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]
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.
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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.
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]
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]
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]
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]
The following is the list of accepted species according to the Checklist of Ferns and Lycophytes of the World as of August 2019:[22]
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]
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]
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]
Aglaomorpha is a genus of ferns in the subfamily Drynarioideae of the family Polypodiaceae. The Pteridophyte Phylogeny Group 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 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.
Drynaria es un género de helechos perteneciente a la familia Polypodiaceae.[2] Es originario del Sudeste de Asia.
Drynaria fue descrito por (Bory) J.Sm. y publicado en Journal of Botany, being a second series of the Botanical Miscellany 4: 60. 1842.[3] La especie tipo es: Drynaria quercifolia (L.) J. Sm.
Drynaria es un género de helechos perteneciente a la familia Polypodiaceae. Es originario del Sudeste de Asia.
Drynaria-slekten (latin: Drynaria) er en slekt av bregner i Drynariaceae-seksjonen av sisselrotfamilien, som igjen på sin side inngår i den meget store, og monofylittiske ordenen (kladen) Polypodiales.[1] gruppen har 19 arter og 1 hybrid.
De fleste artene er oppreiste og av moderat størrelse, ofte inntil om lag 30 cm.
Drynaria-artene vokser overveiende i Asia og da først og fremst Kina. Men det er også arter i Afrika og nordøstlige Australia.
Drynaria-slekten (latin: Drynaria) er en slekt av bregner i Drynariaceae-seksjonen av sisselrotfamilien, som igjen på sin side inngår i den meget store, og monofylittiske ordenen (kladen) Polypodiales. gruppen har 19 arter og 1 hybrid.
De fleste artene er oppreiste og av moderat størrelse, ofte inntil om lag 30 cm.
Drynaria-artene vokser overveiende i Asia og da først og fremst Kina. Men det er også arter i Afrika og nordøstlige Australia.
Drynaria (Bory) J. Sm. – rodzaj roślin należący do rodziny paprotkowatych. Należy do niego ok. 144 epifitycznych gatunków, pochodzących z Afryki, Azji i Północno-Wschodniej Australii[2]. Gatunkiem typowym jest Drynaria quercifolia ((L.) J. Sm.)[3].
Drynaria należy do rodziny paprotkowatych (Polypodiaceae), rząd paprotkowce Polypodiales.
Gromada paprotniki (Polypodiophyta Cronquist), podgromada Polypodiophytina (Reveal), klasa paprocie (Polypodiopsida Cronquist), podklasa paprocie cienkozarodniowe (Polypodiidae Cronquist), rząd paprotkowce (Polypodiales Mett. ex A.B. Frank in Leunis), podrząd Polypodiineae (Bessey), rodzina paprotkowatych (Polypodiaceae Bercht. & J. Presl)[5].
Drynaria (Bory) J. Sm. – rodzaj roślin należący do rodziny paprotkowatych. Należy do niego ok. 144 epifitycznych gatunków, pochodzących z Afryki, Azji i Północno-Wschodniej Australii. Gatunkiem typowym jest Drynaria quercifolia ((L.) J. Sm.).
Дринарії мають оригінальну будову. Складаються з нижніх безплідних лисків та з зелених листків, що формують спори. Нижні листки схожі на листя дуба, але набагато більші за них. Вони формують корзинку, в якій розвиваються коріння. Плодоносні листки Drynaria fortunei досягають до 75 см у довжину.
Епіфітні рослини, що ростуть у заглибленнях між гілками. Проте зустрічаються такі, що можуть прикріплятися до голого стовбура. Нижні безплідні листки формують корзинку, де осідають органічні речовини та пил, для живлення рослини, та збирається дощова волога.
Вирощують як декоративну рослину через оригінальний вигляд.
Це незавершена стаття про Папороті.
本文参照
ハカマウラボシ属 Drynaria は、ウラボシ科のシダ植物。着生で、葉に2形があり、根元に広がる葉に落葉を貯める。
ハカマウラボシは着生シダ植物であり、葉に2形がある。その1つが巣葉と呼ばれ、これは着生した木の幹に密着して上向きに口を開け、そこの上からの落葉を集める。これは樹上という肥料に乏しい環境でそれを集めるための適応と見られる。同様の2形は同科のビカクシダ属にも見られるものであるが、本属では巣葉が胞子葉の形、単羽状と基本的には同じ形を保っているのが特徴である。
園芸的に栽培される場合もあり、その場合には類似のカザリシダ属なども混同されることがある。
着生のシダ植物[1]。根茎は太くて多肉質で、若い部分は細長い鱗片に覆われる。葉は根茎の上に少しずつ距離を置いて出る。葉にははっきりした2形がある。1つは胞子葉で、これは細長く伸び出し、単羽状に裂けるか、あるいは単羽状複葉となり、緑色で光合成をし、葉裏に胞子嚢群をつける。もう1つは比較的短くて幅広く、また柄がない葉で、これを巣葉という。巣葉は縁に大きな鋸歯があって浅く単羽状に裂け、羽状の主脈と側脈がはっきりと見て取れる。またこの葉は成長するとすぐに緑から褐色に変わるが、枯れた姿でそのまま宿在し、着生している樹幹の上にあってその葉と樹皮の隙間の部分に落ち葉を蓄える。またこの葉には胞子嚢は生じない。なお一般にウラボシ科では葉の基部に関節があって葉が枯れるとこの部分で折れて取れるが、本属ではこれが発達せず、胞子葉の場合も落葉はせず、裂片や羽片がそれぞれに枯れて外れ落ち、後に葉柄と中肋だけが残る。
葉脈は独特の複雑な構造を持ち、ドリナリア型とも呼ばれる。裂片の中脈からでるはっきりした主側脈と、このような主側脈間をそれらと直交して結びつける脈によって一次の網目が作られ、その内部に伸びる細脈で二次の網目が形成される。更にこの二次の網目の中に遊離した細脈が入り込む。胞子嚢群は包膜が無く、この遊離細脈か細脈の分岐点に生じ、それぞれの胞子嚢群は散在するものもあれば互いに繋がり合ってパターンを形成する例もあり、その型によって属の細分がなされることもある。
学名は森のニンフであるドリアス(Dryas)に由来するとも、ギリシャ語のdrys（カシワ）に由来し、巣葉がカシワの葉に似ることに由来するとも言われる。
樹幹に着生する様子
D. quercifolia・以下すべて
巣葉と胞子葉の基部
2種の葉・図版
成長途中の巣葉
胞子葉の裏面
この植物は樹幹に着生し、匍匐茎を伸ばすが、無柄で幅広い巣葉は往々に互いに重なり合い、それによって匍匐茎と根をほとんど覆い隠す。この葉の内側に落葉を蓄積するが、その量はビカクシダ属に比べればずっと少ない[2]。
旧世界の熱帯域とオーストラリアに分布し、約20種が知られる[3]。
同じように葉に2形を持つものにビカクシダ属 Platycerium があり、やはり空中へ伸びる葉に胞子嚢をつけ、根本に広がる葉に落ち葉を蓄える。その点では本属と似ているが、本属のものではどちらの葉もこの科本来の葉の形をある程度残しているのに対して、ビカクシダ属ではかなり独特の姿になっている。またカザリシダ属 Agraomorpha などでは葉に2形はないものの、胞子をつける葉の基部が横に広がって落ち葉を受ける構造となっており、同じような適応の方向を示すものとなっている[4]。
ちなみに系統関係の面からは本属とビカクシダ属は縁が遠い。本属ともっとも近縁なのはカザリシダ属と考えられている[5]。
以下に代表的な種をあげておく[6]。
日本には従来は発見されていなかったが、1991年に沖縄本島で発見されたことが報告された。ただしたった1カ所、それも小さなコロニーだけとのことで、絶滅が大いに危惧されている[7]。 ※ただし、この自生地は米軍基地内のため、今のところは乱獲を防ぐことができている。
観賞用に温室で栽培されることがある。その際には多少似た姿を持つカザリシダ属(アグラオモルファ)なども混同され、本属と纏めてドリナリアの名で呼ばれる場合もある[8]。
D. rididula は胞子葉は長さ1-2mに達し、長い葉柄がある。南太平洋ではしばしば現地民族舞踊の際の髪飾りに用いられる[9]。
またハカマウラボシ及び近縁種幾つかの根茎が薬用に用いられる。苦みがあって中国の医術で強精、長期の下痢、リュウマチ痛、歯痛などに用いられ、更に打撲傷や骨折にも効があるとされている[10]。
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