Livingstone | date = 1962 | edition = Second | chapter = Halorrhagidaceae | page = 500 }}</ref>) Such references still cause difficulties in consulting earlier works. However, currently ''Gunnera'' is firmly assigned to the [[Monotypic taxon|monogeneric]] family Gunneraceae.<ref name="WWOK">{{cite journal | vauthors = Wanntorp L, Wanntorp HE, Oxelman B, Källersjö M | title = Phylogeny of ''Gunnera'' | journal = Plant Systematics and Evolution | date = March 2001 | volume = 226 | issue = 1 | pages = 85–107 | doi = 10.1007/s006060170075 | jstor = 23644101 | bibcode = 2001PSyEv.226...85W | s2cid = 42201778 }}</ref> | The genus ''Gunnera'' was named after the Norwegian botanist [[Johann Ernst Gunnerus]]. At first it was assigned to the family [[Haloragaceae]], though that presented difficulties that led to the general recognition of the family Gunneraceae, as had been proposed about the beginning of the 20th century. In the meantime, in many publications it had been referred to as being in the Haloragaceae, variously misspelt (as for example "Halorrhagidaceae".<ref name="WattPP">{{cite book | vauthors = Watt JM, Breyer-Brandwijk MG, Gerdina M | title = The Medicinal and Poisonous Plants of Southern and Eastern Africa | publisher = E & S Livingstone | date = 1962 | edition = Second | chapter = Halorrhagidaceae | page = 500 }}</ref>) Such references still cause difficulties in consulting earlier works. However, currently ''Gunnera'' is firmly assigned to the [[Monotypic taxon|monogeneric]] family Gunneraceae.<ref name="WWOK">{{cite journal | vauthors = Wanntorp L, Wanntorp HE, Oxelman B, Källersjö M | title = Phylogeny of ''Gunnera'' | journal = Plant Systematics and Evolution | date = March 2001 | volume = 226 | issue = 1 | pages = 85–107 | doi = 10.1007/s006060170075 | jstor = 23644101 | bibcode = 2001PSyEv.226...85W | s2cid = 42201778 }}</ref> | ||
The [[type species]] of the genus is ''[[Gunnera perpensa]]'' L.<ref name="IPNI">{{cite web | title=Gunnera | website=International Plant Names Index | url=https://www.ipni.org/n/30006147-2 | access-date=2025-11-14}}</ref> | |||
=== Evolution === | === Evolution === | ||
''Gunnera'' is thought to be a rather ancient group, with a well-documented fossil history due to the presence of fossilized [[pollen]] spores, known by the [[palynotaxon]] ''[[Tricolpites reticulatus]]''. It is a [[Gondwana]]n lineage, having originated in [[South America]] during the [[Cretaceous]]. The earliest fossilized pollen is known from the [[Late Cretaceous]] ([[Turonian]]) of Peru, about 90 million years ago, and within the following 10 million years, ''Gunnera'' had achieved a worldwide distribution, with fossil pollen grains being found in areas where it is not found today, such as western [[North America]], mainland [[Australia]], and [[Antarctica]].<ref>{{Cite journal | vauthors = Jarzen DM |date=1980 |title=The Occurrence of ''Gunnera'' Pollen in the Fossil Record | ''Gunnera'' is thought to be a rather ancient group, with a well-documented fossil history due to the presence of fossilized [[pollen]] spores, known by the [[palynotaxon]] ''[[Tricolpites reticulatus]]''. It is a [[Gondwana]]n lineage, having originated in [[South America]] during the [[Cretaceous]]. The earliest fossilized pollen is known from the [[Late Cretaceous]] ([[Turonian]]) of Peru, about 90 million years ago, and within the following 10 million years, ''Gunnera'' had achieved a worldwide distribution, with fossil pollen grains being found in areas where it is not found today, such as western [[North America]], mainland [[Australia]], and [[Antarctica]].<ref>{{Cite journal | vauthors = Jarzen DM |date=1980 |title=The Occurrence of ''Gunnera'' Pollen in the Fossil Record |journal=Biotropica |volume=12 |issue=2 |pages=117–123 |doi=10.2307/2387727 |jstor=2387727 |bibcode=1980Biotr..12..117J |issn=0006-3606}}</ref><ref name=":0">{{cite journal | vauthors = Bacon CD, Velásquez-Puentes FJ, Hinojosa LF, Schwartz T, Oxelman B, Pfeil B, Arroyo MT, Wanntorp L, Antonelli A | display-authors = 6 | title = Evolutionary persistence in ''Gunnera'' and the contribution of southern plant groups to the tropical Andes biodiversity hotspot | journal = PeerJ | volume = 6 | article-number = e4388 | date = 2018-03-16 | pmid = 29576938 | pmc = 5858603 | doi = 10.7717/peerj.4388 | doi-access = free }}</ref> Based on fossil pollen recovered from drilling cores, ''Gunnera'' is also known to have inhabited the now-submerged islands of the [[Ninety East Ridge|Ninetyeast Ridge]] during the [[Paleocene]], likely having dispersed there from either [[Australasia]] or the then-emergent [[Kerguelen Plateau]] islands.<ref>{{Cite journal | vauthors = Carpenter RJ, Truswell EM, Harris WK |date=2010-03-02 |title=Lauraceae fossils from a volcanic Palaeocene oceanic island, Ninetyeast Ridge, Indian Ocean: ancient long-distance dispersal?: Indian Ocean Lauraceae fossils |journal=Journal of Biogeography |language=en |volume=37 |issue=7 |pages=1202–1213 |doi=10.1111/j.1365-2699.2010.02279.x |s2cid=83060879}}</ref> | ||
Due to the widespread distribution of ''Gunnera'' during the Cretaceous, it was previously thought that the modern disjunct distribution of the genus was a relic of this period. However, phylogenetic analysis indicates that the majority of ''Gunnera'' species, even those found on entirely different continents, diverged from each other during the [[Cenozoic]], indicating that the modern distribution of ''Gunnera'' is a consequence of [[Long distance dispersal|long-distance dispersal]] from South America to other parts of the world, rather than relics of a former cosmopolitan distribution. The only species that diverged prior to the Cenozoic is ''[[Gunnera herteri]]'', described from Uruguay<ref>{{Cite journal | vauthors = Osten C |date=1932 |title=Una Gunnera en el Uruguay: Gunnera Herteri Osten n. sp. |journal= Herbarium Cornelius Osten Comunicaciones (Montevideo) |language=es |volume=1 |issue=2 |pages=33–39 }}</ref> and distributed in Uruguay and southeastern [[Brazil]], which is thought to be the most ancient species of the genus, its lineage having diverged during the Late Cretaceous, roughly concurrent with the oldest ''Gunnera'' fossil pollen from Peru. The persistence of the ''Gunnera'' [[crown group]] since the Cretaceous makes it unique among flowering plants, and may have been facilitated by strong [[Phylogenetic niche conservatism|niche conservatism]], [[Seed dispersal|dispersal]] ability, and being able to aggressively colonize disturbed land.<ref name=":0" /> | Due to the widespread distribution of ''Gunnera'' during the Cretaceous, it was previously thought that the modern disjunct distribution of the genus was a relic of this period. However, phylogenetic analysis indicates that the majority of ''Gunnera'' species, even those found on entirely different continents, diverged from each other during the [[Cenozoic]], indicating that the modern distribution of ''Gunnera'' is a consequence of [[Long distance dispersal|long-distance dispersal]] from South America to other parts of the world, rather than relics of a former cosmopolitan distribution. The only species that diverged prior to the Cenozoic is ''[[Gunnera herteri]]'', described from Uruguay<ref>{{Cite journal | vauthors = Osten C |date=1932 |title=Una Gunnera en el Uruguay: Gunnera Herteri Osten n. sp. |journal= Herbarium Cornelius Osten Comunicaciones (Montevideo) |language=es |volume=1 |issue=2 |pages=33–39 }}</ref> and distributed in Uruguay and southeastern [[Brazil]], which is thought to be the most ancient species of the genus, its lineage having diverged during the Late Cretaceous, roughly concurrent with the oldest ''Gunnera'' fossil pollen from Peru. The persistence of the ''Gunnera'' [[crown group]] since the Cretaceous makes it unique among flowering plants, and may have been facilitated by strong [[Phylogenetic niche conservatism|niche conservatism]], [[Seed dispersal|dispersal]] ability, and being able to aggressively colonize disturbed land.<ref name=":0" /> | ||
==Description== | ==Description== | ||
The 40–50 species vary enormously in [[leaf]] size, with the iconic large-leaved species belonging to the subgenus ''Panke''. The '''giant rhubarb''', or '''Campos des Loges''' (''[[Gunnera manicata]]''), native to the Serra do Mar mountains of southeastern [[Brazil]], is perhaps the largest species, with [[Wiktionary:reniform|reniform]] or sub-reniform leaves typically {{ | The 40–50 species vary enormously in [[leaf]] size, with the iconic large-leaved species belonging to the subgenus ''Panke''. The '''giant rhubarb''', or '''Campos des Loges''' (''[[Gunnera manicata]]''), native to the Serra do Mar mountains of southeastern [[Brazil]], is perhaps the largest species, with [[Wiktionary:reniform|reniform]] or sub-reniform leaves typically {{cvt|1.5|to|2.0|m|ft|0}} long, not including the thick, succulent [[petiole (botany)|petiole]] which may be up to {{cvt|2.5|m|ft|0}} in length. The width of the leaf blade is typically {{cvt|2.5|m|ft|0}}, but on two occasions cultivated specimens (in [[Dorset]], England in 2011<ref>{{cite web|url=https://www.bbc.com/news/uk-england-berkshire-15308919|title=Abbotsbury Gardens celebrates plant's 'monster' leaves |date=14 October 2011 |website=[[BBC]]}}</ref> and at Narrowwater, [[Ulster]], [[Ireland]]<ref>{{cite journal | title = Unknown | journal = The Garden | location = London | volume = 63 | issue = 1631 | date = 21 February 1903 | page = 125 }}</ref> in 1903) produced leaves fully {{cvt|3.3|m|ft|0}} in width. The seeds germinate best in very moist, but not wet, conditions and temperatures of 22–29 °C. | ||
Only slightly smaller is ''[[Gunnera masafuerae]]'' of the [[Juan Fernández Islands|Juan Fernandez Islands]] off the [[Chile]]an coast. They can have leaves up to {{ | Only slightly smaller is ''[[Gunnera masafuerae]]'' of the [[Juan Fernández Islands|Juan Fernandez Islands]] off the [[Chile]]an coast. They can have leaves up to {{cvt|2.9|m|ft|0}} in width on stout leaf stalks {{cvt|1.5|m|ft|0}} long and {{cvt|11|cm|in|frac=2}} thick according to Skottsberg.<ref>{{cite book | vauthors = Skottsberg C | chapter = The Phanerogams of Juan Fernandez Islands | title = The Natural History of Juan Fernandez and Ester Islands | location = Uppsala | publisher = Almquist & Wiksells Boktryckeri AB | date = 1953 | volume = 2 | page = 151 }}</ref> These leaf stalks or [[Petiole (botany)|petiole]]s are the thickest of any [[dicot]], and probably also the most massive. On nearby [[Alejandro Selkirk Island|Isla Más Afuera]], ''[[Gunnera peltata]]'' frequently has an upright trunk to {{cvt|5.5|m|ft|0}} in height by {{cvt|25|–|30|cm|in|frac=2}} thick, bearing leaves up to {{cvt|2|m|ft|0}} wide. The Hoja de Pantano (''[[Gunnera magnifica]]'') of the [[Colombia]]n [[Andes]] bears the largest leaf buds of any plant; up to {{cvt|60|cm|ft|0}} long and {{cvt|40|cm|in|0}} thick.<ref>{{cite journal | vauthors = St John H | title = ''Gunnera'' the Magnificent - Giant Herb of Colombia | journal = Chicago Natural History Museum Bulletin | volume = 30 | issue = 1 | date = January 1959 | page = 3 plus photo on front cover }}</ref> The [[Succulent plant|succulent]] leaf stalks are up to {{cvt|2.7|m|ft|0}} long. The massive [[inflorescence]] of small, reddish [[flower]]s is up to {{cvt|2.3|m|ft|0}} long and weighs about 13 kg. The flowers of ''Gunnera'' species are dimerous (two sepals, two petals (or none), two stamens (or one), and two carpels.<ref>{{cite web | url= https://www.plantsforuse.com/species/clusiaceae-garcinia#more-19 | work = Plants for Use | title = Gunneraceae ''Gunnera'' | access-date= January 20, 2009 }}</ref> Other giant ''Gunnera'' species within the subgenus ''Panke'' are found throughout the [[Neotropics]] and [[Hawaii]]. ''Gunnera insignis'' is also known by the name "poor man's umbrella" in [[Costa Rica]]. | ||
Outside of the subgenus ''Panke'', | Outside of the subgenus ''Panke'', some ''Gunnera'' species have small-to-medium-sized leaves. There are some species with moderately large leaves in [[Africa]] (''G. perpensa'', in the subgenus ''Gunnera'', [[synonym (taxonomy)|syn.]] subgenus ''Perpensum''); and [[Southeast Asia]] (''G. macrophylla'', in the subgenus ''Pseudogunnera''), but the others are low-lying, mat-forming plants with small leaves. There are several small species are found in [[New Zealand]], notably ''G. albocarpa'', with leaves only 1–2 cm long, and also in South America, with ''G. magellanica'' having leaves 5–9 cm wide on stalks 8–15 cm long. ''G. herteri,'' which is [[Sister group|sister]] to all other members of the genus<ref name=lw>{{Cite journal |last=Wanntorp |first=Livia |last2=Wanntorp |first2=Hans‐Erik |date=July 2003 |title=The biogeography of Gunnera L.: vicariance and dispersal |url=https://onlinelibrary.wiley.com/doi/10.1046/j.1365-2699.2003.00895.x |journal=Journal of Biogeography |language=en |volume=30 |issue=7 |pages=979–987 |doi=10.1046/j.1365-2699.2003.00895.x |issn=0305-0270}}</ref> and is found in Uruguay and Brazil, also has small leaves.<ref name=":1">{{Cite journal |last=Wilkinson |first=H |date=October 2000 |title=A revision of the anatomy of Gunneraceae |journal=Botanical Journal of the Linnean Society |volume=134 |issue=1–2 |pages=233–266 |doi=10.1006/bojl.2000.0372 |issn=0024-4074}}</ref><ref>{{Cite journal |last1=Rutishauser |first1=R. |last2=Wanntorp |first2=L. |last3=Pfeifer |first3=E. |date=2004 |title=''Gunnera herteri'' — developmental morphology of a dwarf from Uruguay and S Brazil (Gunneraceae) |journal=Plant Systematics and Evolution |volume=248 |issue=1/4 |pages=219–241 |doi=10.1007/s00606-004-0182-7 |jstor=23654236 |s2cid=24561923 |issn=0378-2697}}</ref> | ||
Some fossil leaf impressions of ''Gunnera'' from the [[Cretaceous]] of [[North America]] have large leaves akin to those of ''Panke'', and | Some fossil leaf impressions of ''Gunnera'' from the [[Cretaceous]] of [[North America]] have large leaves akin to those of ''Panke'', and ''G. mexicana'', which is sister to all other extant species within ''Panke'',<ref name=lw/> is the northernmost member. For this reason, it has been suggested that ''Panke'' originates from South American ''Gunnera'' that colonized North America during the Cretaceous and grew into giant forms, with the remaining South American ''Gunnera'' evolving into the subgenus ''Misandra'', with a low-lying, matlike growth. During the Cenozoic, the North American ''Panke'' would have colonized [[Hawaii]] and retreated southwards on the mainland before recolonizing South America. However, more recent phylogenetic evidence suggests that ''Misandra'' and ''Panke'' diverged only 15 million years ago, much too recent to assign the Cretaceous ''Gunnera'' to ''Panke''. Due to this, the large-leaved Cretaceous ''Gunnera'' from North America may represent a distinct lineage that [[Convergent evolution|convergently evolved]] giant leaves similar to those of ''Panke'', but did not leave any descendants. ''<ref name=":0" />''<ref name=":1" /><ref>{{Cite journal |last1=Wanntorp |first1=Livia |last2=Wanntorp |first2=Hans-Erik |date=July 2003 |title=The biogeography of ''Gunnera'' L.: vicariance and dispersal: Biogeography of ''Gunnera'' |url=http://doi.wiley.com/10.1046/j.1365-2699.2003.00895.x |journal=Journal of Biogeography |language=en |volume=30 |issue=7 |pages=979–987 |doi=10.1046/j.1365-2699.2003.00895.x|s2cid=83697864 |url-access=subscription }}</ref> | ||
==Species== | ==Species== | ||
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| || ''[[Gunnera lobata]]'' {{small|Hook.f.}} || Tierra del Fuego | | || ''[[Gunnera lobata]]'' {{small|Hook.f.}} || Tierra del Fuego | ||
|- | |- | ||
! rowspan="1" style="text-align:center;"| ''Gunnera'' | ! rowspan="1" style="text-align:center;"| ''[[Gunnera (Gunnera)|Gunnera]]'' | ||
|[[File:Gunnera perpensa - indigenous River Pumpkin of South Africa wetlands ponds.jpg|120px]] || ''[[Gunnera perpensa]]'' {{small|L.}} || Africa, Madagascar | |[[File:Gunnera perpensa - indigenous River Pumpkin of South Africa wetlands ponds.jpg|120px]] || ''[[Gunnera perpensa]]'' {{small|L.}} || Africa, Madagascar | ||
|- | |- | ||
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==Cyanobacterial symbiosis== | ==Cyanobacterial symbiosis== | ||
At least some species of ''Gunnera'' host [[Endosymbiont|endosymbiotic]] [[cyanobacteria]] such as ''[[Nostoc punctiforme]]''. The cyanobacteria provide [[Nitrogen fixation#Other plant symbionts|fixed nitrogen]] to the plant, while the plant provides fixed carbon to the microbe.<ref name="WFMJ">{{cite journal | vauthors = Wong FC, Meeks JC | title = Establishment of a functional symbiosis between the cyanobacterium Nostoc punctiforme and the bryophyte Anthoceros punctatus requires genes involved in nitrogen control and initiation of heterocyst differentiation | journal = Microbiology | volume = 148 | issue = Pt 1 | pages = 315–323 | date = January 2002 | pmid = 11782524 | doi = 10.1099/00221287-148-1-315 | doi-access = free }}</ref> The bacteria enter the plant via glands found at the base of each leaf stalk<ref name="Bergman">{{cite journal | vauthors = Bergman B, Johansson C, Söderbäck E | title = The ''Nostoc''-''Gunnera'' symbiosis | journal = The New Phytologist | volume = 122 | issue = 3 | pages = 379–400 | date = November 1992 | pmid = 33874210 | doi = 10.1111/j.1469-8137.1992.tb00067.x | doi-access = free }}</ref> and initiate an intracellular symbiosis which is thought to provide the plant with fixed nitrogen in return for fixed carbon for the bacterium. The Nostoc-filled symbiotic tissue makes up just a small portion of the plant's total biomass. ''Gunnera'' is the only known genus of angiosperms that hosts cyanobacteria, and the only known land plant with intracellular [[cyanobiont]]s. Although the endosymbionts enters the [[cell wall]], they do not penetrate the [[cell membrane]].<ref>{{cite book | url=https://books.google.com/books?id=_RxoEAAAQBAJ&dq=Nostoc+symbiotic+tissue+small+portion+total+Gunnera+biomass&pg=PA52 | title=Microbial Symbionts: Functions and Molecular Interactions on Host | isbn= | At least some species of ''Gunnera'' host [[Endosymbiont|endosymbiotic]] [[cyanobacteria]] such as ''[[Nostoc punctiforme]]''. The cyanobacteria provide [[Nitrogen fixation#Other plant symbionts|fixed nitrogen]] to the plant, while the plant provides fixed carbon to the microbe.<ref name="WFMJ">{{cite journal | vauthors = Wong FC, Meeks JC | title = Establishment of a functional symbiosis between the cyanobacterium Nostoc punctiforme and the bryophyte Anthoceros punctatus requires genes involved in nitrogen control and initiation of heterocyst differentiation | journal = Microbiology | volume = 148 | issue = Pt 1 | pages = 315–323 | date = January 2002 | pmid = 11782524 | doi = 10.1099/00221287-148-1-315 | doi-access = free }}</ref> The bacteria enter the plant via glands found at the base of each leaf stalk<ref name="Bergman">{{cite journal | vauthors = Bergman B, Johansson C, Söderbäck E | title = The ''Nostoc''-''Gunnera'' symbiosis | journal = The New Phytologist | volume = 122 | issue = 3 | pages = 379–400 | date = November 1992 | pmid = 33874210 | doi = 10.1111/j.1469-8137.1992.tb00067.x | doi-access = free }}</ref> and initiate an intracellular symbiosis which is thought to provide the plant with fixed nitrogen in return for fixed carbon for the bacterium. The Nostoc-filled symbiotic tissue makes up just a small portion of the plant's total biomass. ''Gunnera'' is the only known genus of angiosperms that hosts cyanobacteria, and the only known land plant with intracellular [[cyanobiont]]s. Although the endosymbionts enters the [[cell wall]], they do not penetrate the [[cell membrane]].<ref>{{cite book | url=https://books.google.com/books?id=_RxoEAAAQBAJ&dq=Nostoc+symbiotic+tissue+small+portion+total+Gunnera+biomass&pg=PA52 | title=Microbial Symbionts: Functions and Molecular Interactions on Host | isbn=978-0-323-99335-7 | last1=Dharumadurai | first1=Dhanasekaran | date=25 September 2022 | publisher=Elsevier }}</ref><ref>{{cite journal | pmc=1203372 | year=2005 | last1=Chiu | first1=W. L. | last2=Peters | first2=G. A. | last3=Levieille | first3=G. | last4=Still | first4=P. C. | last5=Cousins | first5=S. | last6=Osborne | first6=B. | last7=Elhai | first7=J. | title=Nitrogen Deprivation Stimulates Symbiotic Gland Development in ''Gunnera manicata'' | journal=Plant Physiology | volume=139 | issue=1 | pages=224–230 | doi=10.1104/pp.105.064931 | pmid=16113217 }}</ref> This relationship may provide insights to allow the creation of novel symbioses between crop plants and cyanobacteria, allowing growth in areas lacking fixed nitrogen in the soil. | ||
==Uses== | ==Uses== | ||
The stalks of ''[[Gunnera tinctoria|G. tinctoria]]'' (''nalca''), from southern [[Chile]] and [[Argentina]], are edible.<ref name="TasteAtlas">{{cite web |title=Nalca |url=https://www.tasteatlas.com/nalca |website=TasteAtlas |access-date=6 May 2023}}</ref> Their principal use is fresh consumption, after peeling, but also they are prepared in salads, liquor or marmalade. Leaves of this species are used in covering [[curanto]] (a traditional Chilean food). | The stalks of ''[[Gunnera tinctoria|G. tinctoria]]'' (''nalca''), from southern [[Chile]] and [[Argentina]], are edible.<ref name="TasteAtlas">{{cite web |title=Nalca |url=https://www.tasteatlas.com/nalca |website=TasteAtlas |access-date=6 May 2023}}</ref> Their principal use is fresh consumption, after peeling, but also they are prepared in salads, liquor or marmalade. Leaves of this species are used in covering [[curanto]] (a traditional Chilean food). | ||
''Gunnera perpensa'' is a source of traditional medicine in southern Africa, both in veterinary and human ailments, largely in obstetric and digestive complaints, but also as a wound dressing.<ref name="WattPP"/> It also is eaten in various ways, largely the petioles, flower stalks and leaves, fresh and raw, preferably with skins and fibre removed, which is said to remove bitterness, but also cooked. The plant also is said to be used in making a beer.<ref name="FFVF">{{cite book | vauthors = Fox FW | title = Food from the veld | publisher = Delta Books | date = 1982 | isbn = 978- | ''Gunnera perpensa'' is a source of traditional medicine in southern Africa, both in veterinary and human ailments, largely in obstetric and digestive complaints, but also as a wound dressing.<ref name="WattPP"/> It also is eaten in various ways, largely the petioles, flower stalks and leaves, fresh and raw, preferably with skins and fibre removed, which is said to remove bitterness, but also cooked. The plant also is said to be used in making a beer.<ref name="FFVF">{{cite book | vauthors = Fox FW | title = Food from the veld | publisher = Delta Books | date = 1982 | isbn = 978-0-908387-32-8 }}</ref> | ||
== References == | == References == | ||
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[[Category:Eudicot genera]] | [[Category:Eudicot genera]] | ||
[[Category:Medicinal plants]] | [[Category:Medicinal plants]] | ||
[[Category: | [[Category:Botanical taxa named by Carl Linnaeus]] | ||
Latest revision as of 18:21, 13 December 2025
Template:Short description Template:Automatic taxobox
Gunnera is the sole genus of herbaceous flowering plants in the family Gunneraceae, which contains 63 species. Some species in this genus, namely those in the subgenus Panke, have extremely large leaves. Species in the genus are variously native to Latin America, Australia, New Zealand, Papuasia, Hawaii, insular Southeast Asia, eastern Africa, and Madagascar.[1] The stalks of some species are edible.[2]
Taxonomy
Gunnera is the only genus in the family Gunneraceae.[3] The APG II system of 2003 accepted this family and assigned it to the order Gunnerales in the clade core eudicots. The family then consisted of one or two genera, Gunnera and, optionally, Myrothamnus, the latter optionally segregated as a separate family, Myrothamnaceae. This represents a change from the APG system of 1998, which accepted two separate families, unplaced as to order. The APG III system and APG IV system accept the family Gunneraceae, and places Myrothamnus in Myrothamnaceae; both families are placed in the order Gunnerales in the core eudicots.[4]
The genus Gunnera was named after the Norwegian botanist Johann Ernst Gunnerus. At first it was assigned to the family Haloragaceae, though that presented difficulties that led to the general recognition of the family Gunneraceae, as had been proposed about the beginning of the 20th century. In the meantime, in many publications it had been referred to as being in the Haloragaceae, variously misspelt (as for example "Halorrhagidaceae".[5]) Such references still cause difficulties in consulting earlier works. However, currently Gunnera is firmly assigned to the monogeneric family Gunneraceae.[6]
The type species of the genus is Gunnera perpensa L.[7]
Evolution
Gunnera is thought to be a rather ancient group, with a well-documented fossil history due to the presence of fossilized pollen spores, known by the palynotaxon Tricolpites reticulatus. It is a Gondwanan lineage, having originated in South America during the Cretaceous. The earliest fossilized pollen is known from the Late Cretaceous (Turonian) of Peru, about 90 million years ago, and within the following 10 million years, Gunnera had achieved a worldwide distribution, with fossil pollen grains being found in areas where it is not found today, such as western North America, mainland Australia, and Antarctica.[8][9] Based on fossil pollen recovered from drilling cores, Gunnera is also known to have inhabited the now-submerged islands of the Ninetyeast Ridge during the Paleocene, likely having dispersed there from either Australasia or the then-emergent Kerguelen Plateau islands.[10]
Due to the widespread distribution of Gunnera during the Cretaceous, it was previously thought that the modern disjunct distribution of the genus was a relic of this period. However, phylogenetic analysis indicates that the majority of Gunnera species, even those found on entirely different continents, diverged from each other during the Cenozoic, indicating that the modern distribution of Gunnera is a consequence of long-distance dispersal from South America to other parts of the world, rather than relics of a former cosmopolitan distribution. The only species that diverged prior to the Cenozoic is Gunnera herteri, described from Uruguay[11] and distributed in Uruguay and southeastern Brazil, which is thought to be the most ancient species of the genus, its lineage having diverged during the Late Cretaceous, roughly concurrent with the oldest Gunnera fossil pollen from Peru. The persistence of the Gunnera crown group since the Cretaceous makes it unique among flowering plants, and may have been facilitated by strong niche conservatism, dispersal ability, and being able to aggressively colonize disturbed land.[9]
Description
The 40–50 species vary enormously in leaf size, with the iconic large-leaved species belonging to the subgenus Panke. The giant rhubarb, or Campos des Loges (Gunnera manicata), native to the Serra do Mar mountains of southeastern Brazil, is perhaps the largest species, with reniform or sub-reniform leaves typically Template:Cvt long, not including the thick, succulent petiole which may be up to Template:Cvt in length. The width of the leaf blade is typically Template:Cvt, but on two occasions cultivated specimens (in Dorset, England in 2011[12] and at Narrowwater, Ulster, Ireland[13] in 1903) produced leaves fully Template:Cvt in width. The seeds germinate best in very moist, but not wet, conditions and temperatures of 22–29 °C.
Only slightly smaller is Gunnera masafuerae of the Juan Fernandez Islands off the Chilean coast. They can have leaves up to Template:Cvt in width on stout leaf stalks Template:Cvt long and Template:Cvt thick according to Skottsberg.[14] These leaf stalks or petioles are the thickest of any dicot, and probably also the most massive. On nearby Isla Más Afuera, Gunnera peltata frequently has an upright trunk to Template:Cvt in height by Template:Cvt thick, bearing leaves up to Template:Cvt wide. The Hoja de Pantano (Gunnera magnifica) of the Colombian Andes bears the largest leaf buds of any plant; up to Template:Cvt long and Template:Cvt thick.[15] The succulent leaf stalks are up to Template:Cvt long. The massive inflorescence of small, reddish flowers is up to Template:Cvt long and weighs about 13 kg. The flowers of Gunnera species are dimerous (two sepals, two petals (or none), two stamens (or one), and two carpels.[16] Other giant Gunnera species within the subgenus Panke are found throughout the Neotropics and Hawaii. Gunnera insignis is also known by the name "poor man's umbrella" in Costa Rica.
Outside of the subgenus Panke, some Gunnera species have small-to-medium-sized leaves. There are some species with moderately large leaves in Africa (G. perpensa, in the subgenus Gunnera, syn. subgenus Perpensum); and Southeast Asia (G. macrophylla, in the subgenus Pseudogunnera), but the others are low-lying, mat-forming plants with small leaves. There are several small species are found in New Zealand, notably G. albocarpa, with leaves only 1–2 cm long, and also in South America, with G. magellanica having leaves 5–9 cm wide on stalks 8–15 cm long. G. herteri, which is sister to all other members of the genus[17] and is found in Uruguay and Brazil, also has small leaves.[18][19]
Some fossil leaf impressions of Gunnera from the Cretaceous of North America have large leaves akin to those of Panke, and G. mexicana, which is sister to all other extant species within Panke,[17] is the northernmost member. For this reason, it has been suggested that Panke originates from South American Gunnera that colonized North America during the Cretaceous and grew into giant forms, with the remaining South American Gunnera evolving into the subgenus Misandra, with a low-lying, matlike growth. During the Cenozoic, the North American Panke would have colonized Hawaii and retreated southwards on the mainland before recolonizing South America. However, more recent phylogenetic evidence suggests that Misandra and Panke diverged only 15 million years ago, much too recent to assign the Cretaceous Gunnera to Panke. Due to this, the large-leaved Cretaceous Gunnera from North America may represent a distinct lineage that convergently evolved giant leaves similar to those of Panke, but did not leave any descendants. [9][18][20]
Species
since April 2023[update]Template:Dated maintenance category (articles)Script error: No such module "Check for unknown parameters"., Plants of the World Online accepts the following species[21] separated by subgenus:[22]
| Subgenus | Image | Scientific name | Distribution |
|---|---|---|---|
| Ostenigunnera Mattfeld, 1933Script error: No such module "Check for unknown parameters". | Gunnera herteri OstenScript error: No such module "Check for unknown parameters". | Uruguay, S Brazil | |
| Pseudogunnera Schindler, 1905 Script error: No such module "Check for unknown parameters". | File:COLLECTIE TROPENMUSEUM Bloeiende Gunnera macrophylla op de Karo-hoogvlakte TMnr 10012699.jpg | Gunnera macrophylla BlumeScript error: No such module "Check for unknown parameters". | Papuasia, Indonesia, Philippines |
| Milligania Schindler, 1905 Script error: No such module "Check for unknown parameters". | Gunnera albocarpa (Kirk) CockayneScript error: No such module "Check for unknown parameters". | New Zealand | |
| Gunnera arenaria Cheeseman ex KirkScript error: No such module "Check for unknown parameters". | New Zealand | ||
| Gunnera cordifolia (Hook.f.) Hook.f.Script error: No such module "Check for unknown parameters". | Tasmania | ||
| Gunnera densiflora Hook.f.Script error: No such module "Check for unknown parameters". | New Zealand | ||
| File:Gunnera dentata kz2.jpg | Gunnera dentata KirkScript error: No such module "Check for unknown parameters". | New Zealand | |
| Gunnera flavida ColensoScript error: No such module "Check for unknown parameters". | New Zealand | ||
| File:Gunnera hamiltonii kz03.jpg | Gunnera hamiltonii Kirk ex W.S.Ham.Script error: No such module "Check for unknown parameters". | New Zealand | |
| Gunnera mixta KirkScript error: No such module "Check for unknown parameters". | New Zealand | ||
| File:Gunnera monoica kz5.jpg | Gunnera monoica RaoulScript error: No such module "Check for unknown parameters". | New Zealand incl Chatham Islands | |
| File:Gunnera prorepens Hook.f. (AM AK351347-2).jpg | Gunnera prorepens Hook.f.Script error: No such module "Check for unknown parameters". | New Zealand | |
| Gunnera reniformis Ridl.Script error: No such module "Check for unknown parameters". | New Guinea | ||
| Gunnera strigosa (Kirk) ColensoScript error: No such module "Check for unknown parameters". | New Zealand | ||
| Panke Schindler, 1905 Script error: No such module "Check for unknown parameters". | Gunnera aequatoriensis L.E.MoraScript error: No such module "Check for unknown parameters". | Ecuador | |
| Gunnera annae Schindl.Script error: No such module "Check for unknown parameters". | Peru, Bolivia | ||
| Gunnera antioquensis L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera apiculata Schindl.Script error: No such module "Check for unknown parameters". | Bolivia, Argentina | ||
| Gunnera atropurpurea L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia, Ecuador | ||
| Gunnera berteroi Phil.Script error: No such module "Check for unknown parameters". | Bolivia, Argentina, Chile | ||
| Gunnera bogotana L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera bolivari J.F.Macbr.Script error: No such module "Check for unknown parameters". | Peru, Ecuador | ||
| Gunnera boliviana MorongScript error: No such module "Check for unknown parameters". | Bolivia | ||
| Gunnera bracteata Steud. ex Benn.Script error: No such module "Check for unknown parameters". | Robinson Crusoe Island in Chile | ||
| File:Gunnera brephogera (14726746092).jpg | Gunnera brephogea Linden & AndréScript error: No such module "Check for unknown parameters". | Colombia, Ecuador, Peru | |
| Gunnera caucana L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera colombiana L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia, Ecuador | ||
| Gunnera × cryptica J.M.H.ShawScript error: No such module "Check for unknown parameters". (G. manicata × G. tinctoria) | Cultivated | ||
| Gunnera cuatrecasasii L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera diazii L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera garciae-barrigae L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera hernandezii L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| File:Gunnera insignis.jpg | Gunnera insignis (Oerst.) Oerst.Script error: No such module "Check for unknown parameters". | Panama, Nicaragua, Costa Rica | |
| Gunnera × katherine-wilsoniae L.D.GómezScript error: No such module "Check for unknown parameters". (G. insignis × G. talamancana) | Costa Rica | ||
| Gunnera kauaiensis RockScript error: No such module "Check for unknown parameters". | Kauai in Hawaii | ||
| File:Gunnera killipiana (10620413034).jpg | Gunnera killipiana LundellScript error: No such module "Check for unknown parameters". | Chiapas, Guatemala, Honduras | |
| Gunnera lozanoi L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera magnifica H.St.JohnScript error: No such module "Check for unknown parameters". | Colombia | ||
| File:Gunnera manicata (Mammut-Blatt)-1.JPG | Gunnera manicata Linden ex AndréScript error: No such module "Check for unknown parameters". | S Brazil | |
| Gunnera margaretae Schindl.Script error: No such module "Check for unknown parameters". | Peru, Bolivia | ||
| Gunnera masafuerae Skottsb.Script error: No such module "Check for unknown parameters". | Alejandro Selkirk Island (Isla Mas Afuera) in Chile | ||
| File:IostephaneJBUNAM.JPG | Gunnera mexicana BrandegeeScript error: No such module "Check for unknown parameters". | Veracruz, Chiapas | |
| Gunnera morae Wanntorp & Klack.Script error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera peltata Phil.Script error: No such module "Check for unknown parameters". | Robinson Crusoe Island in Chile | ||
| Gunnera peruviana J.F.Macbr.Script error: No such module "Check for unknown parameters". | Ecuador, Peru | ||
| File:Gunnera petaloïdea (4743768827).jpg | Gunnera petaloidea Gaudich.Script error: No such module "Check for unknown parameters". | Hawaii | |
| Gunnera pilosa KunthScript error: No such module "Check for unknown parameters". | Peru, Bolivia, Ecuador | ||
| Gunnera pittieriana V.M.Badillo & Steyerm.Script error: No such module "Check for unknown parameters". | Venezuela | ||
| Gunnera quitoensis L.E.MoraScript error: No such module "Check for unknown parameters". | Ecuador | ||
| Gunnera saint-johnii (L.E.Mora) L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera sanctae-marthae L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera schindleri L.E.MoraScript error: No such module "Check for unknown parameters". | Bolivia, Argentina | ||
| File:Gunnera schultesii.jpeg | Gunnera schultesii L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | |
| Gunnera silvioana L.E.MoraScript error: No such module "Check for unknown parameters". | Ecuador, Colombia | ||
| Gunnera steyermarkii L.E.MoraScript error: No such module "Check for unknown parameters". | Venezuela | ||
| Gunnera tacueyana L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| Gunnera tajumbina L.E.MoraScript error: No such module "Check for unknown parameters". | Ecuador, Colombia | ||
| File:Gunnera talamancana - Flickr - gailhampshire.jpg | Gunnera talamancana H.Weber & L.E.MoraScript error: No such module "Check for unknown parameters". | Costa Rica, Panama | |
| Gunnera tamanensis L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | ||
| File:Gunnera tayrona (14746906163).jpg | Gunnera tayrona L.E.MoraScript error: No such module "Check for unknown parameters". | Colombia | |
| File:Gunnerales - Gunnera tinctoria - 39.jpg | Gunnera tinctoria (Molina) Mirb.Script error: No such module "Check for unknown parameters". | Chile, Argentina | |
| Gunnera venezolana L.E.MoraScript error: No such module "Check for unknown parameters". | Venezuela | ||
| Misandra Schindler, 1905 Script error: No such module "Check for unknown parameters". | File:Gunnera magellanica - Palmengarten Frankfurt - DSC01945.JPG | Gunnera magellanica Lam.Script error: No such module "Check for unknown parameters". | W + S South America, Falkland Is. |
| Gunnera lobata Hook.f.Script error: No such module "Check for unknown parameters". | Tierra del Fuego | ||
| Gunnera | File:Gunnera perpensa - indigenous River Pumpkin of South Africa wetlands ponds.jpg | Gunnera perpensa L.Script error: No such module "Check for unknown parameters". | Africa, Madagascar |
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In 2022, it was shown that plants in cultivation under the name Gunnera manicata were actually a hybrid, Gunnera × cryptica.[23]
Cyanobacterial symbiosis
At least some species of Gunnera host endosymbiotic cyanobacteria such as Nostoc punctiforme. The cyanobacteria provide fixed nitrogen to the plant, while the plant provides fixed carbon to the microbe.[24] The bacteria enter the plant via glands found at the base of each leaf stalk[25] and initiate an intracellular symbiosis which is thought to provide the plant with fixed nitrogen in return for fixed carbon for the bacterium. The Nostoc-filled symbiotic tissue makes up just a small portion of the plant's total biomass. Gunnera is the only known genus of angiosperms that hosts cyanobacteria, and the only known land plant with intracellular cyanobionts. Although the endosymbionts enters the cell wall, they do not penetrate the cell membrane.[26][27] This relationship may provide insights to allow the creation of novel symbioses between crop plants and cyanobacteria, allowing growth in areas lacking fixed nitrogen in the soil.
Uses
The stalks of G. tinctoria (nalca), from southern Chile and Argentina, are edible.[2] Their principal use is fresh consumption, after peeling, but also they are prepared in salads, liquor or marmalade. Leaves of this species are used in covering curanto (a traditional Chilean food).
Gunnera perpensa is a source of traditional medicine in southern Africa, both in veterinary and human ailments, largely in obstetric and digestive complaints, but also as a wound dressing.[5] It also is eaten in various ways, largely the petioles, flower stalks and leaves, fresh and raw, preferably with skins and fibre removed, which is said to remove bitterness, but also cooked. The plant also is said to be used in making a beer.[28]
References
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- ↑ Kew World Checklist of Selected Plant Families
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External links
- Template:Sister-inline
- Template:Sister-inline
- Gunneraceae Template:Webarchive in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants Template:Webarchive: descriptions, illustrations, identification, information retrieval. Version: 21 May 2006. http://delta-intkey.com Template:Webarchive
- links at CSDL Template:Webarchive
- Gunnera magellanica pictures from Chilebosque
- Global Invasive Species database tinctoria
- Medicinal plant details Gunnera perpensa
- Script error: No such module "citation/CS1".
Template:Angiosperm families Template:Taxonbar Template:Authority control