Gunnera

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Gunnera
Temporal range: Late Cretaceous - present, 90–0  Ma
Gunnera tinctoria 2.jpg
Gunnera tinctoria at the San Francisco Botanical Garden at Strybing Arboretum
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Gunnerales
Family: Gunneraceae
Meisn. [1]
Genus: Gunnera
L.
Gunnera distribution.svg
The range of the genus Gunnera [2]
Synonyms [3]
  • MilliganiaHook.f., rejected name
  • PankeMolina
  • PankeaOerst.
Gunnera manicata, Devon, England Gunnera.manicata.arp.750pix.jpg
Gunnera manicata, Devon, England
Gunnera insignis, Costa Rica Gunnera insignis.jpg
Gunnera insignis, Costa Rica

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, Africa, and Madagascar. [3] The stalks of some species are edible. [4]

Contents

Taxonomy

Gunnera is the only genus in the family Gunneraceae. [5] The APG II system, of 2003, also recognizes this family and assigns 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 firmly recognized two separate families, unplaced as to order. The APG III system and APG IV system recognizes 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]

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. [9] [10] 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. [11]

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 [12] 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. [10]

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 1.5 to 2.0 meters (4 ft 11 in to 6 ft 7 in) long, not including the thick, succulent petiole which may be up to 2.5 meters (8 feet 2 inches) in length. The width of the leaf blade is typically 2.5 meters (8 feet 2 inches), but on two occasions cultivated specimens (in Dorset, England in 2011 [13] and at Narrowwater, Ulster, Ireland [14] in 1903) produced leaves fully 3.3 meters (10 feet 10 inches) in width. The seeds germinate best in very moist, but not wet, conditions and temperatures of 22–29 °C.

Only slightly smaller is G. masafuerae of the Juan Fernandez Islands off the Chilean coast. They can have leaves up to 2.9 m (9 ft 6 in) in width on stout leaf stalks 1.5 m (4 ft 11 in) long and 11 cm (4+1⁄2 in) thick according to Skottsberg. [15] These leafstalks or petioles are the thickest of any dicot, and probably also the most massive. On nearby Isla Más Afuera, G. peltata frequently has an upright trunk to 5.5 m (18 ft 1 in) in height by 25–30 cm (10–12 in) thick, bearing leaves up to 2 m (6 ft 7 in) wide. The Hoja de Pantano (G. magnifica) of the Colombian Andes bears the largest leaf buds of any plant; up to 60 cm (2 ft) long and 40 cm (16 in) thick. [16] The succulent leaf stalks are up to 2.7 m (8 ft 10 in) long. The massive inflorescence of small, reddish flowers is up to 2.3 m (7 ft 7 in) 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. [17] 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, most of the more basal Gunnera species have small-to-medium-sized leaves. There are some species with moderately large leaves in Africa (G. perpensa, in the subgenus Perpensum) and Southeast Asia (G. macrophylla, in the subgenus Pseudogunnera), but the majority of more basal species 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. The most basal species in the genus, G. herteri of 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 the most basal extant species within Panke (G. mexicana) is the most northern 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. [10] [18] [20]

Species

As of April 2023, Plants of the World Online accepts the following species [21] separated by subgenus: [22]

SubgenusImageScientific nameDistribution
OstenigunneraMattfeld, 1933 Gunnera herteri OstenUruguay, S Brazil
PseudogunneraSchindler, 1905 COLLECTIE TROPENMUSEUM Bloeiende Gunnera macrophylla op de Karo-hoogvlakte TMnr 10012699.jpg Gunnera macrophylla BlumePapuasia, Indonesia, Philippines
MilliganiaSchindler, 1905 Gunnera albocarpa (Kirk) CockayneNew Zealand
Gunnera arenaria Cheeseman ex KirkNew Zealand
Gunnera cordifolia (Hook.f.) Hook.f.Tasmania
Gunnera densiflora Hook.f.New Zealand
Gunnera dentata kz2.jpg Gunnera dentata KirkNew Zealand
Gunnera flavida ColensoNew Zealand
Gunnera hamiltonii kz03.jpg Gunnera hamiltonii Kirk ex W.S.Ham.New Zealand
Gunnera mixta KirkNew Zealand
Gunnera monoica kz5.jpg Gunnera monoica RaoulNew Zealand incl Chatham Islands
Gunnera prorepens Hook.f. (AM AK351347-2).jpg Gunnera prorepens Hook.f.New Zealand
Gunnera reniformis Ridl.New Guinea
Gunnera strigosa (Kirk) ColensoNew Zealand
PankeSchindler, 1905 Gunnera aequatoriensis L.E.MoraEcuador
Gunnera annae Schindl.Peru, Bolivia
Gunnera antioquensis L.E.MoraColombia
Gunnera apiculata Schindl.Bolivia, Argentina
Gunnera atropurpurea L.E.MoraColombia, Ecuador
Gunnera berteroi Phil.Bolivia, Argentina, Chile
Gunnera bogotana L.E.MoraColombia
Gunnera bolivari J.F.Macbr.Peru, Ecuador
Gunnera boliviana MorongBolivia
Gunnera bracteata Steud. ex Benn.Robinson Crusoe Island in Chile
Gunnera brephogera (14726746092).jpg Gunnera brephogea Linden & AndréColombia, Ecuador, Peru
Gunnera caucana L.E.MoraColombia
Gunnera colombiana L.E.MoraColombia, Ecuador
Gunnera × cryptica J.M.H.Shaw (G. manicata × G. tinctoria)Cultivated
Gunnera cuatrecasasii L.E.MoraColombia
Gunnera diazii L.E.MoraColombia
Gunnera garciae-barrigae L.E.MoraColombia
Gunnera hernandezii L.E.MoraColombia
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 RockKauai in Hawaii
Gunnera killipiana (10620413034).jpg Gunnera killipiana LundellChiapas, Guatemala, Honduras
Gunnera lozanoi L.E.MoraColombia
Gunnera magnifica H.St.JohnColombia
Gunnera manicata (Mammut-Blatt)-1.JPG Gunnera manicata Linden ex AndréS Brazil
Gunnera margaretae Schindl.Peru, Bolivia
Gunnera masafuerae Skottsb.Alejandro Selkirk Island (Isla Mas Afuera) in Chile
IostephaneJBUNAM.JPG Gunnera mexicana BrandegeeVeracruz, Chiapas
Gunnera morae Wanntorp & Klack.Colombia
Gunnera peltata Phil.Robinson Crusoe Island in Chile
Gunnera peruviana J.F.Macbr.Ecuador, Peru
Gunnera petaloidea (4743768827).jpg Gunnera petaloidea Gaudich.Hawaii
Gunnera pilosa KunthPeru, Bolivia, Ecuador
Gunnera pittieriana V.M.Badillo & Steyerm.Venezuela
Gunnera quitoensis L.E.MoraEcuador
Gunnera saint-johnii (L.E.Mora) L.E.MoraColombia
Gunnera sanctae-marthae L.E.MoraColombia
Gunnera schindleri L.E.MoraBolivia, Argentina
Gunnera schultesii.jpeg Gunnera schultesii L.E.MoraColombia
Gunnera silvioana L.E.MoraEcuador, Colombia
Gunnera steyermarkii L.E.MoraVenezuela
Gunnera tacueyana L.E.MoraColombia
Gunnera tajumbina L.E.MoraEcuador, Colombia
Gunnera talamancana - Flickr - gailhampshire.jpg Gunnera talamancana H.Weber & L.E.MoraCosta Rica, Panama
Gunnera tamanensis L.E.MoraColombia
Gunnera tayrona (14746906163).jpg Gunnera tayrona L.E.MoraColombia
Gunnerales - Gunnera tinctoria - 39.jpg Gunnera tinctoria (Molina) Mirb.Chile, Argentina
Gunnera venezolana L.E.MoraVenezuela
MisandraSchindler, 1905 Gunnera magellanica - Palmengarten Frankfurt - DSC01945.JPG Gunnera magellanica Lam.W + S South America, Falkland Is.
Gunnera lobata Hook.f.Tierra del Fuego
Gunnera 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. [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 [2] 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 landplants with intracellular cyanobionts. Although the endosymbionts enters the cell wall, they do not penetrate the cell membrane. [25] [26] 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. [27]

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<i>Gunnera <span style="font-style:normal;">×</span> cryptica</i> Hybrid between Gunnera manicata and Gunnera tinctoria

Gunnera × cryptica is a hybrid between Gunnera manicata and Gunnera tinctoria. Both species were introduced into cultivation in Western Europe towards the end of the 19th century. The hybrid occurred spontaneously in cultivation, in Europe probably around 1873. Both morphological and molecular analyses have shown that plants widely cultivated in Britain and Ireland under the name Gunnera manicata were all the hybrid, and that the true species G. manicata is no longer found there. This situation may also be the case in Europe and other parts of the world. The hybrid is more resistant to frost and adverse conditions than G. manicata, which is likely to be why it has replaced it in cultivation.

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References

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  13. ↑ "Abbotsbury Gardens celebrates plant's 'monster' leaves". BBC . 14 October 2011.
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  16. ↑ 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.
  17. ↑ "Gunneraceae Gunnera". Plants for Use. Retrieved January 20, 2009.
  18. 1 2 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.
  19. ↑ 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.
  20. ↑ 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.
  21. ↑ "Gunnera L. - Plants of the World Online". Plants of the World Online. 2020-06-22. Retrieved 2023-04-25.
  22. ↑ 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.
  23. ↑ 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.
  24. ↑ 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.
  25. ↑ Dharumadurai, Dhanasekaran (25 September 2022). Microbial Symbionts: Functions and Molecular Interactions on Host. Elsevier. ISBN   9780323993357.
  26. ↑ 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.
  27. ↑ Fox FW (1982). Food from the veld. Delta Books. ISBN   978-0908387328.
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