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Ceratopteris thalictroides (L.) Brongn.

Protologue
Bull. Sci. Soc. Philom. Paris, sér. 3, 8: 186, t. 3–4 (1821).
Family
Pteridaceae
Chromosome number
n = 77, 78, 80; 2n = 154, 156
Synonyms
Acrostichum thalictroides L. (1753), Ceratopteris cornuta (P.Beauv.) Lepr. (1830).
Vernacular names
Water sprite, swamp fern, floating stag’s horn, Sumatra fern, Indian fern, oriental water fern (En). Fougère de Sumatra, fougère aquatique flottante (Fr).
Origin and geographic distribution
Ceratopteris thalictroides is found worldwide in the lowland tropics especially of the Old World. It is found throughout African coastal lowland, and is especially common in West African estuaries and swamps.
Uses
The only record of Ceratopteris thalictroides being eaten in Africa is from Madagascar, where it is used in a similar way as watercress. It is commonly eaten throughout South-East Asia. In Malaysia and Japan it is an established luxury vegetable. The young leaves, before they have uncurled, make excellent greens and when cooked or blanched they can be eaten as a salad.
The plants can be used as a green manure in rice fields. Medicinally, this fern (both leaf and root) is used in Malaysia and the Philippines as a poultice against skin complaints, e.g. as a drawing agent on carbuncles, in China as a styptic to stop bleeding. Ceratopteris species, including Ceratopteris thalictroides, are grown as ornamentals in aquariums, popularly called ‘water sprite’. Ceratopteris thalictroides and especially also Ceratopteris richardii Brongn. serve as model plants in developmental biology and molecular research. They are useful for research because they have independent haploid and diploid life phases, a short life cycle, a simple genetic system, and reproduce by single-celled haploid spores.
Properties
Alkaloids, arbutin and tannin have been found in the green parts of Ceratopteris thalictroides.
Botany
Aquatic or semi-aquatic fern, up to 1 m tall, floating or rooted in soil, with short, erect rhizome and rosette-like tuft of leaves. Leaves dimorphous; sterile leaves with succulent petiole 5–50 cm long, sparsely scaly with broad, pale brown scales, filled with air canals, lamina of sterile leaves 1–3-pinnate, up to 20 cm × 40 cm, membranous, glabrous, with evident anastomosing veins, often provided with adventitious buds in axils of pinnae, pinnae irregularly shaped, ultimate lobes linear-oblong to elliptical; lamina of fertile leaves erect, 2–4-divided, up to 100 cm × 50 cm, ultimate lobes linear, margin revolute, covering the sporangia scattered individually along the veins. Spores tetrahedral, 95–125 μm in diameter, pale yellow, translucent, with irregular reticulum.
Ceratopteris is often classified in Parkeriaceae, a family included in Pteridaceae here. Ceratopteris thalictroides is morphologically very variable. If grown under water, as in aquariums, the lamina of the submersed (sterile) leaves are finely 3–4-pinnate, with ultimate lobes linear.
Ecology
Ceratopteris thalictroides is a floating or loosely rooted aquatic fern. It grows in swampy areas, swamp forests, marshes, natural and man-made ponds, mostly in stagnant water bodies or in still pockets along slow flowing rivers, from sea-level to 1300 m, but mostly below 500 m altitude. It occurs in full sun to moderate shade, sometimes massed on or around logs or other floating vegetation.
Management
Ceratopteris thalictroides has a very short life cycle. The whole cycle from spore to spore can be completed in less than 30 days. Apart from spores, the plant reproduces by bulbils growing on the leaves. In Japan and South-East Asia Ceratopteris thalictroides is sometimes cultivated for food, but rarely commercially. Farmers harvest the plants from rice fields and swampy locations for local consumption. In aquarium culture the mother plants are normally split in 3-leaf segments, but any part of the leaf if left floating in the water will give rise to new plantlets.
Genetic resources and breeding
Ceratopteris thalictroides is distributed throughout the tropics and is not in danger of genetic erosion or extinction. In Africa, the programmes to conserve the natural wetlands will help the species to maintain its genetic diversity. Germplasm collections or breeding programmes are not known to exist, also because it is considered a weed in rice fields.
Prospects
The fact that Ceratopteris thalictroides plants are eaten as a vegetable in Asia and Madagascar indicates a possible similar use for the African continent. It seems worthwhile investigating this potential further. Studies should include the nutritional value, possible negative health indications, unknown local uses, the environmental impact of large-scale gathering from the wild, as well as possibilities for commercial production.
Major references
• Burkill, H.M., 2000. The useful plants of West Tropical Africa. 2nd Edition. Volume 5, Families S–Z, Addenda. Royal Botanic Gardens, Kew, United Kingdom. 686 pp.
• Chanpen Prakongvongs, 2003. Ceratopteris thalictroides (L.) Brongn. In: de Winter, W.P. & Amoroso, V.B. (Editors). Plant Resources of South East Asia No 15(2). Cryptogams: ferns and fern allies. Backhuys Publishers, Leiden, Netherlands. pp. 75–77.
• Copeland, E.B., 1942. Edible ferns. American Fern Journal 32: 121–126.
• Decary, R., 1946. Plantes et animaux utiles de Madagascar. Annales du Musée Colonial de Marseille, 54e année, 6e série, 4e volume, 1er et dernier fascicule. 234 pp.
Other references
• Alston, A.H.G., 1959. The ferns and fern-allies of West Tropical Africa. Crown Agents for Oversea Governments and Administrations, London, United Kingdom. 89 pp.
• Benedict, R.C., 1909. The genus Ceratopteris: a preliminary review. Bulletin of the Torrey Botanical Club 36: 463–476.
• Javalkegar, S.R., 1960. Sporogenesis and prothallial development in Ceratopteris thalictroides. Botanical Gazette 122: 45–50.
• Klekowski, E.J., 1970. Reproductive biology of the Pteridophyta 4. An experimental study of mating systems in Ceratopteris thalictroides (L.) Brongn. Botanical Journal of the Linnean Society 63: 153–169.
• Lloyd, R.M., 1972. Species delimitation in the genus Ceratopteris (Parkeriaceae). American Journal of Botany 59: 676.
• Lloyd, R.M., 1974. Systematics of the genus Ceratopteris Brongn. (Parkeriaceae) 2. Taxonomy. Brittonia 26: 139–160.
• Masuyama, S. & Watano, Y., 1994. Hybrid sterility between two isozymic types of the fern Ceratopteris thalictroides in Japan. Japan Plant Research 107: 269–274.
• Pal, N. & Pal, S., 1969. Studies on the morphology and affinity of the Parkeriaceae. 3. A discussion on the systematic position of the family and specific delimitation of the genus Ceratopteris. Bulletin of the Botanical Society of Bengal 23: 17–25.
• Schelpe, E.A.C.L.E., 1970. Adiantaceae. In: Exell, A.W. & Launert, E. (Editors). Flora Zambesiaca. Pteridophyta. Crown Agents for Oversea Governments and Administrations, London, United Kingdom. pp. 98–138.
• Tardieu-Blot, M.L., 1964. Adiantaceae. Flore du Gabon. Volume 8. Muséum National d’Histoire Naturelle, Paris, France. pp. 97–119.
Sources of illustration
• Chanpen Prakongvongs, 2003. Ceratopteris thalictroides (L.) Brongn. In: de Winter, W.P. & Amoroso, V.B. (Editors). Plant Resources of South East Asia No 15(2). Cryptogams: ferns and fern allies. Backhuys Publishers, Leiden, Netherlands. pp. 75–77.
Author(s)
W.J. van der Burg
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
Based on PROSEA 15(2): ‘Cryptogams: Ferns and fern allies’.

Editors
G.J.H. Grubben
Prins Hendriklaan 24, 1401 AT Bussum, Netherlands
O.A. Denton
National Horticultural Research Institute, P.M.B. 5432, Idi-Ishin, Ibadan, Nigeria
Associate Editors
C.-M. Messiaen
Bat. B 3, Résidence La Guirlande, 75, rue de Fontcarrade, 34070 Montpellier, France
R.R. Schippers
De Boeier 7, 3742 GD Baarn, Netherlands
General editors
R.H.M.J. Lemmens
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
L.P.A. Oyen
PROTA Network Office Europe, Wageningen University, P.O. Box 341, 6700 AH Wageningen, Netherlands
Illustrator
PROSEA
PROSEA Network Office, Herbarium Bogoriense, P.O. Box 234, Bogor 16122, Indonesia

Correct citation of this article:
van der Burg, W.J., 2004. Ceratopteris thalictroides (L.) Brongn. In: Grubben, G.J.H. & Denton, O.A. (Editors). PROTA 2: Vegetables/Légumes. [CD-Rom]. PROTA, Wageningen, Netherlands.
1, habit; 2, petiole scale; 3, fertile part leaf segment; 4, sporangia along veins; 5, sporangium; 6, spores.
Source: PROSEA