Drynaria
Encyclopedia
Drynaria, commonly known as basket ferns, is a genus
of fern
s in the family Polypodiaceae
. It contains around 16 species and one natural hybrid.
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.
s, fertile foliage fronds and sterile nest fronds. The dark green foliage fronds are large, 2 – 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. 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 Drynaria are creeping and densely covered in brown scales.
(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 rainforest
s. Their native range extends from equatorial Africa to tropical South
and East Asia, Southeast Asia, Australia, and Oceania.
, Drynaria 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 spore
s released by mature sporophytes; while sporophytes, in turn, develop from the fusion of gametes produced by mature prothalli.
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
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 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
) gamete
s. The gametes fuse, forming the diploid sporophyte, the 'fern' part of the life cycle.
Drynaria 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.
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 sugar
s and amino acid
s.
Their function may be to attract ant
s (or other organisms) for protection or for spore
dispersal. They may also be simply excretory organs (hydathode
s), used for exuding surplus metabolic products. The ant species Iridomyrmex cordatus is commonly associated with D. quercifolia, in addition to other epiphytic plants.
In Australia, Drynaria 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 D. rigidula located about 17 – above the ground. Snakes seek shelter in D. rigidula more frequently during the colder seasons.
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 D. rigidula.
The large rhizome mass of Drynaria can also serve as growing substrates for other plants like the ribbon fern (Ophioglossum pendulum). Due to their ability to preserve moisture and persistence even after death, the nest leaves of Drynaria are also fertile hosts to a large number of water-borne fungi
.
Drynarieae
, subfamily Polypodioideae
, of the family Polypodiaceae
. Species belonging to Drynaria were once classified under the genus Polypodium
(rockcap ferns), under the subgenus
Drynaria.
Drynaria callispora Su, Zhou et Liu
. In China, Taiwan, Vietnam
, Thailand, and Laos
, the rhizomes of Gu-Sui-Bu Drynaria roosii (more frequently cited by Asian authors by its 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 (Drynaria quercifolia) is used similarly in South Asia and Maritime Southeast Asia
.
studies have been conducted into the properties of Drynaria. In agreement with their use in traditional medicine, several studies have shown that basket ferns (D. roosii in particular) are effective in preventing resorption
of bone cell
s and osteoporosis
, increases bone density, and have therapeutic effects on bone healing. They have also been shown to possess a wide range of antimicrobial activity.
industry.
Drynaria are also considered endangered in some areas (like in New South Wales
, Australia), due to threats of habitat loss and low population numbers.
specimens of Drynaria 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).
The fossil record of Drynaria and drynarioids is not very well documented due to the typically poor preservation of fossil
s 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.
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.
Below is a reconstruction of the evolution
of species of Drynaria and closely related genera Aglaomorpha and Selliguea
based on combined morphological and molecular phylogenetic data (Janssen & Schneider, 2005). Drynaria is likely to be paraphyletic
, in contrast to Aglaomorpha, another genus which possess specialized humus-collecting structures.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of fern
Fern
A fern is any one of a group of about 12,000 species of plants belonging to the botanical group known as Pteridophyta. Unlike mosses, they have xylem and phloem . They have stems, leaves, and roots like other vascular plants...
s in the family Polypodiaceae
Polypodiaceae
Polypodiaceae is a family of polypod ferns, which includes more than 60 genera divided into several tribes and containing around 1,000 species. Nearly all are epiphytes, but some are terrestrial.-Description:...
. It contains around 16 species and one natural hybrid.
Basket ferns are epiphytic or epipetric
Epipetric
An epipetric plant, is one that is found growing on rocks. Many ferns fall into this category of plant habitat, including Asplenium . Epipetric plants are called Lithophytes....
and are native to tropical Africa, South Asia, East Asia, Southeast Asia, Australia, and Oceania. Some species are economically important as medicinal plants.
Description
Basket ferns are characterized by the presence of two types of frondFrond
The term frond refers to 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...
s, fertile foliage fronds and sterile nest fronds. The dark green foliage fronds are large, 2 – long, with elongated stalks. They are deeply lobed or pinnate
Pinnate
Pinnate is a term used to describe feather-like or multi-divided features arising from both sides of a common axis in plant or animal structures, and comes from the Latin word pinna meaning "feather", "wing", or "fin". A similar term is pectinate, which refers to a comb-like arrangement of parts...
, 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. 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 Drynaria are creeping and densely covered in brown scales.
Habitat and distribution
Basket ferns are epiphytic (growing on trees) or epipetricEpipetric
An epipetric plant, is one that is found growing on rocks. Many ferns fall into this category of plant habitat, including Asplenium . Epipetric plants are called Lithophytes....
(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 rainforest
Rainforest
Rainforests are forests characterized by high rainfall, with definitions based on a minimum normal annual rainfall of 1750-2000 mm...
s. Their native range extends from equatorial Africa to tropical South
South Asia
South Asia, also known as Southern Asia, is the southern region of the Asian continent, which comprises the sub-Himalayan countries and, for some authorities , also includes the adjoining countries to the west and the east...
and East Asia, Southeast Asia, Australia, and Oceania.
Life cycle
Like other spore-bearing plantsPteridophyte
The pteridophytes are vascular plants that produce neither flowers nor seeds, and are hence called vascular cryptogams. Instead, they reproduce and disperse only via spores. Pteridophytes include horsetails, ferns, club mosses, and quillworts...
, Drynaria exhibits metagenesis or the alternation of generations
Alternation of generations
Alternation of generations is a term primarily used in describing the life cycle of plants . A multicellular sporophyte, which is diploid with 2N paired chromosomes , alternates with a multicellular gametophyte, which is haploid with N unpaired chromosomes...
. One generation being the diploid multicellular sporophyte
Sporophyte
All land plants, and some algae, have life cycles in which a haploid gametophyte generation alternates with a diploid sporophyte, the generation of a plant or algae that has a double set of chromosomes. A multicellular sporophyte generation or phase is present in the life cycle of all land plants...
(the phase where the plant is most familiar), and the other being the haploid multicellular gametophyte
Gametophyte
A gametophyte is the haploid, multicellular phase of plants and algae that undergo alternation of generations, with each of its cells containing only a single set of chromosomes....
(the phase where the plant is known as a prothallus). Gametophytes develop from spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
s released by mature sporophytes; while sporophytes, in turn, develop from the fusion of gametes produced by mature prothalli.
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
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...
cells with one or more rhizoid
Rhizoid
Rhizoids are thread-like growths from the base or bottom of a plant, found mainly in lower groups such as algae, fungi, bryophytes and pteridophytes, that function like roots of higher plants ....
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 Polypodiaceae
Polypodiaceae
Polypodiaceae is a family of polypod ferns, which includes more than 60 genera divided into several tribes and containing around 1,000 species. Nearly all are epiphytes, but some are terrestrial.-Description:...
. 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
Antheridium
An antheridium or antherida is a haploid structure or organ producing and containing male gametes . It is present in the gametophyte phase of lower plants like mosses and ferns, and also in the primitive vascular psilotophytes...
), and female (archegonium
Archegonium
An archegonium , from the ancient Greek ἀρχή and γόνος , is a multicellular structure or organ of the gametophyte phase of certain plants, producing and containing the ovum or female gamete. The archegonium has a long neck canal and a swollen base...
) gamete
Gamete
A gamete is a cell that fuses with another cell during fertilization in organisms that reproduce sexually...
s. The gametes fuse, forming the diploid sporophyte, the 'fern' part of the life cycle.
Drynaria also naturally exhibits apospory
Vegetative reproduction
Vegetative reproduction is a form of asexual reproduction in plants. It is a process by which new individuals arise without production of seeds or spores...
, 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
Drynaria, like some other generaGenera
Genera is a commercial operating system and development environment for Lisp machines developed by Symbolics. It is essentially a fork of an earlier operating system originating on the MIT AI Lab's Lisp machines which Symbolics had used in common with LMI and Texas Instruments...
of ferns (including Polybotrya
Polybotrya
Polybotrya is a genus of ferns in family Dryopteridaceae.Species include:* Polybotrya andina, C.Chr....
and Polypodium
Polypodium
Polypodium is a genus of 75–100 species of true ferns, widely distributed throughout the world, with the highest species diversity in the tropics. The name is derived from Ancient Greek poly "many" + podion "little foot", on account of the foot-like appearance of the rhizome and its branches...
), 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 sugar
Sucrose
Sucrose is the organic compound commonly known as table sugar and sometimes called saccharose. A white, odorless, crystalline powder with a sweet taste, it is best known for its role in human nutrition. The molecule is a disaccharide composed of glucose and fructose with the molecular formula...
s and amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...
s.
Their function may be to attract ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s (or other organisms) for protection or for spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
dispersal. They may also be simply excretory organs (hydathode
Hydathode
A hydathode is a type of secretory tissue in leaves, usually of Angiosperms, that secretes water through pores in the epidermis or margin of leaves, typically at the tip of a marginal tooth or serration. They probably evolved from modified stomata...
s), used for exuding surplus metabolic products. The ant species Iridomyrmex cordatus is commonly associated with D. quercifolia, in addition to other epiphytic plants.
In Australia, Drynaria 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 D. rigidula located about 17 – above the ground. Snakes seek shelter in D. rigidula more frequently during the colder seasons.
In the 19th century, Indigenous Australians were documented by the Norwegian explorer Carl Sofus Lumholtz
Carl Sofus Lumholtz
Carl Sofus Lumholtz was a Norwegian explorer and ethnographer, best known for his meticulous field research and ethnographic publications on indigenous cultures of Australia and Mesoamerican central Mexico.-Biography:...
to have hunted pythons regularly during the winter months by climbing up to individuals of D. rigidula.
The large rhizome mass of Drynaria can also serve as growing substrates for other plants like the ribbon fern (Ophioglossum pendulum). Due to their ability to preserve moisture and persistence even after death, the nest leaves of Drynaria are also fertile hosts to a large number of water-borne fungi
Hyphomycetes
Hyphomycetes is an obsolete class of fungi in the equally obsolete phylum Deuteromycota that lack fruiting bodies. Most hyphomycetes have now been assigned to the Ascomycota, mainly as a result of DNA sequencing, but many remain unassigned...
.
Classification
Basket ferns are classified under the tribeTribe (biology)
In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...
Drynarieae
Drynarieae
Drynarieae is a tribe of ferns belonging to the subfamily Polypodioideae of the family Polypodiaceae. Its members are characterized by dimorphic fronds. It contains around 35 species and two genera:* Aglaomorpha* Drynaria...
, subfamily Polypodioideae
Polypodioideae
Polypodioideae is a subfamily belonging to the family Polypodiaceae. They possess scales and/or hairs. Never stellate....
, of the family Polypodiaceae
Polypodiaceae
Polypodiaceae is a family of polypod ferns, which includes more than 60 genera divided into several tribes and containing around 1,000 species. Nearly all are epiphytes, but some are terrestrial.-Description:...
. Species belonging to Drynaria were once classified under the genus Polypodium
Polypodium
Polypodium is a genus of 75–100 species of true ferns, widely distributed throughout the world, with the highest species diversity in the tropics. The name is derived from Ancient Greek poly "many" + podion "little foot", on account of the foot-like appearance of the rhizome and its branches...
(rockcap ferns), under the subgenus
Subgenus
In biology, a subgenus is a taxonomic rank directly below genus.In zoology, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the generic name and the specific epithet: e.g. the Tiger Cowry of the Indo-Pacific, Cypraea tigris Linnaeus, which...
Drynaria.
Species
The following is the list of accepted species classified under the genus. Species marked with are extinct.- Drynaria bonii H. Christ
-
- Native to India, Southeast Asia, and China. One of the smallest species of Drynaria, it can be distinguished by rounded nest fronds. The foliage fronds are pinnatesectPinnatePinnate is a term used to describe feather-like or multi-divided features arising from both sides of a common axis in plant or animal structures, and comes from the Latin word pinna meaning "feather", "wing", or "fin". A similar term is pectinate, which refers to a comb-like arrangement of parts...
with few widely separated lobes. The sori are arranged in two irregular rows between the secondary veins.
- Native to India, Southeast Asia, and China. One of the smallest species of Drynaria, it can be distinguished by rounded nest fronds. The foliage fronds are pinnatesect
Drynaria callispora Su, Zhou et Liu
-
- Extinct species from the PiacenzianPiacenzianThe Piacenzian is in the international geologic timescale the upper stage or latest age of the Pliocene. It spans the time between 3.6 ± 0.005 Ma and 2.588 ± 0.005 Ma...
age of China. See section below.
- Extinct species from the Piacenzian
- Drynaria delavayi H. Christ
-
- Native to Tibet, China, Bhutan, and Myanmar. Nest fronds pinnatilobed with wavy margins. Foliage fronds are deeply pinnatifid, with 7 to 9 pinnae. Sori are large in comparison to the size of the lobes and arranged in two rows on both sides of the main rib and in between the secondary veins. Closely related to D. sinica.
- Drynaria descensa Copel.
-
- Native to the PhilippinesPhilippinesThe Philippines , officially known as the Republic of the Philippines , is a country in Southeast Asia in the western Pacific Ocean. To its north across the Luzon Strait lies Taiwan. West across the South China Sea sits Vietnam...
. Similar to D. bonii but has smaller nest fronds, less than 2.5 cm (0.984251968503937 in) wide, with slightly scalloped margins. The foliage fronds are wavy.
- Native to the Philippines
- Drynaria involuta Alderw.
-
- Native to BorneoBorneoBorneo is the third largest island in the world and is located north of Java Island, Indonesia, at the geographic centre of Maritime Southeast Asia....
and JavaJavaJava is an island of Indonesia. With a population of 135 million , it is the world's most populous island, and one of the most densely populated regions in the world. It is home to 60% of Indonesia's population. The Indonesian capital city, Jakarta, is in west Java...
.
- Native to Borneo
- Drynaria laurentii (H. Christ) Hieron.
-
- Commonly known as ibandibandi. It is native to equatorial Africa, from CameroonCameroonCameroon, officially the Republic of Cameroon , is a country in west Central Africa. It is bordered by Nigeria to the west; Chad to the northeast; the Central African Republic to the east; and Equatorial Guinea, Gabon, and the Republic of the Congo to the south. Cameroon's coastline lies on the...
to the Democratic Republic of the CongoDemocratic Republic of the CongoThe Democratic Republic of the Congo is a state located in Central Africa. It is the second largest country in Africa by area and the eleventh largest in the world...
.
- Commonly known as ibandibandi. It is native to equatorial Africa, from Cameroon
- Drynaria mollis Bedd.
-
- Native to TibetTibetTibet is a plateau region in Asia, north-east of the Himalayas. It is the traditional homeland of the Tibetan people as well as some other ethnic groups such as Monpas, Qiang, and Lhobas, and is now also inhabited by considerable numbers of Han and Hui people...
, Nepal, BhutanBhutanBhutan , officially the Kingdom of Bhutan, is a landlocked state in South Asia, located at the eastern end of the Himalayas and bordered to the south, east and west by the Republic of India and to the north by the People's Republic of China...
, and northern IndiaIndiaIndia , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...
. Characterized by deeply lobed pinnatifid nest fronds. Foliage fronds are deeply pinnatifid. The sori are small and arranged in one row along the primary vein.
- Native to Tibet
- Drynaria parishii (Bedd.) Bedd.
-
- Native to China and IndochinaIndochinaThe Indochinese peninsula, is a region in Southeast Asia. It lies roughly southwest of China, and east of India. The name has its origins in the French, Indochine, as a combination of the names of "China" and "India", and was adopted when French colonizers in Vietnam began expanding their territory...
. Commonly known as xiao hu jue ('small oak fern') and tắc kè đá parít.
- Native to China and Indochina
- Drynaria propinqua (Wall. ex Mett.) Bedd.
-
- Native to Nepal, Bhutan, India, Southeast Asia, and China. A medium-sized species, it can be distinguished by deeply pinnatifidPinnatePinnate is a term used to describe feather-like or multi-divided features arising from both sides of a common axis in plant or animal structures, and comes from the Latin word pinna meaning "feather", "wing", or "fin". A similar term is pectinate, which refers to a comb-like arrangement of parts...
on both the foliage and nest fronds, with lobes that taper to a sharp point. The sori are arranged in two rows on each side of the primary rib of each lobe and in between the secondary veins.
- Native to Nepal, Bhutan, India, Southeast Asia, and China. A medium-sized species, it can be distinguished by deeply pinnatifid
- Drynaria quercifolia (L.) J. Sm.
-
- Commonly known as the oak-leaf fern, pakpak lawin, gurar, koi hin, ashvakatri, or uphatkarul. It is native to India, Southeast Asia, Malaysia, Indonesia, the Philippines, New GuineaNew GuineaNew Guinea is the world's second largest island, after Greenland, covering a land area of 786,000 km2. Located in the southwest Pacific Ocean, it lies geographically to the east of the Malay Archipelago, with which it is sometimes included as part of a greater Indo-Australian Archipelago...
, and Australia. It is a large species with deeply pinnatifid foliage fronds. The nest fronds resemble the leaves of oakOakAn oak is a tree or shrub in the genus Quercus , of which about 600 species exist. "Oak" may also appear in the names of species in related genera, notably Lithocarpus...
s, hence the common name. The sori are either scattered or arranged in two regular rows in between the secondary veins.
- Commonly known as the oak-leaf fern, pakpak lawin, gurar, koi hin, ashvakatri, or uphatkarul. It is native to India, Southeast Asia, Malaysia, Indonesia, the Philippines, New Guinea
- Drynaria rigidula (Sw.) Bedd.
-
- Commonly known simply as the basket fern. It is native to Southeast Asia, China, Malaysia, Indonesia, the Philippines, Australia, New Guinea, and the southeastern Pacific IslandsPacific IslandsThe Pacific Islands comprise 20,000 to 30,000 islands in the Pacific Ocean. The islands are also sometimes collectively called Oceania, although Oceania is sometimes defined as also including Australasia and the Malay Archipelago....
. Easily recognizable by being the only species in the genus with 1-pinnate foliage fronds. The lobes are narrow, taper to a sharp point, and are narrowly-winged at their attachment to the rachisRachisRachis is a biological term for a main axis or "shaft".-In zoology:In vertebrates a rachis can refer to the series of articulated vertebrae, which encase the spinal cord. In this case the rachis usually form the supporting axis of the body and is then called the spine or vertebral column...
. Sori are arranged in two rows along the primary vein of each lobe. Includes the cultivar Drynaria rigidula 'Whitei'.
- Commonly known simply as the basket fern. It is native to Southeast Asia, China, Malaysia, Indonesia, the Philippines, Australia, New Guinea, and the southeastern Pacific Islands
- Drynaria roosii Nakaike
-
- Commonly known as Gu-Sui-Bu. It is native to East Asia. Sterile nest fronds are rounded shallowly-lobed reddish-brown fronds overlapping each other. The fertile fronds are larger and deeply lobed. They bear 1 to 3 sori arranged on both sides of the central rib. Formerly known as Drynaria fortunei.
- Drynaria sinica Diels
-
- Commonly known as rgyal po re ral. Native to TibetTibetTibet is a plateau region in Asia, north-east of the Himalayas. It is the traditional homeland of the Tibetan people as well as some other ethnic groups such as Monpas, Qiang, and Lhobas, and is now also inhabited by considerable numbers of Han and Hui people...
.
- Commonly known as rgyal po re ral. Native to Tibet
- Drynaria sparsisora (Desv.) T. Moore
-
- Commonly known as simbar layangan. Native to Southeast Asia, Malaysia, Indonesia, New Guinea, and Australia. A large species, it is similar to D. quercifolia but has smaller and stiffer foliage fronds. The sori are also smaller and arranged irregularly.
- Drynaria volkensii Hieron.
-
- Native to tropical Africa including Cameroon, RwandaRwandaRwanda or , officially the Republic of Rwanda , is a country in central and eastern Africa with a population of approximately 11.4 million . Rwanda is located a few degrees south of the Equator, and is bordered by Uganda, Tanzania, Burundi and the Democratic Republic of the Congo...
, Kenya, BurundiBurundiBurundi , officially the Republic of Burundi , is a landlocked country in the Great Lakes region of Eastern Africa bordered by Rwanda to the north, Tanzania to the east and south, and the Democratic Republic of the Congo to the west. Its capital is Bujumbura...
, Tanzania, SudanSudanSudan , officially the Republic of the Sudan , is a country in North Africa, sometimes considered part of the Middle East politically. It is bordered by Egypt to the north, the Red Sea to the northeast, Eritrea and Ethiopia to the east, South Sudan to the south, the Central African Republic to the...
, Ethiopia, SomaliaSomaliaSomalia , officially the Somali Republic and formerly known as the Somali Democratic Republic under Socialist rule, is a country located in the Horn of Africa. Since the outbreak of the Somali Civil War in 1991 there has been no central government control over most of the country's territory...
, Zambia, MalawiMalawiThe Republic of Malawi is a landlocked country in southeast Africa that was formerly known as Nyasaland. It is bordered by Zambia to the northwest, Tanzania to the northeast, and Mozambique on the east, south and west. The country is separated from Tanzania and Mozambique by Lake Malawi. Its size...
, and MozambiqueMozambiqueMozambique, officially the Republic of Mozambique , is a country in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi and Zambia to the northwest, Zimbabwe to the west and Swaziland and South Africa to the southwest...
. Nest fronds are pinnatifid with acute to obtuse narrow lobes. Foliage fronds are pinnatifid and deeply and narrowly lobed. Sori arranged close to the main rib, about 3 mm (0.118110236220472 in) apart.
- Native to tropical Africa including Cameroon, Rwanda
- Drynaria willdenowii (Bory) T. Moore
-
- Native to MauritiusMauritiusMauritius , officially the Republic of Mauritius is an island nation off the southeast coast of the African continent in the southwest Indian Ocean, about east of Madagascar...
and MadagascarMadagascarThe Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...
.
- Native to Mauritius
- Drynaria × dumicola Bostock
-
- A natural hybrid of D. sparsisora and D. rigidula Found in QueenslandQueenslandQueensland is a state of Australia, occupying the north-eastern section of the mainland continent. It is bordered by the Northern Territory, South Australia and New South Wales to the west, south-west and south respectively. To the east, Queensland is bordered by the Coral Sea and Pacific Ocean...
, Australia.
- A natural hybrid of D. sparsisora and D. rigidula Found in Queensland
Traditional medicine
Extracts from the rhizomes of some Drynaria species are used extensively in traditional medicineTraditional medicine
Traditional medicine comprises unscientific knowledge systems that developed over generations within various societies before the era of modern medicine...
. In China, Taiwan, Vietnam
Vietnam
Vietnam – sometimes spelled Viet Nam , officially the Socialist Republic of Vietnam – is the easternmost country on the Indochina Peninsula in Southeast Asia. It is bordered by China to the north, Laos to the northwest, Cambodia to the southwest, and the South China Sea –...
, Thailand, and Laos
Laos
Laos Lao: ສາທາລະນະລັດ ປະຊາທິປະໄຕ ປະຊາຊົນລາວ Sathalanalat Paxathipatai Paxaxon Lao, officially the Lao People's Democratic Republic, is a landlocked country in Southeast Asia, bordered by Burma and China to the northwest, Vietnam to the east, Cambodia to the south and Thailand to the west...
, the rhizomes of Gu-Sui-Bu Drynaria roosii (more frequently cited by Asian authors by its 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 (Drynaria quercifolia) is used similarly in South Asia and Maritime Southeast Asia
Maritime Southeast Asia
Maritime Southeast Asia refers to the maritime region of Southeast Asia as opposed to mainland Southeast Asia and includes the modern countries of Indonesia, Malaysia, the Philippines, Brunei, East Timor and Singapore....
.
Medical research
Numerous ethnopharmacologicalEthnopharmacology
Ethnopharmacology is the scientific study of ethnic groups and their use of drugs.Ethnopharmacology is distinctly linked to plant use, botany, as this is the main delivery of pharmaceuticals. It is also often associated with ethnopharmacy...
studies have been conducted into the properties of Drynaria. In agreement with their use in traditional medicine, several studies have shown that basket ferns (D. roosii in particular) are effective in preventing resorption
Bone resorption
Bone resorption is the process by which osteoclasts break down bone and release the minerals, resulting in a transfer of calcium from bone fluid to the blood....
of bone cell
Bone cell
There are five types of bone cells.*Osteoblasts - These are commonly called bone-forming cells. They secrete osteoid, which forms the bone matrix. They also begin mieralization, and are unable to divide....
s and osteoporosis
Osteoporosis
Osteoporosis is a disease of bones that leads to an increased risk of fracture. In osteoporosis the bone mineral density is reduced, bone microarchitecture is deteriorating, and the amount and variety of proteins in bone is altered...
, increases bone density, and have therapeutic effects on bone healing. They have also been shown to possess a wide range of antimicrobial activity.
Conservation
Species of Drynaria commonly used in traditional medicine like D. roosii and D. quercifolia are in danger of being overexploited. None of the species are currently cultivated for the alternative medicineAlternative medicine
Alternative medicine is any healing practice, "that does not fall within the realm of conventional medicine." It is based on historical or cultural traditions, rather than on scientific evidence....
industry.
Drynaria are also considered endangered in some areas (like in New South Wales
New South Wales
New South Wales is a state of :Australia, located in the east of the country. It is bordered by Queensland, Victoria and South Australia to the north, south and west respectively. To the east, the state is bordered by the Tasman Sea, which forms part of the Pacific Ocean. New South Wales...
, Australia), due to threats of habitat loss and low population numbers.
Evolution and fossil record
In 2010, twelve well-preserved fossilFossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
specimens of Drynaria were described from the Sanying Formation of the Yangjie coal mine of China. Named Drynaria callispora, it comes from the Piacenzian
Piacenzian
The Piacenzian is in the international geologic timescale the upper stage or latest age of the Pliocene. It spans the time between 3.6 ± 0.005 Ma and 2.588 ± 0.005 Ma...
age of the Pliocene
Pliocene
The Pliocene Epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present. It is the second and youngest epoch of the Neogene Period in the Cenozoic Era. The Pliocene follows the Miocene Epoch and is followed by the Pleistocene Epoch...
epoch
Epoch (geology)
An epoch is a subdivision of the geologic timescale based on rock layering. In order, the higher subdivisions are periods, eras and eons. We are currently living in the Holocene epoch...
(about 3.6 to 2.5 million years ago).
The fossil record of Drynaria and drynarioids is not very well documented due to the typically poor preservation of fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
s recovered. Previous fossil species assigned to Drynaria include Drynaria astrostigma, D. dura, and D. tumulosa from the Cenomanian
Cenomanian
The Cenomanian is, in the ICS' geological timescale the oldest or earliest age of the Late Cretaceous epoch or the lowest stage of the Upper Cretaceous series. An age is a unit of geochronology: it is a unit of time; the stage is a unit in the stratigraphic column deposited during the corresponding...
of the Czech Republic
Czech Republic
The Czech Republic is a landlocked country in Central Europe. The country is bordered by Poland to the northeast, Slovakia to the east, Austria to the south, and Germany to the west and northwest....
; 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.
Outside the genus, Protodrynaria takhtajani from the Eocene
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...
-Oligocene
Oligocene
The Oligocene is a geologic epoch of the Paleogene Period and extends from about 34 million to 23 million years before the present . As with other older geologic periods, the rock beds that define the period are well identified but the exact dates of the start and end of the period are slightly...
boundary of Kursk Oblast
Kursk Oblast
Kursk Oblast is a federal subject of Russia . Its administrative center is the city of Kursk.-Geography:The oblast occupies the southern slopes of the middle-Russian plateau, and its average elevation is from 177 to 225 meters . The surface is hilly, and intersected by ravines...
, 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
Late Miocene
The Late Miocene is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch....
(23.03 to 5.332 million years ago) of Palembang
Palembang
Palembang is the capital city of the South Sumatra province in Indonesia. Palembang is one of the oldest cities in Indonesia, and has a history of being a capital of a maritime empire. Located on the Musi River banks on the east coast of southern Sumatra island, it has an area of 400.61 square...
, Indonesia
Indonesia
Indonesia , officially the Republic of Indonesia , is a country in Southeast Asia and Oceania. Indonesia is an archipelago comprising approximately 13,000 islands. It has 33 provinces with over 238 million people, and is the world's fourth most populous country. Indonesia is a republic, with an...
. These were later transferred to the living species Aglaomorpha heraclea. It remains, as of 2011, the oldest known drynarioid.
Below is a reconstruction of the evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
of species of Drynaria and closely related genera Aglaomorpha and Selliguea
Selliguea
Selliguea is a fern genus in the family Polypodiaceae.-Species list:* Selliguea albicaula M.Kato & M.G.Price; Acta Phytotax. Geobot. 41: 72...
based on combined morphological and molecular phylogenetic data (Janssen & Schneider, 2005). Drynaria is likely to be paraphyletic
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...
, in contrast to Aglaomorpha, another genus which possess specialized humus-collecting structures.
See also
- PlatyceriumPlatyceriumPlatycerium is a genus of about 18 fern species in the polypod family, Polypodiaceae. Ferns in this genus are widely known as staghorn or elkhorn ferns due to their uniquely-shaped fronds...
– staghorn or elkhorn ferns - AspleniumAspleniumAsplenium is a genus of about 700 species of ferns, often treated as the only genus in the family Aspleniaceae, though other authors consider Hymenasplenium separate, based on molecular phylogenetic analysis of DNA sequences, a different chromosome count, and structural differences in the rhizomes...
– spleenworts, bird's-nest ferns, and walking ferns - PolypodiumPolypodiumPolypodium is a genus of 75–100 species of true ferns, widely distributed throughout the world, with the highest species diversity in the tropics. The name is derived from Ancient Greek poly "many" + podion "little foot", on account of the foot-like appearance of the rhizome and its branches...
– rockcap ferns