''Aglaomorpha'' is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...

s in the subfamily

Drynarioideae Drynarioideae is a subfamily of the fern family Polypodiaceae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The subfamily is also treated as the tribe Drynarieae within a very broadly defined family Polypodiaceae ''sen ...

of the family

Polypodiaceae Polypodiaceae is a family of ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family includes around 65 genera and an estimated 1,650 species and is placed in the order Polypodiales, suborder Polypodiineae. A broade ...

. The

Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group, or PPG, is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relatio ...

classification of 2016 (PPG I) uses this genus name, while other sources use ''Drynaria'' to include ''Aglaomorpha''. Species are commonly known as basket ferns. As circumscribed in PPG I, the genus contains around 50 species. Basket ferns are

epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...

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

medicinal plant Medicinal plants, also called medicinal herbs, have been discovered and used in traditional medicine practices since prehistoric times. Plants synthesize hundreds of chemical compounds for various functions, including defense and protection ag ...

Description

Basket ferns are characterized by the presence of two types of

frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...

s, fertile foliage fronds and sterile nest fronds. The dark green foliage fronds are large, long, with elongated stalks. They are deeply lobed or

pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...

, winged, and bear sori (structures producing and containing

spores In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...

) 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. The collected debris decompose into humus, providing the plants with nutrients it would otherwise not have received from being suspended above the ground. Both frond types grow from rhizomes typically anchored to a tree or a rock. The rhizomes of basket ferns are creeping and densely covered in brown scales.

Habitat and distribution

Basket ferns are

(growing on trees) or epipetric (growing on rocks). They can also sometimes be found in man-made structures like brick walls. They are found in wet tropical environments, usually in rainforests. Their native range extends from equatorial Africa to tropical South and East Asia, Southeast Asia, Australia, and Oceania.

Life cycle

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 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 ...

(the phase where the plant is known as a

prothallus A prothallus, or prothallium, (from Latin ''pro'' = forwards and Greek ''θαλλος'' (''thallos'') = twig) is usually the gametophyte stage in the life of a fern or other pteridophyte. Occasionally the term is also used to describe the young ...

). Gametophytes develop from spores released by mature sporophytes; while sporophytes, in turn, develop from the fusion of gametes produced by mature prothalli. 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

rhizoid Rhizoids are protuberances that extend from the lower epidermal cells of bryophytes and algae. They are similar in structure and function to the root hairs of vascular land plants. Similar structures are formed by some fungi. Rhizoids may be uni ...

s 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). The cordate prothallus are usually smaller with thinner midribs than that of other members of

. 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 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. Androecium is also ...

), and female ( archegonium)

gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...

s. The gametes fuse, forming the diploid sporophyte, the 'fern' part of the life cycle. ''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.

Ecology

''Aglaomorpha'', like some other genera of ferns (including ''

Polybotrya ''Polybotrya'' is a genus of ferns in the family Dryopteridaceae, subfamily Polybotryoideae Polybotryoideae is a subfamily of the fern family Dryopteridaceae. Genera The Pteridophyte Phylogeny Group classification of 2016 (PPG I) accepts the ...

'' and ''

Polypodium ''Polypodium'' is a genus of ferns in the family Polypodiaceae, subfamily Polypodioideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The genus is widely distributed throughout the world, with the highest ...

''), possess specialized nectar-secreting structures (

nectaries Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...

) on the bases of the frond lobes or the underside of the fronds. The produced nectar is rich in sugars and

amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha a ...

s. Their function may be to attract

ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...

s (or other organisms) for protection or for spore dispersal. They may also be simply excretory organs (

hydathode A hydathode is a type of pore, commonly found in angiosperms, that secretes water through pores in the epidermis or leaf margin, typically at the tip of a marginal tooth or serration. Hydathodes occur in the leaves of submerged aquatic plants suc ...

s), used for exuding surplus metabolic products. The ant species ''

Iridomyrmex cordatus ''Philidris cordata'' is a species of ant in the genus '' Philidris''. Described by Smith in 1859, the species is endemic to Indonesia and New Guinea. This species is a frequent inhabitant of the ant plant genera: '' Myrmecodia'' and ''Hydnophyt ...

'' is commonly associated with ''A. quercifolia'', in addition to other epiphytic plants. In Australia, '' Aglaomorpha rigidula'' serve as shelter for amethystine pythons ('' Morelia amethistina'') and scrub pythons ('' Morelia kinghorni''). As much as 81% of sightings of the snakes in one study were in large individuals of ''A. rigidula'' located about above the ground. Snakes seek shelter in ''A. rigidula'' more frequently during the colder seasons. In the 19th century,

Indigenous Australian Indigenous Australians or Australian First Nations are people with familial heritage from, and membership in, the ethnic groups that lived in Australia before British colonisation. They consist of two distinct groups: the Aboriginal peoples ...

s were documented by the

Norwegian Norwegian, Norwayan, or Norsk may refer to: *Something of, from, or related to Norway, a country in northwestern Europe * Norwegians, both a nation and an ethnic group native to Norway * Demographics of Norway *The Norwegian language, including ...

explorer Exploration refers to the historical practice of discovering remote lands. It is studied by geographers and historians. Two major eras of exploration occurred in human history: one of convergence, and one of divergence. The first, covering most ...

Carl Sofus Lumholtz to have hunted pythons regularly during the winter months by climbing up to individuals of ''A. rigidula''. The large rhizome mass of ''Aglaomorpha'' can also serve as growing substrates for other plants like the ribbon fern (''

Ophioglossum pendulum ''Ophioderma pendulum'' is sometimes known as the old-world adder's-tongue. In Malaysia, it is known as daun rambu. It is a fern in the family Ophioglossaceae, and is the type species of the genus ''Ophioderma''. It is most noteworthy for the len ...

''). 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.

Classification

Basket ferns are classified under the subfamily

of the family

. Species belonging to ''Aglaomorpha'' were once classified under the genus ''

'' (rockcap ferns), under the subgenus ''Aglaomorpha''.

Species

The following is the list of accepted species according to the ''Checklist of Ferns and Lycophytes of the World'' : *'' Aglaomorpha baronii'' (Diels) Hovenkamp & S. Linds. *'' Aglaomorpha bonii'' (Christ) Hovenkamp & S. Linds. *'' Aglaomorpha brooksii'' Copel. *'' Aglaomorpha cornucopia'' (Copel.) M. C. Roos *'' Aglaomorpha coronans'' (Wall. ex Mett.) Copel. *'' Aglaomorpha delavayi'' (Christ) Hovenkamp & S. Linds. *'' Aglaomorpha descensa'' (Copel.) Hovenkamp & S. Linds. *'' Aglaomorpha drynarioides'' (Hook.) M. C. Roos * ''Aglaomorpha'' × ''dumicola'' (Bostock) comb. ined. *'' Aglaomorpha fortunei'' (Kunze ex Mett.) Hovenkamp & S. Linds. *'' Aglaomorpha heraclea'' (Kunze) Copel. *'' Aglaomorpha hieronymi'' (Brause) Copel. *'' Aglaomorpha involuta'' (Alderw.) Hovenkamp & S. Linds. *'' Aglaomorpha latipinna'' (C. Chr.) M. C. Roos *'' Aglaomorpha laurentii'' (Christ ex De Wild. & Durand) Hovenkamp & S. Linds. *'' Aglaomorpha meeboldii'' (Rosenst.) comb. ined. *'' Aglaomorpha meyeniana'' Schott *'' Aglaomorpha mollis'' (Bedd.) Hovenkamp & S. Linds. *'' Aglaomorpha nectarifera'' (Becc. ex Baker) M. C. Roos *'' Aglaomorpha novoguineensis'' (Brause) C. Chr. *'' Aglaomorpha parishii'' (Bedd.) Hovenkamp & S. Linds. *'' Aglaomorpha parkinsonii'' (Baker) Parris & M. C. Roos *'' Aglaomorpha pilosa'' (J. Sm. ex Kunze) Copel. *'' Aglaomorpha pleuridioides'' (Mett.) Hovenkamp & S. Linds. *'' Aglaomorpha propinqua'' (Wall. ex Mett.) Hovenkamp & S. Linds. *''

Aglaomorpha quercifolia ''Aglaomorpha quercifolia'' (synonym ''Drynaria 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' ...

'' (L.) Hovenkamp & S. Linds. *'' Aglaomorpha rigidula'' (Sw.) Hovenkamp & S. Linds. *'' Aglaomorpha sagitta'' (Christ) Hovenkamp & S. Linds. *'' Aglaomorpha sparsisora'' (Desv.) Hovenkamp & S. Linds. *'' Aglaomorpha speciosa'' (Blume) M. C. Roos *'' Aglaomorpha splendens'' (J. Sm.) Copel. *'' Aglaomorpha tricuspis'' (Hook.) Hovenkamp & S. Linds. *'' Aglaomorpha volkensii'' (Hieron.) Hovenkamp & S. Linds. *'' Aglaomorpha willdenowii'' (Bory) Hovenkamp & S. Linds.

Traditional medicine

Extracts from the rhizomes of some ''Aglaomorpha'' species are used extensively in

traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...

. In China, Taiwan,

Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...

, Thailand, and Laos, the rhizomes of gu-sui-bu, ''Aglaomorpha fortunei'' (more frequently cited by Asian authors by its

illegitimate Legitimacy, in traditional Western common law, is the status of a child born to parents who are legally married to each other, and of a child conceived before the parents obtain a legal divorce. Conversely, ''illegitimacy'', also known as '' ...

synonym ''Drynaria fortunei''), are commonly used to treat bone injuries. Its common name literally means "mender of shattered bones" in Chinese. Another species, the oak-leaf fern (''Aglaomorpha quercifolia'') is used similarly in South Asia and

Maritime Southeast Asia Maritime Southeast Asia comprises the countries of Brunei, Indonesia, Malaysia, the Philippines, Singapore, and East Timor. Maritime Southeast Asia is sometimes also referred to as Island Southeast Asia, Insular Southeast Asia or Oceanic Sout ...

Conservation

Basket fern at Ravensbourne National Park 2817529

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. ''Aglaomorpha'' are also considered

endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and in ...

in some areas (like in

New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...

, Australia), due to threats of

habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...

and low population numbers.

Evolution and fossil record

Fossil species have not been transferred to ''Aglaomorpha'' in

PPG I The Pteridophyte Phylogeny Group, or PPG, is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes (lycophytes and ferns) that reflects knowledge about plant relati ...

, and are still recorded in the genus ''Drynaria''. In 2010, twelve well-preserved

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

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 The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58epoch In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...

(about 3.6 to 2.5 million years ago). The

fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

of drynarioids is not very well documented due to the typically poor preservation of

s recovered. Previous fossil species assigned to ''Drynaria'' include ''Drynaria astrostigma'', ''D. dura'', and ''D. tumulosa'' from the Cenomanian of the

Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...

; 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 Matoniaceae is one of the three families of ferns in the Gleicheniales order of the Polypodiopsida class. Fossil records reveal that Matoniaceae ferns were abundant during the Mesozoic era (about 250-million to 66-million years ago), during which ...

instead. Outside the genus, '' Protodrynaria takhtajani'' from the

Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...

- Oligocene boundary of

Kursk Oblast Kursk Oblast ( rus, Курская область, r=Kurskaya oblast, p=ˈkurskəjə ˈobləsʲtʲ) is a federal subject of Russia (an oblast). Its administrative center is the city of Kursk. As of the 2010 Census, Kursk Oblast has a populati ...

, 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 Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...

. These were later transferred to the living species '' Aglaomorpha heraclea''. It remains, as of 2011, the oldest known drynarioid.

References

{{Taxonbar, from=Q3020925, from2=Q4692851 Aglaomorpha Epiphytes Medicinal plants Fern genera