''Lophosoria quadripinnata'' (

J.F.Gmel. Johann Friedrich Gmelin (8 August 1748 – 1 November 1804) was a German natural history, naturalist, chemist, botanist, entomologist, herpetologist, and malacologist. Education Johann Friedrich Gmelin was born as the eldest son of Philipp F ...

) C.Chr. is a species of fern that, according to DNA molecular analysis, belongs to the family Dicksoniaceae, where it is placed in the genus ''

Lophosoria Lophosoria is a genus of tree ferns within the family Dicksoniaceae. Although it is confined to tropical America in modern times, there is fossil evidence that it was once spread throughout Gondwana with the exception of New Zealand. Species ...

''. It is found in

the Americas The Americas, sometimes collectively called America, are a landmass comprising the totality of North America and South America.'' Webster's New World College Dictionary'', 2010 by Wiley Publishing, Inc., Cleveland, Ohio. When viewed as a sin ...

spanning from

Cuba Cuba, officially the Republic of Cuba, is an island country, comprising the island of Cuba (largest island), Isla de la Juventud, and List of islands of Cuba, 4,195 islands, islets and cays surrounding the main island. It is located where the ...

and

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

Chile Chile, officially the Republic of Chile, is a country in western South America. It is the southernmost country in the world and the closest to Antarctica, stretching along a narrow strip of land between the Andes, Andes Mountains and the Paci ...

. In Chile it is present in the area between

Talca Talca () is a city and commune in Chile located about south of Santiago, and is the capital of both Talca Province and Maule Region (7th Region of Chile). As of the 2012 census, the city had a population of 201,142. The city is an important ...

and Aysén including

Juan Fernández Islands The Juan Fernández Islands () are a sparsely inhabited series of islands in the South Pacific Ocean, reliant on tourism and fishing. Situated off the coast of Chile, they are composed of three main volcanic islands: Robinson Crusoe Island, R ...

. In Argentina it grows only in the humid valleys of western

Neuquén Neuquén (; ) is the capital city of the Argentine province of Neuquén and of the Confluencia Department, located in the east of the province. It occupies a strip of land west of the confluence of the Limay and Neuquén rivers which form t ...

and

Río Negro Province Río Negro (, ''Black River'') is a province of Argentina, located in northern Patagonia. Neighboring provinces are from the south clockwise Chubut, Neuquén, Mendoza, La Pampa and Buenos Aires. To the east lies the Atlantic Ocean. Its cap ...

. Diamondleaf fern is a common name. In Spanish it is known as 'ampe' (from the

Mapudungun Mapuche ( , ; from 'land' and 'people', meaning 'the people of the land') or Mapudungun (from 'land' and 'speak, speech', meaning 'the speech of the land'; also spelled Mapuzugun and Mapudungu) is either a language isolate or member of the s ...

añpe) or palmilla, but one has to remember that there are several species of ferns called "palmillas" that have larger or smaller fronds, and which grow in colder climates. It is a medium-sized plant, growing to about 4–5 feet (though 10–12 feet in a sheltered place at Arduaine Garden in Argyll, Scotland) and even though the

rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...

does not grow a trunk, it is clearly related to the other tree ferns due to features that were apparently already present in their common ancestor, like 'pneumathodes', and the

which changed from the dorsiventral symmetry typical of the other ferns, to a radial symmetry typical of tree ferns. Their large and multiple pinnate fronds, with the petiole raised adaxially, and the hairs on the rhizome and lower part of the petioles, also resemble those of tree ferns. To identify the species, use the position and characteristics of the spores found on the fertile fronds. The genus already existed in the

Cretaceous Period The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 77.1 million years, it is the ninth and longest geologi ...

in southern

Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...

according to fossil remains found in

Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...

. The species is well known as an ornamental plant.

Description

''Lophosoria quadripinnata'' is a

vascular plant Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...

with two alternating generations, a

sporophyte A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...

and a

gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...

, multicellular and independent; with spores as a means of dispersion and survival. The gametophyte is a "thallus" (undifferentiated tissue), and the sporophyte is a "corm" (with roots, shoots, and a vascular system). Due to these characteristics it is usually classified as a "

pteridophyte A pteridophyte is a vascular plant (with xylem and phloem) that reproduces by means of spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as " cryptogams", meaning that their means of reproduction is ...

". They have sporophytes with megaphylls or "fronds" (

Euphyllophytina The euphyllophytes are a clade of plants within the tracheophytes (the vascular plants). The group may be treated as an unranked clade, a division under the name Euphyllophyta or a subdivision under the name Euphyllophytina. The euphyllophytes ...

). The rhizome is massive, with hairs, not growing a trunk (not arborescent), and with radial symmetry instead of horizontal; a characteristic apparently originating in the ancestor of the tree ferns. The fronds are large in size, 2-3 pinnations, with hairs on the under side of the petioles, and they are high on its abaxial part, all common characteristics of the

Cyatheales The order Cyatheales, which includes most tree ferns, is a Taxonomy (biology), taxonomic order of the fern class, Polypodiopsida. No clear morphology (biology), morphological features characterize all of the Cyatheales, but DNA sequence data indi ...

order (the tree ferns clade). Also present are the pneumathodes (discrete vent lines, or patches, present on the rachis, petiole and rhizome, whose function is facilitating gas exchange in the dense tissue), which are characteristic of the tree fern clade. ---- The sori, without indusia (covering), are located on the abaxial (lower) surface, on the veins of the last pinnae. The leaf margin does not form part of an indusium, as in other Dicksoniaceae. The sori have numerous hairs (called "filiform

paraphyses Paraphyses are erect sterile filament-like support structures occurring among the reproductive apparatuses of fungi, ferns, bryophytes and some thallophytes. The singular form of the word is paraphysis. In certain fungi, they are part of the f ...

") between the

sporangia A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...

. The characteristics and position of the sori are unique and can be used to identify the species. Like all Polypodiopsidas, the sporangia are leptosporangiate (with a foot, capsule with a single-cell wall thickness, and dehiscent ring in the capsule). As in all the tree ferns, the ring is oblique and complete, not interrupted by the foot of the sporangium. The spores have trilete marks. This species has spores with a very unusual morphology, with a central belt ("cincture") that meant for a long time the species was assigned its own family, Lophosoriaceae. The germination of the gametophyte corresponds to the genus ''Cyathea'', giving rise to short strands of from two to six cells in both varieties. The prothallus development is of the ''Adiantum'' type. When raised in the laboratory, gametophytes of the quadripinnata variety are always chordates, however gametophytes of the ''contracta'' variety possess three morphs, depending on the density of spores that are developed (Dyer 1979Dyer, A.F. 1979. "The culture of fern gametophytes for experimental investigation." p. 253-305. In A.F. Dyer (ed.) ''The experimental biology of ferns.'' Academic Press, Londres.). At low density, they are long spatulates with a central meristem and a slightly more developed wing than the other; at medium densities they are cordiforms and at high densities are band-like with a large amount of antheridia. The gametophyte is protandric (a hermaphrodite, first

antheridia An antheridium is a haploid structure or organ producing and containing male gametes (called ''antherozoids'' or sperm). The plural form is antheridia, and a structure containing one or more antheridia is called an androecium. The androecium is al ...

are developed that give antherozoids, and then the archegonia that gives the oospheres). The number of x

chromosome A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...

s = 65.

Taxonomy

:''Theoretical introduction to

Phylogenetics In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

and

Taxonomy image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...

Genetic analysis Genetic analysis is the overall process of studying and researching in fields of science that involve genetics and molecular biology. There are a number of applications that are developed from this research, and these are also considered parts ...

has placed the species unequivocally in the family

Dicksoniaceae Dicksoniaceae is a group of tropical, subtropical and warm temperate ferns, treated as a family in the Pteridophyte Phylogeny Group classification of 2016 (PPG I), and counting 30-40 species. Alternatively, the family may be sunk into a very broa ...

, but the story of its taxonomic placement is long. Pichi Sermolli (1970Pichi Sermolli R. E. G. 1970. "Fragmenta Pteridologiae". -II. ''Webbia.'' 24: 699-722.) placed it in its own family, Lophosoriaceae Pichi Sermolli, where it was located for a long time. At first it was thought to be a relation of the

Cyatheaceae The Cyatheaceae are a family of ferns, the scaly tree ferns, one of eight families in the order Cyatheales in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Alternatively, the family may defined much more broadly (Cyatheaceae ...

because of the characteristic abaxial sori, or the pattern of gametophyte germination. In the 1990s a close relationship was hypothesized with

Metaxyaceae ''Metaxya'' is a neotropical genus of ferns in the order Cyatheales. It is the only genus in the family Metaxyaceae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Alternatively, the genus may be placed in the subfamily Me ...

because of characteristics of the petiole and stem morphology but subsequent analyzes suggested that these families were not related (D. S. Conant, unpublished data, cited in Wolf ''et al.'' 1999Wolf ''et al.'' 1999. "Phylogenetic relationships of the enigmatic fern families Hymenophyllopsidaceae and Lophosoriaceae: evidences from ''rbc''L nucleotide sequences". ''Plant Syst. Evol.'' 219: 263-270). On the other hand, ultrastructural studies (using

scanning electron microscopy A scanning electron microscope (SEM) is a type of electron microscope that produces images of a sample by scanning the surface with a focused beam of electrons. The electrons interact with atoms in the sample, producing various signals that ...

) of the spores (Gastony y Tyron 1976Gastony G. J. y Tyron R. 1976. "Spore morphology in the Cyatheaceae". II. The genera ''Lophosoria'', ''Metaxya'', ''Sphaeropteris'', ''Alsophila'', and ''Nephelea''. ''Amer J. Bot.'' 63: 738-758) and recent studies of gametophyte development (Pérez-García ''et al.'' 1995Pérez-García B., Fraile M. E., Mendoza A. 1995. "Desarrollo del gametofito del ''Lophosoria quadripinnata'' (Filicales: Lophosoriaceae)". ''Revista Biol. Trop.'' 43: 55-60.) have concluded that ''Lophosoria'' differs significantly from the characteristics of other tree ferns, so their relationship to the others remains unknown. Other researchers, such as Kubitzki in Kramer (1990Kubitzki, K. 1990. Pteridophytes and Gymnosperms. en: K. V. Kramer. P. S. Green (Eds.) ''The families and genera of vascular plants.'' Vol 1. Springer-Verlag, Berlín, Alemania.) have it nested within Dicksoniaceae along with other genera that today are classified in other families of tree ferns. As molecular studies of the species continued, there was an increasingly obvious affinity with the family

, such as having the same number of chromosomes, and molecular DNA analysis done on ''rbc''L sequence in 1999 also placed it within that family (Wolf ''et al.'' 1999), a relationship that was confirmed in an extensive molecular study of ferns made by Korall et al. (2006,Korall ''et al.'' 2006. "Tree ferns: Monophiletic groups and their relationships as revealed by four protein-coding plastid loci". ''Molecular phylogenetics and Evolution'' 39: 830-845. on the sequences atpA, atpB, rps4 and the aforementioned). Because of that the species was placed in the family Dicksoniaceae in its own monotypic genus in Smith's 2006 classification system of monilophytes.A. R. Smith, K. M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, P. G. Wolf. 2006. "A classification for extant ferns". ''Taxonomy'' 55(3), 705-731 (pd
here
) Circumscription: the species consists of at least two varieties, sometimes placed as species of the genus: * ''Lophosoria quadripinnata'' (J.F.Gmel.) C.Chr. variety ''quadripinnata'' ** wide distribution in the tropical parts of

, from the mild and humid regions of South America, between 1000 and 3000

meters above sea level Height above mean sea level is a measure of a location's vertical distance (height, elevation or altitude) in reference to a vertical datum based on a historic mean sea level. In geodesy, it is formalized as orthometric height. The zero level vari ...

*''Lophosoria quadripinnata'' variety ''contracta'' ( Hieron.) R.M.Tryon & A.F.Tryon ** found in Ecuador, at 2900

, and on

Robinson Crusoe Island Robinson Crusoe Island (, ) is the second largest of the Juan Fernández Islands, situated 670 km (362 nmi; 416 mi) west of San Antonio, Chile, San Antonio, Chile, in the South Pacific Ocean. It is the more populous of the inhabit ...

, Juan Fernandez Archipelago, Chile The spores of the ''contracta'' variety are slightly bigger (70 x 75 μm) than those of the ''quadripinnata'' variety (50 × 60 μm). Furthermore, the ''quadripinnata'' variety's antheridia are made up of 5 cells: 2 baseline, 2 annular, and a small elliptical operculum, while those of the variety ''contracta'' are formed by three cells, one basal an annular and operculum (Stockey 1930Stokey, A.G. 1930. "Prothallia of the Cyatheaceae." ''Bot. Gaz.'' (Crawfordsville) 90: 1-45.). They also differ in the morphology of their gametophytes. Another variety ''Lophosoria quadripinnata'' variety ''quesadae'' A.Rojas has also been described in Costa Rica and Panama (Rojas-Alvarado 1996, it was described as ''Lophosoria quesadae'' speciesRojas Alvarado A. F. 1996. "Aportes a la flora Pteridophyta costarricense". II. Taxones nuevos. ''Brenesia'' 45-46: 33-50).

Evolution

Dettmann (1986Dettmann M. E. 1986. "Significance of the Cretaceous-Tertiary spore genus ''Cyatheacidites'' in tracing the origin and migration of ''Lophosoria'' (Filicopsida). ''Special Papers in Palaeontology'' 35: 63-94.) has remarked that the spores of the extinct genus ''Cyatheacidites'' correspond very well with living ''Lophosoria''. The distribution of the ''Cyatheacidites'' has been used to infer that ''Lophosoria'' was located in the southern part of Gondwana during the Early

, and then migrated to

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

(Dettmann 1986). More recently, Cantrill (1998Cantrill D. J. 1998. "Early Cretaceous fern foliage from President Head, South Island, Antarctica. ''Alcheringa'' 22: 241-258) described fossilized leaves in the early Cretaceous layer (more specifically in the Aptian) in Antarctica, which contained spores of ''Cyatheacidites''. The preservation of the leaves and spores was so good that Cantrill could identify the species, and described them by the name of ''Lophosoria cupulata'' D.J.Cantrill (published as ''Lophosoria cupulatus'') with the genus change, as he considered ''Lophosoria quadripinnata'' (J.F.Gmel.) C.Chr. was the living species more closely related to the species found.

Economic importance

It is used as an

ornamental plant Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...

in parks and gardens, as well as cut foliage in flower bouquets.Smith-Ramírez C. 1994. "Usos artesanales del bosque nativo. La extracción silenciosa", Revista Ambiente y Desarrollo, X, (2), 71-7
pdf here
The plants used in latter are removed from their natural environment, creating certain local conservation issues. Some local markets in the south of Chile sell their sprouts, called ''perritos'', for consuming in salads.Hoffmann J. 1999. "El bosque chilote. Historia natural del archipiélago de Chiloé. Conservación y desarrollo sustentable de sus bosques y biodiversidad", Santiago: Defensores del Bosque Chileno,

References

External links

{{Taxonbar, from=Q6678562 Dicksoniaceae Ferns of Chile Flora of the Juan Fernández Islands Flora of the Valdivian temperate forests Flora of the Neotropical realm