- Protologue: in Schrader, J. Bot. 1800(2): 106 (1801).
- Family: Schizaeaceae
- Chromosome number: x= 28, 29, 30;L. auriculatum: 2n= 140;L. circinnatum: 2n= 58, 116;L. flexuosum: 2n= 112;L. japonicum: 2n= 58, 116;L. microphyllum: 2n= 60, 120
Major species and synonyms
- Lygodium auriculatum (Willd.) Alston, Reinwardtia 5: 16 (1959), synonyms: L. semihastatum Desv. (1827, nom. illeg.), L. circinnatum (Burm.f.) Swartz var. semihastatum Fosb. (1941).
- Lygodium circinnatum (Burm.f.) Swartz, Syn. fil.: 153 (1806), synonyms: L. dichotomum (Cav.) Swartz (1806), L. pedatum (Burm.f.) Swartz (1806), L. basilanicum Christ. (1907).
- Lygodium flexuosum (L.) Swartz, in Schrader, J. Bot. 1800(2): 106 (1801), synonyms: L. pinnatifidum Swartz (1801), L. serrulatum Blume (1828).
- Lygodium japonicum (Thunb.) Swartz, in Schrader, J. Bot. 1800(2): 106 (1801), synonyms: L. dissectum Desv. (1811), L. tenue Blume (1828), L. mearnsii Copel. (1908).
- Lygodium microphyllum (Cav.) R. Br., Prodr.: 162 (1810), synonym: L. scandens Swartz (1801, excl. syn. L.).
- climbing fern, snake fern (En)
- Philippines: nito.
- Philippines: nito, nitong puti (Tagalog), balanitu (Bontoc).
- Indonesia: pakis rambat (Javanese), paku hata (Sundanese), kapai gorita (Moluccas)
- Malaysia: ribu-ribu dudok, ribu-ribu bukit, paku jari merah (Malay)
- Philippines: nito (Tagalog), agsam (Albay), nitong puti (Tayabas)
- Thailand: yaan phi phek, li-phao haang kai (peninsular).
- Indonesia: hata kembang (Sundanese), paku ribu-ribu (Asahan), durhawa (Alor)
- Malaysia: ribu-ribu gajah, akar sidin (Malay), haî chîn shâ shù (Chinese)
- Papua New Guinea: tatan (Orokawa Mumuni), zangi (Orokawa Horata)
- Philippines: nito (Tagalog), nito a dadakkel (Iloko), kalulung (Ibanag)
- Thailand: kra chok (Prachin Buri), kuut kong (northern), lee phao (peninsular)
- Vietnam: thong bong, bong bong, duong vong.
- Japanese climbing fern (En)
- Indonesia: pakis kembang (Javanese), paku areuy, hata kawat (Sundanese)
- Malaysia: ribu-ribu, selada, capay alus (Malay)
- Philippines: nito (Tagalog), nitong puti (Rizal, Cavite, Batangas), kerekai (Ibanag)
- Thailand: kuut ngo ngae, phak ngo ngae (northern).
- Small leaved climbing fern, climbing maidenhair (En)
- Indonesia: paku tali (general), paku kawat (western Sumatra), paku hata leutik (Sundanese)
- Malaysia: ribu-ribu, selada, kapai alus (Malay)
- Philippines: nito, nitong puti (Tagalog), nitong parang (Rizal)
- Thailand: kachot nuu (south-eastern), ree-bun paa dee, liphao yung (peninsular).
Origin and geographic distribution
Lygodium is a pantropical to southern temperate genus. Most of the about 40 species are found in South-East Asia and Central America (with one species extending into south-eastern United States).
L. auriculatum is distributed in Indo-China, eastern Borneo, the Philippines and Micronesia.
L. circinnatum is found from Sri Lanka and north-eastern India to southern China, throughout South-East Asia to Vanuatu and Solomon Islands. In the Philippines it is the most common species.
L. flexuosum occurs from Sri Lanka and the Himalayas to southern China, Hong Kong, Ryukyu Islands, throughout South-East Asia to northern Queensland (Australia).
L. japonicum is distributed from Sri Lanka and the Himalayas to northern China, Korea, Japan and throughout most of seasonal South-East Asia to New Guinea but absent in areas with a uniformly humid climate as in Peninsular Malaysia, Sumatra and Borneo. It has become naturalized in Florida and Texas (United States) and is also cultivated as an ornamental.
L. microphyllum is distributed in tropical Africa, throughout South-East Asia to Bangladesh and Hong Kong, Australia and Melanesia. It has naturalized in the southern United States where locally it has become a nuisance.
The very young leaves of all Lygodium species are eaten as a cooked vegetable, especially in Indonesia when other vegetables are lacking, but also in Malaysia. All over South-East Asia the winding rachises are used for plaiting and weaving, sometimes in combination with rattan and bamboo, to make hats, boxes and baskets. The splints of the rachises are freed and dried in the sun (known as "nito" in the Philippines). Everywhere the cord-like rachises are used for tying rice sheaves as they are usually ready to hand. In some areas it played a role at initiation ceremonies for new houses, fishing stations and, especially, new rice fields. In the latter case, the attributes used in the ceremony such as the broom (for sprinkling consecrated rice paste) and the baskets might be made of Lygodium rachises. In the Philippines bracelets made of L. auriculatum are believed to protect against poison. Medicinally, Lygodium is used in the treatment of various ailments, especially those related to the skin. In Malaysia L. circinnatum was used as a childbirth medicine and in New Guinea as a contraceptive. In Indonesia, the pounded leaves of L. circinnatum were applied to wounds. In Indonesia and the Philippines, the roots and rachises were chewed and applied to bites of venomous reptiles and insects to neutralize the poison. Likewise in Bangladesh the juice of the whole plant of a Lygodium sp. is applied to insect bites after squeezing out a little blood. L. flexuosum was used in Malaysia in external applications for skin complaints, including ringworm, and in an extract prepared with many other ingredients drunk for fever. In Vietnam the plant is boiled and taken as diuretic or crushed to cover wounds. In Nepal, the aerial parts of L. japonicum are made into a paste to treat scabies, the juice to treat herpes and wounds. L. microphyllum was used for external application in poultices or emollients to bruises, burns and sprains and it is also supposed to reduce bleeding. In Malaysia it was processed into an astringent decoction to treat dysentery and spitting of blood. The decoction may be sweetened or combined with cordyline and some salt and taken before the meals. Pills made of the leaves were variously used: taken internally for skin complaints, applied externally as poultices against skin diseases, measles and swellings and as an ingredient of lotions to cool fever. In Java (Indonesia), L. microphyllum was used as a remedy for sprue and in Ivory Coast to cure hiccup. In the eastern highlands of New Guinea, fresh leaves of L. longifolium (Willd.) Swartz are chewed with ash salt for stomach-ache and diarrhoea. The Kofanes and other Indian tribes of Colombia added a handful of leaves of L. venustum Swartz to the "ayahuasca", a hallucinogenic drink made of Banisteriopsis caapi (Spruce ex Griseb.) Morton.
Production and international trade
No international trade in Lygodium exists. Lygodium is not cultivated commercially except for L. japonicum which is widely available as an ornamental. All other species are only collected in the wild and used locally.
L. flexuosum has been reported to show antifertility activity. It contains O-p-coumaryl-dryocrassol, dryocrassol, tectoquinone, kaempferol, kaempferol-3-β-D-glucoside, β-sitosterol and stigmasterol. Methanol extracts of L. japonicum exhibit antiviral activity against the Sindbis virus and antibiotic activity against Bacillus subtilis , Mycobacterium phlei , and the dermatophytic mould Trichophyton mentagrophytes . Water extracts of L. japonicum showed antimicrobial activity against Bacillus subtilis, Escherichia coli, Micrococcus luteus , Pseudomonas aeruginosa and Staphylococcus aureus . L. circinnatum and L. flexuosum have also been investigated for their antimicrobial property. Rachises of L. longifolium have a higher tensile strength than those of L. circinnatum , L. flexuosum and L. microphyllum .
Ferns with extremely long leaves that climb, twining through the vegetation. Rhizome creeping, hairy but without scales. Leaves monostichous, twining, of indefinite growth, usually a few m long; the climbing rachis bears alternate, short primary branches with a hairy, dormant apex, and a pair of apparently opposite secondary branches; secondary branches may bear leaflets or tertiary branches which are pinnately arranged or once or more times dichotomously branched; sterile leaflets entire, toothed or lobed, veins free or less often reticulate; fertile leaflets fringed along their margins with short narrow lobes (sorophores), each lobe bearing 2 rows of sporangia, each sporangium attached to a short vein and covered by a small indusium (each sorus consists of 1 sporangium only). Spores tetrahedral and trilete.
- L. auriculatum . Rhizome short-creeping, bearing leaves close together, its apex and bases of petioles densely covered with dark hairs. Juvenile leaves once dichotomous, each branch bearing a palmatisect leaflet, usually 5-lobed, with truncate base, edges closely and irregularly serrate; rachis of climbing leaves hardly 2 mm in diameter, usually glabrous; apex of primary rachis-branches covered with pale brown hairs having a swollen base; secondary rachis-branches rarely bearing a simple leaflet, most commonly once dichotomous, one branch with a simple, one with a forked leaflet, less often each branch with a simple leaflet; sterile leaflets suboblong, 12-20 cm × 1-3 cm, base usually asymmetric and strongly cordate-auriculate on the outer side, margins not thickened and very shallowly serrate, apex subacute; lamina of fertile leaflets 1-2(-3) cm wide. Sorophores 3-9 mm long, constricted at the base, at the apices of short triangular lobes of the lamina; indusia glabrous; spores irregularly warty, very variable.
- L. circinnatum . Rhizome short-creeping, bearing leaves very close together, its apex and bases of petioles densely covered with black hairs. Juvenile leaves once dichotomous, each branch bearing a pedato-palmatisect leaflet; lobes 4-5, subequal, up to 25 cm × 3.5 cm, the midrib of an outer lobe arising near base of the next inner lobe, margins entire but often somewhat crisped, pale and much thickened (translucent when living), apex acute to acuminate, surface glabrous but conspicuously warty when dry (not when living), veins uniting with the thickened margin; rachis of climbing leaf up to10 m long and 2-5 mm in diameter, glabrous; apex of primary rachis-branches with sunken dormant apex covered with pale hairs which are not thickened at the base; secondary rachis-branches unbranched, 2-6 cm long, or once dichotomous with each branch 1-2 cm long beyond the fork; sterile leaflets usually with 2-6 subequal diverging lobes which are separate to within 2 cm from the base, base cuneate or truncate, margin entire, pale and thickened, surfaces nearly always warty when dry; fertile secondary rachis-branches unbranched or 1-3 times dichotomous (rarely sub-pinnate); fertile leaflets usually sessile in pairs at the end of the ultimate branches, or members of a pair partly fused at the base, less often 3-5 lobed (always so if the secondary rachis is unbranched), lamina more or less reduced and commonly 3-6 mm wide. Sorophores 2-5 mm long, sessile; spores finely and evenly verrucose all over the surface.
- L. flexuosum . Rhizome short-creeping, densely covered with roots, the petioles very close together, apex covered with brown-black hairs. Juvenile leaves once or twice dichotomous, each branch bearing a single leaflet which is deeply palmately 3-7-lobed; lobes almost equal, base cordate, margins serrate; rachis of scandent leaves narrowly winged, flattened and puberulous on the upper surface between the wings; primary rachis-branches up to 3 mm long, dormant apex covered with pale brown hairs; secondary rachis-branches pinnate; sterile leaflets of lower branches palmately 5-lobed; higher secondary branches bearing 3-5(-7) leaflets on each side and an apical one, the apical and lower leaflets asymmetrical or more or less lobed at the base, the lowest often with 2 or 3 (exceptionally up to 6) separate quaternary leaflets at its base; sterile leaflets 3-10 cm × 8-15 mm, margins serrate, apex subacute, lower leaflets stalked, upper sessile, lamina rather thin, costae usually bearing scattered long hairs, less often densely short-hairy, veins often with scattered short hairs on the lower surface; fertile leaflets smaller than sterile ones. Sorophores 3-5(-10) mm long at the apices of small triangular lobes; indusia subglabrous; spores finely verrucose.
- L. japonicum . Rhizome wide-creeping, dichotomously branched, 2-5 mm in diameter, densely clothed with dark brown hairs, leaves usually 5-10 mm apart. Juvenile leaves erect, the first branching an unequal dichotomy, the two main branches of large leaves bipinnate, deltoid in outline, with palmatisect leaflets, their margins doubly serrate. Rachis of climbing leaves hardly 2 mm in diameter, glabrous apart from minute hairs on the flattened adaxial surface between the narrow wings; primary rachis-branches 3-10 mm long; secondary branches of leaves on young or stunted plants pinnate, on well-grown plants leaves 2-3-pinnate, deltoid in outline, commonly 12 cm × 12 cm, rachises densely short hairy on the upper surface and bearing fewer longer hairs elsewhere; sterile tertiary leaflets of lower rachis-branches palmate with 5-7 lobes, the middle lobe much longer than laterals; tertiary leaflets higher up the leaf trilobed with an elongate middle lobe or pinnate with small oblique and often lobed quaternary leaflets and a usually deltoid-pinnatisect terminal leaflet about 3 cm long, margins acutely biserrate, apex obtuse or subacute; stalks of leaflets up to 3 mm long, never articulate or thickened at apex, costae usually bearing long scattered hairs, veins and surfaces usually glabrous; fertile secondary branches tripinnate, the leaflets smaller than sterile ones. Sorophores 2-12 mm long; indusia glabrous or with a few hairs if the lamina is hairy; spores sparsely and finely verrucose.
- L. microphyllum . Rhizome wide-creeping, dichotomously branched, 2.5 mm in diameter, densely clothed with short brown-black hairs. Juvenile leaves small, commonly once dichotomous (the stipe distinctly winged below the dichotomy), each branch bearing a 4-lobed leaflet not jointed at the base; lobes 3-5 cm × 5 mm, thin, glabrous, margins crenately toothed; rachis of climbing leaves glabrous, commonly 2-3 m long, hardly 1.5 mm in diameter; primary branches about 4 mm long; secondary rachis-branches pinnate, in all up to about 15 cm long, with 3-6 stalked leaflets on each side (stalks 2-4 mm long) and a similar or geminate leaflet; leaflets quite glabrous, mostly ovate (sterile leaflets often elongate with broader base on young plants), 1-4(-6) cm × 6-18 mm, margins of sterile ones minutely crenate, a joint always present at base of lamina, where the wing, which in other species connects stalk and lamina, is constricted; fertile leaflets usually shorter than sterile ones but lamina hardly narrowed. Sorophores 4-6 mm long; spores faintly granulose with a raised reticulum on the outer surface.
Growth and development
The gametophytes of Lygodium are more or less cordate, sometimes asymmetrical, and glabrous. The antheridia are larger, and of a more complex, primitive, pluricellular type than in most leptosporangiate ferns. The Lygodium leaves are borne on the dorsal surface of a subterranean rhizome and undergo a twining growth to form the aerial part of the shoot. These leaves have continuous apical growth, circumnutation, a delay in leaflet expansion below the leaf apex and budlike resting leaflet apices. The determinate primary leaves and the indeterminate climbing leaves arise from a single cell on the flank of the apical meristem; they are strictly foliar in nature and structurally homologous with each other. The climbing leaves of L. japonicum mature in a period of more than 70 days during which about 36 primary rachis-branches are produced alternately on the main rachis. The crozier remains compact throughout the growing period. The lamina segments on secondary rachis-branches expand simultaneously; in basal segments it takes about 23-28 days. In addition to apical growth, the length of the main rachis is increased by elongation of the internodes. The growth of the rachis is maximal when growing upward, but retarded when the vine grows either upside down or horizontally; the number of primary rachis-branches on such leaves is relatively low. Damaged leaves may activate dormant buds.
Other botanical information
Lygodium stands apart from the rest of the Schizaeaceae (which comprises 4 genera) through its asymmetrical sporangia, each covered by a kind of indusium, and also through its twining rachis. Furthermore it is biochemically divergent. It fits very well in the subfamily Lygodioideae . Other authors give it a family on its own, the Lygodiaceae .
Lygodium plants are twining climbers, mainly in secondary vegetation, producing fertile leaflets on parts exposed to the brightest light. L. auriculatum usually grows in light shade in tropical forests. L. circinnatum occurs in light shade in tropical evergreen forest, from sea-level up to 1500 m altitude. It grows in wet soils and never in locations where the soil becomes seasonally very dry. L. flexuosum climbs on shrubs or on branches of tall trees in open locations and in deciduous or mixed forest, from sea-level up to 1000 m altitude. It is a vigorous fern which may form thickets along forest margins and in swampy areas. L. japonicum occurs in regions with a pronounced dry season, on dry open grass fields or mountain slopes in deciduous forest, from sea-level up to 2500 m altitude. L. microphyllum grows in edges of secondary forest or is a climber on woody plants, bushes or on branches of tall trees, usually on dry slopes in open areas. It prefers clay soils and frequently forms thickets in open, swampy locations in regions with a dry season, from sea-level up to 1300 m altitude. Its naturalization in south-eastern Florida (United States) was first detected in 1965. Growing in tropical and subtropical wetlands and areas with moist soils, it is well adapted to most parts of Florida where its current rate of spread and environmental impact is serious. It has become an aggressive invader of natural vegetation in many different habitats that are frequently dominated by the species. Currently, increasing densities and continued expansion of its distribution are observed. Moreover, it has become a serious fire hazard.
Propagation and planting
All Lygodium species grow from spores. Those with long-creeping rhizomes can be propagated by rooted rhizome cuttings.
L. circinnatum is suitable for planting in a tropical garden, requiring warmth and plenty of moisture. It can also be grown in a pot but the roots are sensitive to drying. L. flexuosum can be grown easily. It should be planted in the ground and trained on supports such as trees and trellises because it can be very vigorous. L. japonicum can be grown in a pot or basket in a warm garden position and is cultivated commercially as an ornamental. L. microphyllum adapts readily to cultivation and will succeed in pots, baskets or in a semi-shady garden position. Because of its rampant climbing habit it needs regular trimming.
Diseases and pests
The thrips Octothrips lygodii was widely found in South-East Asia causing severe damage to L. microphyllum .
All used parts of Lygodium are harvested whenever needed.
Handling after harvest
Particularly the lower parts of rachises of Lygodium are used for cording. They are cut into pieces of about 1.5-2 m length and split to take out the green inner part. The bast is then split into pieces of the required length and can be stored for later use. Sometimes the rachis parts are first dried before being split. Entire plant parts, stored dry for 6 years at room temperature, had retained at least part of their antibiotic activity.
All Lygodium species are rampant in common, non-vulnerable habitats. There are no known germplasm collections, but none of the species is threatened with genetic erosion or extinction.
Lygodium ferns are used as medicinal plants in many countries, in addition to their uses as fibres and vegetables. Their widespread medicinal application on wounds and infections suggests antimicrobial properties, which have been partly confirmed experimentally. Further pharmacological research is needed to explore these possibilities. The use in plaiting will persist in touristic and traditional niche markets.
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- Bodner, C.C. & Gereau, R.E., 1988. A contribution to Bontoc ethnobotany. Economic Botany 42(3): 307-369.
- Griggs, J.K., Manadhar, N.P., Towers, G.H.N. & Taylor, R.S.L., 2001. The effect of storage on the biological activity of medicinal plants from Nepal. Journal of Ethnopharmacology 77: 247-252.
- Holttum, R.E., 1959. Schizaeaceae. In: van Steenis, C.G.G.J. & Holttum, R.E. (General editors): Flora Malesiana, Series 2. Pteridophyta (Ferns & fern allies). Vol. 1, part 1. Martinus Nijhoff / Dr W. Junk Publishers, The Hague, The Netherlands. pp. 37-61.
- Punetha, N., 2000. Phenology of the climbing leaf of Lygodium japonicum (Thunb.) Sw. (Schizaeaceae). Phytomorphology 50(1): 75-85.
- Sahi, A.N. & Tiwari, D.N., 1998. Pot culture technique for propagating the tropical fern Lygodium japonicum (Filicales; Lygodiaceae). National Academy of Science and Letters India 21(11-12): 309.
- Taylor, R.S.L, Manandhar, N.P., Hudson, J.B. & Towers, G.H.N., 1996. Antiviral activities of Nepalese medicinal plants. Journal of Ethnopharmacology 52(3): 157-163.
Titien Ngatinem Praptosuwiryo