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Aglaomorpha(plant)

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Aglaomorpha
Temporal range:Piacenzianto Present
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Drynaria quercifoliaat theLincoln Park Conservatory
Scientific classificationEdit this classification
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Suborder: Polypodiineae
Family: Polypodiaceae
Subfamily: Drynarioideae
Genus: Aglaomorpha
Schott
Species

Seetext

Synonyms[1]
  • ChristiopterisCopel.
  • DrynariaJ.Sm.
  • DryostachyumJ.Sm.
  • MerinthosorusCopel.
  • PhotinopterisJ.Sm.
  • PseudodrynariaC.Chr.
  • ThayeriaCopel.

Aglaomorphais agenusoffernsin the subfamilyDrynarioideaeof the familyPolypodiaceae.[1]ThePteridophyte Phylogeny Groupclassification of 2016 (PPG I) uses this genus name,[1]while other sources useDrynariato includeAglaomorpha.[2]Species are commonly known asbasket ferns.Ascircumscribedin PPG I, the genus contains around 50 species.[1]

Basket ferns areepiphyticorepipetricand are native to tropical Africa, South Asia, East Asia, Southeast Asia, Australia, and Oceania. Some species are economically important asmedicinal plants.

Description

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Aglaomorpha rigidulagrowing on rocks in Australia. The fertile foliagefrondsare large and dark green, the smaller brown sterile nest fronds are clustered at their bases.
Sorion theabaxialsurface of the foliagefrondofAglaomorpha quercifolia.

Basket ferns are characterized by the presence of two types offronds,fertile foliage fronds and sterile nest fronds. The dark green foliage fronds are large, 2–4 feet (0.61–1.22 m) long, with elongated stalks. They are deeply lobed orpinnate,winged, and bearsori(structures producing and containingspores) on the bottom surfaces.

The nest fronds are smaller rounded leaves basal to the foliage fronds. They do not bear sori and are persistent, not being shed after turning brown and dying. They form a characteristic 'basket' that collect litter and organic debris, hence the common name.[3]The collected debris decompose intohumus,providing the plants with nutrients it would otherwise not have received from being suspended above the ground.[4][5]

Both frond types grow fromrhizomestypically anchored to a tree or a rock.[6][7]The rhizomes of basket ferns are creeping and densely covered in brown scales.

Habitat and distribution

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Basket ferns areepiphytic(growing on trees) orepipetric(growing on rocks). They can also sometimes be found in man-made structures like brick walls.[8]They are found in wet tropical environments, usually inrainforests.[3]Their native range extends from equatorial Africa to tropicalSouthand East Asia, Southeast Asia, Australia, and Oceania.[9]

Life cycle

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The "Drynaria"type prothallial germination.

Like otherspore-bearing plants,Aglaomorphaexhibits metagenesis or thealternation of generations.One generation being thediploidmulticellularsporophyte(the phase where the plant is most familiar), and the other being thehaploidmulticellulargametophyte(the phase where the plant is known as aprothallus). Gametophytes develop fromsporesreleased by mature sporophytes; while sporophytes, in turn, develop from the fusion of gametes produced by mature prothalli.[8][10]

The synonymDrynarialends its name to a certain type of prothallial germination, the 'Drynariatype', observed in several other ferns. In this type, the spores germinate into a germ filament composed of barrel-shapedchlorophyllouscells with one or morerhizoidsat the base cell. The tipmost cell divides repeatedly by cross-walls, forming a broad spatulate (spoon-shaped) prothallial plate. One of the cells at the top margin of the prothallus then divides obliquely when it has 5, 10, or more cells across its width. This results in anobconicalmeristematic cell.Division by this type of cell is parallel to each other and perpendicular to the rest of the cells, forming rows. This eventually results in the formation of a notch at the anterior edge of the prothallus, giving it a roughly heart-shaped appearance (cordate).[11]

The cordate prothallus are usually smaller with thinner midribs than that of other members ofPolypodiaceae.They are also usually more sparsely haired, with some prothalli rarely having multicellular hair. They mature after six to nine months, and finish their life cycle at around a year. The gametophytes produce male (antheridium), and female (archegonium)gametes.The gametes fuse, forming the diploid sporophyte, the 'fern' part of the life cycle.[8][10][11]

Aglaomorphaalso naturally exhibitsapospory,the production of a gametophyte not from spores, but directly from the vegetative cells of the sporophytes. Their leaves can develop prothalli under dim light and sporophytic buds in strong light.[12]

Ecology

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Antsfeeding onnectarsecreted by the lower surface of a youngAglaomorpha quercifoliafrond.

Aglaomorpha,like some othergeneraof ferns (includingPolybotryaandPolypodium), possess specializednectar-secreting structures (nectaries) on the bases of the frond lobes or the underside of the fronds. The produced nectar is rich insugarsandamino acids.[13]

Their function may be to attractants(or other organisms) for protection or forsporedispersal. They may also be simply excretory organs (hydathodes), used for exuding surplus metabolic products.[13]The ant speciesIridomyrmex cordatusis commonly associated withA. quercifolia,in addition to other epiphytic plants.[14]

In Australia,Aglaomorpha rigidulaserve as shelter for amethystine pythons (Morelia amethistina) and scrub pythons (Morelia kinghorni).[15]As much as 81% of sightings of the snakes in one study were in large individuals ofA. rigidulalocated about 17–40 m (56–131 ft) above the ground. Snakes seek shelter inA. rigidulamore frequently during the colder seasons.[16]

In the 19th century,Indigenous Australianswere documented by theNorwegianexplorerCarl Sofus Lumholtzto have hunted pythons regularly during the winter months by climbing up to individuals ofA. rigidula.[17] The large rhizome mass ofAglaomorphacan also serve as growing substrates for other plants like the ribbon fern (Ophioglossum pendulum).[18]Due to their ability to preserve moisture and persistence even after death, the nest leaves ofAglaomorphaare also fertile hosts to a large number ofwater-borne fungi.[19]

Classification

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Basket ferns are classified under the subfamilyDrynarioideaeof the familyPolypodiaceae.[1]Species belonging toAglaomorphawere once classified under the genusPolypodium(rockcap ferns), under thesubgenusAglaomorpha.[20][21]

Species

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The following is the list of accepted species according to theChecklist of Ferns and Lycophytes of the Worldas of August 2019:[22]

Traditional medicine

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Extracts from the rhizomes of someAglaomorphaspecies are used extensively intraditional medicine.[23][24]In China, Taiwan,Vietnam,Thailand, andLaos,the rhizomes of gu-sui-bu,Aglaomorpha fortunei(more frequently cited by Asian authors by itsillegitimatesynonymDrynaria fortunei),[25]are commonly used to treat bone injuries.[8][26]Its common name literally means "mender of shattered bones" in Chinese.[27]Another species, the oak-leaf fern (Aglaomorpha quercifolia) is used similarly in South Asia andMaritime Southeast Asia.[10][28]

Conservation

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A basket fern inRavensbourne National Park,Australia

Species ofAglaomorphacommonly used in traditional medicine likeA. roosiiandA. quercifoliaare in danger of being overexploited. None of the species are currently cultivated for thealternative medicineindustry.[8][10]

Aglaomorphaare also consideredendangeredin some areas (like inNew South Wales,Australia), due to threats ofhabitat lossand low population numbers.[10][29]

Evolution and fossil record

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Fossil species have not been transferred toAglaomorphainPPG I,and are still recorded in the genusDrynaria.In 2010, twelve well-preservedfossilspecimens were described from theSanying Formationof theYangjie coal mineof China. NamedDrynaria callispora,it comes from thePiacenzianage of thePlioceneepoch(about 3.6 to 2.5 million years ago).[30]

Thefossil recordof drynarioids is not very well documented due to the typically poor preservation offossilsrecovered. Previous fossil species assigned toDrynariaincludeDrynaria astrostigma,D. dura,andD. tumulosafrom theCenomanianof theCzech Republic;andD. durum,all assigned tentatively to the genus in 1899. The arrangement and type of their sori, however, indicate that they are members of the familyMatoniaceaeinstead.[30]

Outside the genus,Protodrynaria takhtajanifrom theEocene-Oligoceneboundary ofKursk Oblast,Russia shows some affinities toDrynariabut only distantly. The only other reasonably convincing fossil remains of drynarioids aside fromD. callisporawas a specimen namedPolypodium quercifoliarecovered in 1985 from theLate Miocene(23.03 to 5.332 million years ago) ofPalembang,Indonesia.These were later transferred to the living speciesAglaomorpha heraclea.It remains, as of 2011, the oldest known drynarioid.[31][30]

See also

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References

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  1. ^abcdePPG I (2016). "A community-derived classification for extant lycophytes and ferns".Journal of Systematics and Evolution.54(6): 563–603.doi:10.1111/jse.12229.S2CID39980610.
  2. ^"AglaomorphaSchott ".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved2019-08-14.
  3. ^abRussell Cumming; Jan Sked (August 21, 2006)."Drynaria&Platycerium:Interesting Fern Genera ".Society for Growing Australian Plants. Archived fromthe originalon August 23, 2011.RetrievedJuly 27,2011.
  4. ^Frederick Garrett Dickason (1946)."The Ferns of Burma"(PDF).The Ohio Journal of Science.46(3): 109–141.RetrievedJuly 28,2011.
  5. ^Cliver Jermy; Roger Chapman (2002).Tropical Forest Expeditions(PDF).RGS-IBG Expedition Advisory Centre, Royal Geographical Society. p. 15.ISBN0-907649-84-X.
  6. ^William Jackson Hooker (1862).Garden ferns; or, Coloured figures and descriptions of a selection of exotic ferns adapted for cultivation in the garden.Illustrated by Walter Hood Fitch. p.5.
  7. ^Robert Lee Riffle (1998).The tropical look: an encyclopedia of dramatic landscape plants.Timber Press. p. 152.ISBN978-0-88192-422-0.
  8. ^abcdeH.C. Chang; D.C. Agrawal; C.L. Kuo; J.L. Wen; C.C. Chen; H.S. Tsay (2007). "In VitroCulture ofDrynaria fortunei,a Fern Species Source of Chinese Medicine "Gu-Sui-Bu"".In Vitro Cellular & Developmental Biology - Plant.43(2). Springer: 133–139.doi:10.1007/s11627-007-9037-6.ISSN1475-2689.S2CID10349428.
  9. ^"Drynaria(Bory) J.Sm ".Global Biodiversity Information Facility.RetrievedJuly 27,2011.
  10. ^abcdeP.B. Mazumder; Bonani Mazumder; M. Dutta Choudhury; G.D. Sharma (2011)."In VitroPropagation ofDrynaria quercifolia(L.) J. Sm., a Medicinal Fern ".Assam University Journal of Science & Technology: Biological and Environmental Sciences.7(1). Assam University: 79–83.ISSN0975-2773.Archived fromthe originalon September 29, 2011.RetrievedJuly 31,2011.
  11. ^abB. K. Nayar; S. Kaur (1971)."Gametophytes of Homosporous Ferns"(PDF).The Botanical Review.37(3). The New York Botanical Garden: 295–396.Bibcode:1971BotRv..37..295N.doi:10.1007/bf02859157.ISSN0006-8101.S2CID12733899.RetrievedJuly 31,2011.
  12. ^"The Opening Face of Vascular Plants: Pteridophyta".Competition Science Vision.10(113). Pratiyogita Darpan: 650–661. 2007.RetrievedJuly 31,2011.
  13. ^abSuzanne Koptur; Alan R. Smith; Irene Baker (1982)."Nectaries in Some Neotropical Species ofPolypodium(Polypodiaceae): Preliminary Observations and Analyses "(PDF).Biotropica.14(2). Association for Tropical Biology: 108–113.Bibcode:1982Biotr..14..108K.doi:10.2307/2387739.JSTOR2387739.Archived fromthe original(PDF)on April 6, 2012.RetrievedJuly 28,2011.
  14. ^Victor Rico-Gray; Paulo S. Oliveria (2007).The Ecology and Evolution of Ant-Plant Interactions.Interspecific Interactions. The University of Chicago Press. p. 48.ISBN978-0-226-71348-9.
  15. ^Jason Cummings; Michelle Martin; Anne Rogers (2006)."Quantifying the abundance of four large epiphytic fern species in remnant complex notophyll vine forest on the Atherton Tableland, north Queensland, Australia"(PDF).Cunninghamia.9(4). Royal Botanic Gardens, Sydney & the National Herbarium of New South Wales: 521–527. Archived fromthe original(PDF)on March 29, 2011.RetrievedJuly 30,2011.
  16. ^Alastair Freeman; Amanda Freeman (2009)."Habitat Use in a Large Rainforest Python (Morelia kinghorni) in the Wet Tropics of North Queensland, Australia "(PDF).Herpetological Conservation and Biology.4(2): 252–260.RetrievedJuly 30,2011.
  17. ^Alastair Freeman; Amanda Freeman (2007)."Giants in the Rainforest: A Radiotelemetry Study of the Amethystine Python in North Queensland, Australia"(PDF).Iguana.14(4). International Reptile Conservation Foundation: 215–221. Archived fromthe original(PDF)on October 2, 2011.RetrievedJuly 30,2011.
  18. ^Elvianna Dorante-Day."Ferns and Fern Allies".Bush Medicine Woman. Archived fromthe originalon March 28, 2012.RetrievedJuly 31,2011.
  19. ^Kishore Karamchand; Kandikerere R. Sridhar (2009)."Association of water-borne conidial fungi with epiphytic tree fern (Drynaria quercifolia) ".Acta Mycologica.44(1). Swiss Association for International Cooperation: 19–27.doi:10.5586/am.2009.004.
  20. ^Michael Hassler; Brian Swale."Family Drynariaceae, genusDrynaria;world species list ".Checklist of World Ferns. Archived fromthe originalon October 17, 2008.RetrievedJuly 27,2011.
  21. ^D. J. Mabberley (2008).Mabberley's plant-book: a portable dictionary of plants, their classification and uses.Cambridge University Press. p. 690.ISBN978-0-521-82071-4.
  22. ^Hassler, Michael & Schmitt, Bernd (June 2019). "Aglaomorpha".Checklist of Ferns and Lycophytes of the World.Vol. 8. Archived fromthe originalon 2017-09-02.Retrieved2019-08-14.
  23. ^Subhuti Dharmananda (July 2010)."DrynariaandDipsacus:yang tonifying herbs for bones, tendons, and brains ".Institute for Traditional Medicine.RetrievedJuly 30,2011.
  24. ^Karin Hilfiker (2008)."Improve livelihoods through market assessment and sustainable development of non-timber forest products (NTFPs) in two selected villages in the northern uplands of Vietnam"(PDF).Helvetas.Swiss Association for International Cooperation. Archived fromthe original(PDF)on September 29, 2011.RetrievedJuly 31,2011.
  25. ^Stuart Lindsay; David J. Middleton; Thaweesakdi Boonkerd; Somran Suddee (2009)."Towards a stable nomenclature for Thai ferns"(PDF).Thai Forest Bulletin (Botany)(37): 64–106. Archived fromthe original(PDF)on March 16, 2012.RetrievedJuly 31,2011.
  26. ^Eun-Kyung Jung (2007)."Antimicrobial Activity of Extract and Fractions fromDrynaria fortuneiAgainst Oral Bacteria "(PDF).Journal of Bacteriology and Virology.37(2): 61–68.doi:10.4167/jbv.2007.37.2.61.RetrievedJuly 28,2011.
  27. ^Christopher Hobbs; Kathi Keville (2007).Women's Herbs, Women's Health.Book Publishing Company. p. 287.ISBN978-1-57067-152-4.
  28. ^Godofredo Stuart."Pakpak lawin:Drynaria quercifoliaLinn ".Philippine Medicinal Plants.RetrievedJuly 27,2011.
  29. ^NSW Scientific Committee (October 4, 1998)."Drynaria rigidula(a fern) – endangered species listing ".Office of Environment & Heritage, NSW Government.RetrievedJuly 31,2011.
  30. ^abcTao Su; Frédéric M.B. Jacques; Yu-Sheng (Christopher) Liu; Jianying Xiang; Yaowu Xing; Yongjiang Huang & Zhekun Zhou (2010)."A newDrynaria(Polypodiaceae) from the Upper Pliocene of Southwest China "(PDF).Review of Palaeobotany and Palynology.164(2011). Elsevier: 521–527.doi:10.1016/j.revpalbo.2010.11.011.ISSN0034-6667.RetrievedJuly 30,2011.
  31. ^Thomas Janssen; Hans-Peter Kreier; Harald Schneider (2007)."Origin and diversification of African ferns with special emphasis on Polypodiaceae"(PDF).Brittonia.59(2). The New York Botanical Garden Press: 159–181.doi:10.1663/0007-196x(2007)59[159:oadoaf]2.0.co;2.S2CID23364840.Archived fromthe original(PDF)on March 19, 2012.RetrievedJuly 31,2011.
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