Aglaomorpha (plant)

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Aglaomorpha
Temporal range: Piacenzian to Present
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Drynaria quercifolia 'Pinto' kz01.jpg
Drynaria quercifolia at the Lincoln Park Conservatory
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Suborder: Polypodiineae
Family: Polypodiaceae
Subfamily: Drynarioideae
Genus: Aglaomorpha
Schott
Species

See text

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

Aglaomorpha is a genus of ferns in the subfamily Drynarioideae of the family Polypodiaceae. [1] The Pteridophyte Phylogeny Group classification of 2016 (PPG I) uses this genus name, [1] while other sources use Drynaria to include Aglaomorpha. [2] Species are commonly known as basket ferns. As circumscribed in PPG I, the genus contains around 50 species. [1]

Contents

Basket ferns are epiphytic or epipetric and are native to tropical Africa, South Asia, East Asia, Southeast Asia, Australia, and Oceania. Some species are economically important as medicinal plants.

Description

Aglaomorpha rigidula growing on rocks in Australia. The fertile foliage fronds are large and dark green, the smaller brown sterile nest fronds are clustered at their bases. Basket Fern.jpg
Aglaomorpha rigidula growing on rocks in Australia. The fertile foliage fronds are large and dark green, the smaller brown sterile nest fronds are clustered at their bases.
Sori on the abaxial surface of the foliage frond of Aglaomorpha quercifolia. Drynaria quercifolia (Oakleaf fern) sporangia.jpg
Sori on the abaxial surface of the foliage frond of Aglaomorpha quercifolia .

Basket ferns are characterized by the presence of two types of fronds, 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 or pinnate, winged, and bear sori (structures producing and containing spores) 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 into humus, providing the plants with nutrients it would otherwise not have received from being suspended above the ground. [4] [5]

Both frond types grow from rhizomes typically 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

Basket ferns are epiphytic (growing on trees) or epipetric (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 in rainforests. [3] Their native range extends from equatorial Africa to tropical South and East Asia, Southeast Asia, Australia, and Oceania. [9]

Life cycle

The "Drynaria" type prothallial germination. Drynaria type prothallial development.png
The "Drynaria" type prothallial germination.

Like other spore-bearing plants, Aglaomorpha exhibits metagenesis or the alternation of generations. One generation being the diploid multicellular sporophyte (the phase where the plant is most familiar), and the other being the haploid multicellular gametophyte (the phase where the plant is known as a prothallus). Gametophytes develop from spores released by mature sporophytes; while sporophytes, in turn, develop from the fusion of gametes produced by mature prothalli. [8] [10]

The synonym Drynaria lends its name to a certain type of prothallial germination, the 'Drynaria type', observed in several other ferns. In this type, the spores germinate into a germ filament composed of barrel-shaped chlorophyllous cells with one or more rhizoids at 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 an obconical meristematic 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 of Polypodiaceae. 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]

Aglaomorpha also naturally exhibits apospory, 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

Ants feeding on nectar secreted by the lower surface of a young Aglaomorpha quercifolia frond. Ants on Drynaria.jpg
Ants feeding on nectar secreted by the lower surface of a young Aglaomorpha quercifolia frond.

Aglaomorpha, like some other genera of ferns (including Polybotrya and Polypodium ), possess specialized nectar-secreting structures (nectaries) on the bases of the frond lobes or the underside of the fronds. The produced nectar is rich in sugars and amino acids. [13]

Their function may be to attract ants (or other organisms) for protection or for spore dispersal. They may also be simply excretory organs (hydathodes), used for exuding surplus metabolic products. [13] The ant species Iridomyrmex cordatus is commonly associated with A. quercifolia, in addition to other epiphytic plants. [14]

In Australia, Aglaomorpha rigidula serve 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 of A. rigidula located about 17–40 m (56–131 ft) above the ground. Snakes seek shelter in A. rigidula more frequently during the colder seasons. [16]

In the 19th century, Indigenous Australians were documented by the Norwegian explorer Carl Sofus Lumholtz to have hunted pythons regularly during the winter months by climbing up to individuals of A. rigidula. [17] The large rhizome mass of Aglaomorpha can 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 of Aglaomorpha are also fertile hosts to a large number of water-borne fungi. [19]

Classification

Basket ferns are classified under the subfamily Drynarioideae of the family Polypodiaceae. [1] Species belonging to Aglaomorpha were once classified under the genus Polypodium (rockcap ferns), under the subgenus Aglaomorpha. [20] [21]

Species

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

Traditional medicine

Extracts from the rhizomes of some Aglaomorpha species are used extensively in traditional medicine. [23] [24] In China, Taiwan, Vietnam, Thailand, and Laos, the rhizomes of gu-sui-bu, Aglaomorpha fortunei (more frequently cited by Asian authors by its illegitimate synonym Drynaria 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 and Maritime Southeast Asia. [10] [28]

Conservation

A basket fern in Ravensbourne National Park, Australia Basket fern at Ravensbourne National Park 2817529.jpg
A basket fern in Ravensbourne National Park, Australia

Species of Aglaomorpha commonly used in traditional medicine like A. roosii and A. quercifolia are in danger of being overexploited. None of the species are currently cultivated for the alternative medicine industry. [8] [10]

Aglaomorpha are also considered endangered in some areas (like in New South Wales, Australia), due to threats of habitat loss and low population numbers. [10] [29]

Evolution and fossil record

Fossil species have not been transferred to Aglaomorpha in PPG I, and are still recorded in the genus Drynaria. In 2010, twelve well-preserved fossil specimens were described from the Sanying Formation of the Yangjie coal mine of China. Named Drynaria callispora , it comes from the Piacenzian age of the Pliocene epoch (about 3.6 to 2.5 million years ago). [30]

The fossil record of drynarioids is not very well documented due to the typically poor preservation of fossils recovered. Previous fossil species assigned to Drynaria include Drynaria astrostigma, D. dura, and D. tumulosa from the Cenomanian of the Czech Republic; and D. durum, all assigned tentatively to the genus in 1899. The arrangement and type of their sori, however, indicate that they are members of the family Matoniaceae instead. [30]

Outside the genus, Protodrynaria takhtajani from the Eocene-Oligocene boundary of Kursk Oblast, Russia shows some affinities to Drynaria but only distantly. The only other reasonably convincing fossil remains of drynarioids aside from D. callispora was a specimen named Polypodium quercifolia recovered in 1985 from the Late Miocene (23.03 to 5.332 million years ago) of Palembang, Indonesia. These were later transferred to the living species Aglaomorpha heraclea . It remains, as of 2011, the oldest known drynarioid. [31] [30]

See also

Related Research Articles

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<i>Aglaomorpha fortunei</i> Species of plant

Aglaomorpha fortunei, commonly known as gu-sui-bu, is a species of basket fern of the family Polypodiaceae. The plant is native to Eastern Asia, including eastern China.

<i>Grammitis</i> Genus of plants

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<i>Lophosoria quadripinnata</i> Species of fern

Lophosoria quadripinnata(J.F.Gmel.) 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. It is found in the Americas spanning from Cuba and Mexico to Chile. In Chile it is present in the area between Talca and Aysén including Juan Fernández Islands. In Argentina it grows only in the humid valleys of western Neuquén and Río Negro Province. Diamondleaf fern is a common name. In Spanish it is known as 'ampe' 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 and even though the rhizome 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 rhizome 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 in southern Gondwana according to fossil remains found in Antarctica. The species is well known as an ornamental plant.

<i>Zealandia pustulata</i> Species of fern

Zealandia pustulata is a species of fern native to eastern Australia and New Zealand. It is commonly referred to as kangaroo fern because of its mature leaves tend to resemble the shape of a kangaroo foot. It is also referred to as hound's tongue, and as kōwaowao and pāraharaha in Māori.

<i>Parablechnum wattsii</i> Species of plant

Parablechnum wattsii, synonym Blechnum wattsii, is a common terrestrial fern growing in rainforest and open forest. It is often seen near creeks in much of south eastern Australia, including Victoria, Tasmania, South Australia, New South Wales and Queensland. The specific epithet wattsii honours William Walter Watts (1856-1920). Watts was considered an authority on mosses and ferns and has more than 30 species named for him. Common names by which the species may be called are hard water fern - from its stiff leathery fronds, leech fern - as forest workers often encounter leaches while working in clusters of these ferns, hard hill fern - from the fern's habit and habitat, and red cabbage fern - from the bronze-pink colour of the young fronds resembling cooked red cabbage.

<i>Hymenophyllum australe</i> Species of fern

Hymenophyllum australe, commonly known as austral filmy fern, is a relatively large rupestral and epiphytic fern, indigenous to eastern Australia and New Zealand. It belongs to the unique Hymenophyllum genus, which are characterised by their thin membranous fronds that are seldom more than one cell thick, with the exception of regions over and around veins. Hymenophyllum australe is distinctive in that the fronds are typically thicker than other Hymenophyllum species, often being up to 2-3 cells thick.

<i>Pyrrosia</i> Genus of ferns

Pyrrosia is a genus of about 100 fern species in the polypod family, Polypodiaceae. Like other species in Polypodiaceae, the species of Pyrrosia are generally epiphytic on trees or rocks, a few species are terrestrial. The Latin name of Pyrrosia comes from the Greek pyrrhos (red), which refers to its leaves that are red due to the sporangia.

<i>Aglaomorpha quercifolia</i> Species of fern

Aglaomorpha quercifolia, commonly known as the oakleaf fern or oakleaf basket fern, is a species of Aglaomorpha in the family Polypodiaceae. Other common names for the fern are pakpak lawin, gurar, koi hin, ashvakatri, kabkab, kabkaban, or uphatkarul.

<i>Platycerium grande</i> Species of fern

Platycerium grande, the giant staghorn fern, capa de leon, and dapong repolyo, is a species of epiphytic fern in the family Polypodiaceae. It is one of the two staghorn ferns native to the Philippines, along with P. coronarium, and is endemic to the island of Mindanao, in the provinces of Zamboanga, Lanao and Davao. P. grande is often collected from the forests and sold as a highly prized ornamental plant. Due to overcollection and the difficulty of the spores to germinate under natural conditions, in vitro technique is necessary to ensure mass production of this plant species. The local government categorized it as critically endangered species.

<i>Aglaomorpha coronans</i> Species of plant

Aglaomorpha coronans is a species of tropical fern widely distributed throughout Southeast Asia, from Nepal to Peninsular Malaysia. Like other Aglaomorpha, it is an epiphyte and its scientific name refers to the circular, crown-like growth habit. One of the species of basket fern more common in cultivation as a houseplant, it has as thick rhizome that soaks up moisture and fronds that are up to 1.8 metres long and 40 centimetres (16 in) wide.

<i>Aglaomorpha rigidula</i> Species of plant

Aglaomorpha rigidula is a species of tropical fern in genus Aglaomorpha widely distributed throughout Southeast Asia, from Southwest China to Queensland and New South Wales in Australia. Growing as an epiphyte or lithophyte, its green fronds are up to 1.22 metres in length. This fern develops two types of fronds, some of which near the base of the plant are shorter, sterile, and rust-colored which form a basket-like shape that trap moisture and organic components that nourish it. Especially in colder weather, this basket acts as prime habitat for animals such as the Amethystine python.

<i>Aglaomorpha willdenowii</i> Species of plant

Aglaomorpha willdenowii is a species of subtropical fern native to the islands of Comoros, Madagascar, and Mauritius. Drynarioid ferns are largely native to Asia and Oceania, but A. willdenowii is the only species of fern in the genus native to Madagascar. Like other Aglaomorpha, its caudex is covered in dry, scale-like sterile fronds and fine hairs, while the larger green fronds are the fertile ones bearing spores.

References

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  3. 1 2 Russell Cumming; Jan Sked (August 21, 2006). "Drynaria & Platycerium: Interesting Fern Genera". Society for Growing Australian Plants. Archived from the original on August 23, 2011. Retrieved July 27, 2011.
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  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. ISBN   978-0-88192-422-0.
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  13. 1 2 Suzanne Koptur; Alan R. Smith; Irene Baker (1982). "Nectaries in Some Neotropical Species of Polypodium (Polypodiaceae): Preliminary Observations and Analyses" (PDF). Biotropica. 14 (2). Association for Tropical Biology: 108–113. Bibcode:1982Biotr..14..108K. doi:10.2307/2387739. JSTOR   2387739. Archived from the original (PDF) on April 6, 2012. Retrieved July 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. ISBN   978-0-226-71348-9.
  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 from the original (PDF) on March 29, 2011. Retrieved July 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. Retrieved July 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 from the original (PDF) on October 2, 2011. Retrieved July 30, 2011.
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  22. ↑ Hassler, Michael & Schmitt, Bernd (June 2019). "Aglaomorpha". Checklist of Ferns and Lycophytes of the World. Vol. 8. Archived from the original on 2017-09-02. Retrieved 2019-08-14.
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  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 from the original (PDF) on September 29, 2011. Retrieved July 31, 2011.
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  26. ↑ Eun-Kyung Jung (2007). "Antimicrobial Activity of Extract and Fractions from Drynaria fortunei Against Oral Bacteria" (PDF). Journal of Bacteriology and Virology. 37 (2): 61–68. doi:10.4167/jbv.2007.37.2.61 . Retrieved July 28, 2011.
  27. ↑ Christopher Hobbs; Kathi Keville (2007). Women's Herbs, Women's Health. Book Publishing Company. p. 287. ISBN   978-1-57067-152-4.
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  29. ↑ NSW Scientific Committee (October 4, 1998). "Drynaria rigidula (a fern) – endangered species listing". Office of Environment & Heritage, NSW Government. Retrieved July 31, 2011.
  30. 1 2 3 Tao Su; Frédéric M.B. Jacques; Yu-Sheng (Christopher) Liu; Jianying Xiang; Yaowu Xing; Yongjiang Huang & Zhekun Zhou (2010). "A new Drynaria (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. ISSN   0034-6667 . Retrieved July 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. S2CID   23364840. Archived from the original (PDF) on March 19, 2012. Retrieved July 31, 2011.
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