Gunnera
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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.[2] The stalks of some species are edible.[4]
Taxonomy
Gunnera is the only genus in the family Gunneraceae.[5] 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.[6]
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".[7]) Such references still cause difficulties in consulting earlier works. However, currently Gunnera is firmly assigned to the monogeneric family Gunneraceae.[8]
The type species of the genus is Gunnera perpensa L.[9]
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.[10][11] 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.[12]
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[13] 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.[11]
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[14] and at Narrowwater, Ulster, Ireland[15] 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.[16] 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.[17] 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.[18] 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[19] and is found in Uruguay and Brazil, also has small leaves.[20][21]
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,[19] 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. [11][20][22]
Species
As of April 2023[update], Plants of the World Online accepts the following species[23] separated by subgenus:[24]
| Subgenus | Image | Scientific name | Distribution |
|---|---|---|---|
| Ostenigunnera Mattfeld, 1933 | Gunnera herteri Osten | Uruguay, S Brazil | |
| Pseudogunnera Schindler, 1905 | File:COLLECTIE TROPENMUSEUM Bloeiende Gunnera macrophylla op de Karo-hoogvlakte TMnr 10012699.jpg | Gunnera macrophylla Blume | Papuasia, Indonesia, Philippines |
| Milligania Schindler, 1905 | Gunnera albocarpa (Kirk) Cockayne | New Zealand | |
| Gunnera arenaria Cheeseman ex Kirk | New Zealand | ||
| Gunnera cordifolia (Hook.f.) Hook.f. | Tasmania | ||
| Gunnera densiflora Hook.f. | New Zealand | ||
| File:Gunnera dentata kz2.jpg | Gunnera dentata Kirk | New Zealand | |
| Gunnera flavida Colenso | New Zealand | ||
| File:Gunnera hamiltonii kz03.jpg | Gunnera hamiltonii Kirk ex W.S.Ham. | New Zealand | |
| Gunnera mixta Kirk | New Zealand | ||
| File:Gunnera monoica kz5.jpg | Gunnera monoica Raoul | New Zealand incl Chatham Islands | |
| File:Gunnera prorepens Hook.f. (AM AK351347-2).jpg | Gunnera prorepens Hook.f. | New Zealand | |
| Gunnera reniformis Ridl. | New Guinea | ||
| Gunnera strigosa (Kirk) Colenso | New Zealand | ||
| Panke Schindler, 1905 | Gunnera aequatoriensis L.E.Mora | Ecuador | |
| Gunnera annae Schindl. | Peru, Bolivia | ||
| Gunnera antioquensis L.E.Mora | Colombia | ||
| Gunnera apiculata Schindl. | Bolivia, Argentina | ||
| Gunnera atropurpurea L.E.Mora | Colombia, Ecuador | ||
| Gunnera berteroi Phil. | Bolivia, Argentina, Chile | ||
| Gunnera bogotana L.E.Mora | Colombia | ||
| Gunnera bolivari J.F.Macbr. | Peru, Ecuador | ||
| Gunnera boliviana Morong | Bolivia | ||
| Gunnera bracteata Steud. ex Benn. | Robinson Crusoe Island in Chile | ||
| File:Gunnera brephogera (14726746092).jpg | Gunnera brephogea Linden & André | Colombia, Ecuador, Peru | |
| Gunnera caucana L.E.Mora | Colombia | ||
| Gunnera colombiana L.E.Mora | Colombia, Ecuador | ||
| Gunnera × cryptica J.M.H.Shaw (G. manicata × G. tinctoria) | Cultivated | ||
| Gunnera cuatrecasasii L.E.Mora | Colombia | ||
| Gunnera diazii L.E.Mora | Colombia | ||
| Gunnera garciae-barrigae L.E.Mora | Colombia | ||
| Gunnera hernandezii L.E.Mora | Colombia | ||
| File:Gunnera insignis.jpg | Gunnera insignis (Oerst.) Oerst. | Panama, Nicaragua, Costa Rica | |
| Gunnera × katherine-wilsoniae L.D.Gómez (G. insignis × G. talamancana) | Costa Rica | ||
| Gunnera kauaiensis Rock | Kauai in Hawaii | ||
| File:Gunnera killipiana (10620413034).jpg | Gunnera killipiana Lundell | Chiapas, Guatemala, Honduras | |
| Gunnera lozanoi L.E.Mora | Colombia | ||
| Gunnera magnifica H.St.John | Colombia | ||
| File:Gunnera manicata (Mammut-Blatt)-1.JPG | Gunnera manicata Linden ex André | S Brazil | |
| Gunnera margaretae Schindl. | Peru, Bolivia | ||
| Gunnera masafuerae Skottsb. | Alejandro Selkirk Island (Isla Mas Afuera) in Chile | ||
| File:IostephaneJBUNAM.JPG | Gunnera mexicana Brandegee | Veracruz, Chiapas | |
| Gunnera morae Wanntorp & Klack. | Colombia | ||
| Gunnera peltata Phil. | Robinson Crusoe Island in Chile | ||
| Gunnera peruviana J.F.Macbr. | Ecuador, Peru | ||
| File:Gunnera petaloïdea (4743768827).jpg | Gunnera petaloidea Gaudich. | Hawaii | |
| Gunnera pilosa Kunth | Peru, Bolivia, Ecuador | ||
| Gunnera pittieriana V.M.Badillo & Steyerm. | Venezuela | ||
| Gunnera quitoensis L.E.Mora | Ecuador | ||
| Gunnera saint-johnii (L.E.Mora) L.E.Mora | Colombia | ||
| Gunnera sanctae-marthae L.E.Mora | Colombia | ||
| Gunnera schindleri L.E.Mora | Bolivia, Argentina | ||
| File:Gunnera schultesii.jpeg | Gunnera schultesii L.E.Mora | Colombia | |
| Gunnera silvioana L.E.Mora | Ecuador, Colombia | ||
| Gunnera steyermarkii L.E.Mora | Venezuela | ||
| Gunnera tacueyana L.E.Mora | Colombia | ||
| Gunnera tajumbina L.E.Mora | Ecuador, Colombia | ||
| File:Gunnera talamancana - Flickr - gailhampshire.jpg | Gunnera talamancana H.Weber & L.E.Mora | Costa Rica, Panama | |
| Gunnera tamanensis L.E.Mora | Colombia | ||
| File:Gunnera tayrona (14746906163).jpg | Gunnera tayrona L.E.Mora | Colombia | |
| File:Gunnerales - Gunnera tinctoria - 39.jpg | Gunnera tinctoria (Molina) Mirb. | Chile, Argentina | |
| Gunnera venezolana L.E.Mora | Venezuela | ||
| Misandra Schindler, 1905 | File:Gunnera magellanica - Palmengarten Frankfurt - DSC01945.JPG | Gunnera magellanica Lam. | W + S South America, Falkland Is. |
| Gunnera lobata Hook.f. | Tierra del Fuego | ||
| Gunnera | File:Gunnera perpensa - indigenous River Pumpkin of South Africa wetlands ponds.jpg | Gunnera perpensa L. | Africa, Madagascar |
In 2022, it was shown that plants in cultivation under the name Gunnera manicata were actually a hybrid, Gunnera × cryptica.[25]
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.[26] The bacteria enter the plant via glands found at the base of each leaf stalk[3] 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.[27][28] 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.[4] 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.[7] 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.[29]
References
Page Template:Reflist/styles.css has no content.
- ^ Page Module:Citation/CS1/styles.css has no content.Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. hdl:10654/18083.
- ^ a b Kew World Checklist of Selected Plant Families
- ^ a b Page Module:Citation/CS1/styles.css has no content.Bergman B, Johansson C, Söderbäck E (November 1992). "The Nostoc-Gunnera symbiosis". The New Phytologist. 122 (3): 379–400. doi:10.1111/j.1469-8137.1992.tb00067.x. PMID 33874210.
- ^ a b Page Module:Citation/CS1/styles.css has no content."Nalca". TasteAtlas. Retrieved 6 May 2023.
- ^ Page Module:Citation/CS1/styles.css has no content.Christenhusz MJ, Byng JW (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
- ^ Page Module:Citation/CS1/styles.css has no content.Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385. ISSN 0024-4074.
- ^ a b Page Module:Citation/CS1/styles.css has no content.Watt JM, Breyer-Brandwijk MG, Gerdina M (1962). "Halorrhagidaceae". The Medicinal and Poisonous Plants of Southern and Eastern Africa (Second ed.). E & S Livingstone. p. 500.
- ^ Page Module:Citation/CS1/styles.css has no content.Wanntorp L, Wanntorp HE, Oxelman B, Källersjö M (March 2001). "Phylogeny of Gunnera". Plant Systematics and Evolution. 226 (1): 85–107. Bibcode:2001PSyEv.226...85W. doi:10.1007/s006060170075. JSTOR 23644101. S2CID 42201778.
- ^ Page Module:Citation/CS1/styles.css has no content."Gunnera". International Plant Names Index. Retrieved 2025-11-14.
- ^ Page Module:Citation/CS1/styles.css has no content.Jarzen DM (1980). "The Occurrence of Gunnera Pollen in the Fossil Record". Biotropica. 12 (2): 117–123. Bibcode:1980Biotr..12..117J. doi:10.2307/2387727. ISSN 0006-3606. JSTOR 2387727.
- ^ a b c Page Module:Citation/CS1/styles.css has no content.Bacon CD, Velásquez-Puentes FJ, Hinojosa LF, Schwartz T, Oxelman B, Pfeil B, et al. (2018-03-16). "Evolutionary persistence in Gunnera and the contribution of southern plant groups to the tropical Andes biodiversity hotspot". PeerJ. 6 e4388. doi:10.7717/peerj.4388. PMC 5858603. PMID 29576938.
- ^ Page Module:Citation/CS1/styles.css has no content.Carpenter RJ, Truswell EM, Harris WK (2010-03-02). "Lauraceae fossils from a volcanic Palaeocene oceanic island, Ninetyeast Ridge, Indian Ocean: ancient long-distance dispersal?: Indian Ocean Lauraceae fossils". Journal of Biogeography. 37 (7): 1202–1213. doi:10.1111/j.1365-2699.2010.02279.x. S2CID 83060879.
- ^ Page Module:Citation/CS1/styles.css has no content.Osten C (1932). "Una Gunnera en el Uruguay: Gunnera Herteri Osten n. sp". Herbarium Cornelius Osten Comunicaciones (Montevideo) (in español). 1 (2): 33–39.
- ^ Page Module:Citation/CS1/styles.css has no content."Abbotsbury Gardens celebrates plant's 'monster' leaves". BBC. 14 October 2011.
- ^ Page Module:Citation/CS1/styles.css has no content."Unknown". The Garden. 63 (1631). London: 125. 21 February 1903.
- ^ Page Module:Citation/CS1/styles.css has no content.Skottsberg C (1953). "The Phanerogams of Juan Fernandez Islands". The Natural History of Juan Fernandez and Ester Islands. Vol. 2. Uppsala: Almquist & Wiksells Boktryckeri AB. p. 151.
- ^ Page Module:Citation/CS1/styles.css has no content.St John H (January 1959). "Gunnera the Magnificent - Giant Herb of Colombia". Chicago Natural History Museum Bulletin. 30 (1): 3 plus photo on front cover.
- ^ Page Module:Citation/CS1/styles.css has no content."Gunneraceae Gunnera". Plants for Use. Retrieved January 20, 2009.
- ^ a b Page Module:Citation/CS1/styles.css has no content.Wanntorp, Livia; Wanntorp, Hans‐Erik (July 2003). "The biogeography of Gunnera L.: vicariance and dispersal". Journal of Biogeography. 30 (7): 979–987. doi:10.1046/j.1365-2699.2003.00895.x. ISSN 0305-0270.
- ^ a b Page Module:Citation/CS1/styles.css has no content.Wilkinson, H (October 2000). "A revision of the anatomy of Gunneraceae". Botanical Journal of the Linnean Society. 134 (1–2): 233–266. doi:10.1006/bojl.2000.0372. ISSN 0024-4074.
- ^ Page Module:Citation/CS1/styles.css has no content.Rutishauser, R.; Wanntorp, L.; Pfeifer, E. (2004). "Gunnera herteri — developmental morphology of a dwarf from Uruguay and S Brazil (Gunneraceae)". Plant Systematics and Evolution. 248 (1/4): 219–241. doi:10.1007/s00606-004-0182-7. ISSN 0378-2697. JSTOR 23654236. S2CID 24561923.
- ^ Page Module:Citation/CS1/styles.css has no content.Wanntorp, Livia; Wanntorp, Hans-Erik (July 2003). "The biogeography of Gunnera L.: vicariance and dispersal: Biogeography of Gunnera". Journal of Biogeography. 30 (7): 979–987. doi:10.1046/j.1365-2699.2003.00895.x. S2CID 83697864.
- ^ Page Module:Citation/CS1/styles.css has no content."Gunnera L. - Plants of the World Online". Plants of the World Online. 2020-06-22. Retrieved 2023-04-25.
- ^ Page Module:Citation/CS1/styles.css has no content.WANNTORP, LIVIA; WANNTORP, HANS-ERIK; RUTISHAUSER, ROLF (2003). "On the homology of the scales in Gunnera (Gunneraceae)". Botanical Journal of the Linnean Society. 142 (3). Oxford University Press (OUP): 301–308. doi:10.1046/j.1095-8339.2003.00185.x. ISSN 1095-8339.
- ^ Page Module:Citation/CS1/styles.css has no content.Shaw, Julian M.H.; Edwards, Dawn & David, John (2022). "A new spontaneous hybrid in Gunnera subgenus Panke (Gunneraceae) widespread in the British Isles, with notes on the typification of G. manicata". British & Irish Botany. 4 (3): 364–384. doi:10.33928/bib.2022.04.364. S2CID 252243597.
- ^ Page Module:Citation/CS1/styles.css has no content.Wong FC, Meeks JC (January 2002). "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". Microbiology. 148 (Pt 1): 315–323. doi:10.1099/00221287-148-1-315. PMID 11782524.
- ^ Page Module:Citation/CS1/styles.css has no content.Dharumadurai, Dhanasekaran (25 September 2022). Microbial Symbionts: Functions and Molecular Interactions on Host. Elsevier. ISBN 978-0-323-99335-7.
- ^ Page Module:Citation/CS1/styles.css has no content.Chiu, W. L.; Peters, G. A.; Levieille, G.; Still, P. C.; Cousins, S.; Osborne, B.; Elhai, J. (2005). "Nitrogen Deprivation Stimulates Symbiotic Gland Development in Gunnera manicata". Plant Physiology. 139 (1): 224–230. doi:10.1104/pp.105.064931. PMC 1203372. PMID 16113217.
- ^ Page Module:Citation/CS1/styles.css has no content.Fox FW (1982). Food from the veld. Delta Books. ISBN 978-0-908387-32-8.
External links
- Page Template:Sister-inline/styles.css has no content.Script error: No such module "Sister project logo". Media related to Script error: No such module "Commons link". at Wikimedia Commons
- Template:Wikispecies-inline
- Gunneraceae Script error: No such module "webarchive". in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants Script error: No such module "webarchive".: descriptions, illustrations, identification, information retrieval. Version: 21 May 2006. http://delta-intkey.com Script error: No such module "webarchive".
- links at CSDL Script error: No such module "webarchive".
- Gunnera magellanica pictures from Chilebosque
- Global Invasive Species database tinctoria
- Medicinal plant details Gunnera perpensa
- Page Module:Citation/CS1/styles.css has no content."Gunnera on Achill Island". Achill Journal. Archived from the original on 2007-09-29.
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