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Gunnera is the sole genus of 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 , , , , , insular Southeast Asia, eastern , and . Kew World Checklist of Selected Plant Families The stalks of some species are edible.


Taxonomy
Gunnera is the only genus in the family Gunneraceae. The APG II system of 2003 accepted this family and assigned it to the order Gunnerales in the clade . The family then consisted of one or two genera, Gunnera and, optionally, , the latter optionally segregated as a separate family, . This represents a change from the 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.

The genus Gunnera was named after the Norwegian botanist Johann Ernst Gunnerus. At first it was assigned to the family , 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".) Such references still cause difficulties in consulting earlier works. However, currently Gunnera is firmly assigned to the family Gunneraceae.

The of the genus is L.


Evolution
Gunnera is thought to be a rather ancient group, with a well-documented fossil history due to the presence of fossilized spores, known by the Tricolpites reticulatus. It is a lineage, having originated in during the . The earliest fossilized pollen is known from the () 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 , mainland , and . 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 , likely having dispersed there from either or the then-emergent Kerguelen Plateau islands.

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 , 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 , described from Uruguay and distributed in Uruguay and southeastern , 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 since the Cretaceous makes it unique among flowering plants, and may have been facilitated by strong niche conservatism, ability, and being able to aggressively colonize disturbed land.


Description
The 40–50 species vary enormously in size, with the iconic large-leaved species belonging to the subgenus Panke. The giant rhubarb, or Campos des Loges ( ), native to the Serra do Mar mountains of southeastern , is perhaps the largest species, with or sub-reniform leaves typically long, not including the thick, succulent petiole which may be up to in length. The width of the leaf blade is typically , but on two occasions cultivated specimens (in , England in 2011 and at Narrowwater, , in 1903) produced leaves fully 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 coast. They can have leaves up to in width on stout leaf stalks long and thick according to Skottsberg. These leaf stalks or petioles are the thickest of any , and probably also the most massive. On nearby Isla Más Afuera, frequently has an upright trunk to in height by thick, bearing leaves up to wide. The Hoja de Pantano ( Gunnera magnifica) of the bears the largest leaf buds of any plant; up to long and thick. The leaf stalks are up to long. The massive of small, reddish is up to 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. Other giant Gunnera species within the subgenus Panke are found throughout the and . Gunnera insignis is also known by the name "poor man's umbrella" in .

Outside of the subgenus Panke, some Gunnera species have small-to-medium-sized leaves. There are some species with moderately large leaves in ( G. perpensa, in the subgenus Gunnera, syn. subgenus Perpensum); and ( 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 , 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 to all other members of the genus and is found in Uruguay and Brazil, also has small leaves.

Some fossil leaf impressions of Gunnera from the of have large leaves akin to those of Panke, and G. mexicana, which is sister to all other extant species within Panke, 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 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.


Species
, Plants of the World Online accepts the following species separated by subgenus:
New Zealand
Tasmania
New Zealand
New Zealand
New Zealand
New Zealand
New Zealand
New Zealand incl Chatham Islands
New Zealand
New Guinea
New Zealand
Peru, Bolivia
Colombia
Bolivia, Argentina
Colombia, Ecuador
Bolivia, Argentina, Chile
Colombia
Peru, Ecuador
Bolivia
Robinson Crusoe Island in Chile
Colombia, Ecuador, Peru
Colombia
Colombia, Ecuador
Cultivated
Colombia
Colombia
Colombia
Colombia
Panama, Nicaragua, Costa Rica
Costa Rica
Kauai in Hawaii
Chiapas, Guatemala, Honduras
Colombia
Colombia
S Brazil
Peru, Bolivia
Alejandro Selkirk Island (Isla Mas Afuera) in Chile
Veracruz, Chiapas
Colombia
Robinson Crusoe Island in Chile
Ecuador, Peru
Hawaii
Peru, Bolivia, Ecuador
Venezuela
Ecuador
Colombia
Colombia
Bolivia, Argentina
Colombia
Ecuador, Colombia
Venezuela
Colombia
Ecuador, Colombia
Costa Rica, Panama
Colombia
Colombia
Chile, Argentina
Venezuela
Tierra del Fuego

In 2022, it was shown that plants in cultivation under the name Gunnera manicata were actually a hybrid, Gunnera × cryptica.


Cyanobacterial symbiosis
At least some species of Gunnera host such as Nostoc punctiforme. The cyanobacteria provide fixed nitrogen to the plant, while the plant provides fixed carbon to the microbe. The bacteria enter the plant via glands found at the base of each leaf stalk 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 . Although the endosymbionts enters the , they do not penetrate the .
(2022). 9780323993357, Elsevier. .
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 and , are edible. 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 (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. 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.

(1982). 9780908387328, Delta Books.


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