Marsilea minuta (pepperwort)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Marsilea minuta L.
Preferred Common Name
Other Scientific Names
- Marsilea crenata Presl, 1828
- Marsilea diffusa Lepr. ex A. Br.
- Marsilea erosa
- Marsilea perrieriana
- Marsilea senegalensis
International Common Names
- English: airy pepperwort
- French: marsilea a quatre feuilles
Local Common Names
- Indonesia: semanggi
- Japan: nangokudenjiso
- Malaysia: tapak itek
- Philippines: kaya-kayapuan
- Thailand: phakwaen
- MASCR (Marsilea crenata)
- MASMI (Marsilea minuta)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Pteridophyta
- Class: Pteridopsida
- Family: Marsileaceae
- Genus: Marsilea
- Species: Marsilea minuta
Notes on Taxonomy and NomenclatureTop of page M. minuta is a perennial water fern that belongs to the family Marsileaceae and was first described by Presl in 1825. The Marsileaceae are sometimes presented in the subclass Hydropterides (water ferns). However, this is not a homogeneous group because its members merely represent heterosporous derivatives of old plant groups, adapted to aquatic life. This family consists of the genera Marsilea, Pilularia and Regnellidium.
The genus Marsilea includes approximately 65 species, which are aquatic or semi-aquatic (see Braun 1871, 1873; Launert, 1968; Cook et al., 1974). The Marsilea species are difficult to distinguish from each other. Most species display a large phenotypic plasticity, which is exhibited more in the vegetative parts of the plant than in the sporocarps. Consequently, the classification into species is almost exclusively on the basis of the sporocarp.
The names M. minuta and M. crenata have often been treated as distinct species on the basis of small differences in the structure of the sporocarps, but they are widely confused and/or treated as the same, especially in South-East Asia. Hence, for the purposes of this Compendium they are treated as synonymous, and the fern is referred to by the older name, M. minuta.
DescriptionTop of page M. minuta is a perennial fern with slender, rooted, creeping, branching rhizomes bearing erect leaves (sterile fronds) along their length (Kostermans et al., 1987). The leaves, which consist of four, 'clover-like' leaflets at the apex of a slender erect stalk, arise along the length of each rhizome. At the base of the petioles the sporocarps are formed on about 5 mm long stalks. The sporocarps are 3-4 mm long, oblong with rounded ends (when seen laterally), with their long axis at right angles to the stalk.
The bean-shaped sporocarps contain both megasporangia and microsporangia. The features of the sporocarps are important for distinguishing the species. A typical characteristic of M. minuta is that the sporocarps are unribbed and have two basal teeth. The upper tooth is sharply pointed, the lower tooth is shorter and obtuse (see Aston, 1977; Pancho and Soerjani, 1978).
DistributionTop of page According to Aston (1977), M. crenata occurs in Australia in the tropical North, Central Australia and Queensland.
Merlier and Montagu (1982) include M. minuta as a weed of 'Afrique tropicale'. Holm et al. (1979) have few records for M. crenata or M. minuta but many for M. quadrifolia.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|China||Present||Present based on regional distribution.|
|-Hong Kong||Present||Pancho and Soerjani, 1978|
|India||Present||Holm et al., 1979|
|-Kerala||Present||Bhat and Kaveriappa, 2003|
|Indonesia||Widespread||Soerjani et al., 1987|
|Malaysia||Widespread||Holttum, 1954; Pancho and Soerjani, 1978|
|Philippines||Widespread||Pancho and Soerjani, 1978|
|Taiwan||Present||Pancho and Soerjani, 1978|
|Thailand||Widespread||Peudpaichit et al., 1987|
|Algeria||Present||Merlier and Montagu, 1982|
|Comoros||Present||Merlier and Montagu, 1982|
|Madagascar||Present||Merlier and Montagu, 1982|
|Papua New Guinea||Present||Aston, 1977|
Risk of IntroductionTop of page M. minuta is a very troublesome weed in rice in South-East Asia and Australia. Further dispersal of M. minuta by human activities, for example via market gardens or the aquarium trade, should be prevented. Although closely related weedy Marsilea species occur in other areas of the world where rice is a major crop, it is conceivable that M. minuta is a more aggressive species, at least under certain environmental conditions.
HabitatTop of page Marsilea species are found in pools, pans, ditches, swamps and lowland ricefields. M. minuta can grow under water.
Hosts/Species AffectedTop of page M. minuta is a very troublesome weed in lowland ricefields and ditches, and can grow under water: during the first half of the growth period of rice plants, it is a severe competitor, in particular during the first two weeks after transplantation of young rice plants.
It may also affect vegetables that are cultivated in water, such as Ipomoea aquatica.
Host Plants and Other Plants AffectedTop of page
|Oryza sativa (rice)||Poaceae||Main|
Biology and EcologyTop of page In shallow water, temporarily flooded depressions and moist soils, M. minuta is rooted: plants growing in shallow water or wet mud have erect petioles with emergent leaflets. In deeper water, the leaflets float on the water surface.
The primary mode of reproduction is vegetatively by rhizomes. A network of slender, creeping, branched rhizomes can produce numerous erect petioles. Sexual reproduction is connected with the formation of two types of spores: large female spores (megaspores) produced in megasporangia (one spore per megasporangium) and tiny male spores (microspores) produced in microsporangia (numerous spores per microsporangium) (Sculthorpe, 1967). Both macro- and microsporangia occur together in each sporocarp. The stalked sporocarps burst when the female spores are ripe and swollen. A macrospore produces a very reduced prothallus bearing a solitary simple archegonium. The microspores produce a small number of male germ cells. The male germ cells will fertilize the female cells in the archegonia. Sporocarps of Marsilea species have been reported to remain viable for many years (Smith, 1938; Rodgers, 1993).
The leaves of Marsilea species, along with the sporocarps, are consumed by several species of waterfowl. The sporocarps pass through the intestinal tracts of water birds, which therefore provide an efficient means of distribution.
If conditions are favourable, vegetative growth and development are very fast. During the first half of the growth period of rice plants, M. minuta is a severe competitor, in particular during the first two weeks after transplantation of young rice plants.
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
Notes on Natural EnemiesTop of page The insect Nymphula responsalis is able to complete its life cycle on M. minuta (Subagyo, 1975). For its survival, this insect needs relatively deep water. Host-specificity tests on 35 plant species belonging to 19 plant families showed that, apart from M. minuta, the larvae fed and developed on Salvinia spp., Lemna perpusilla, Spirodela polyrhiza, Monochoria vaginalis, Azolla pinnata and Pistia stratiotes (Handayani and Syed, 1976). There are no reports indicating that N. responsalis causes much damage to M. minuta.
A survey for natural enemies attacking M. minuta in Africa is required before the prospects for its biological control in South-East Asia can be evaluated (Waterhouse, 1993)
ImpactTop of page M. minuta is a very troublesome weed in lowland ricefields and ditches in South-East Asia and Australia. In contrast to many other weeds in rice, it can grow when temporarily submerged. It will emerge when it is shallowly buried by soil. The earlier the emergence, the greater the advantage of the weed over rice and the greater the competitive effect. When Marsilea emerges sooner than two weeks after rice has been transplanted, the competitive effect is very severe. Yield reduction of 70% has been recorded (Ampong-Nyarko and de Datta, 1991).
In general, fertilizer application stimulates the growth of rice more than that of M. minuta. Consequently, fertilizer application leads to a lower percentage of yield reduction due to competition with M. minuta. In a pot experiment, the dry weight of rice plants affected by M. minuta varied from 50% (as a percentage of the weed-free control) in the absence of urea to 75% at 6 g urea per pot (Pons et al., 1987).
Uses ListTop of page
Similarities to Other Species/ConditionsTop of page M. minuta may be confused with M. quadrifolia.
Prevention and ControlTop of page Apart from manual control, M. minuta in rice can be controlled using various herbicides. These include applications (after sowing or transplanting of the rice) of bensulfuron-methyl (Peudpaichit et al., 1987), cinosulfuron (Achlderon et al., 1987; Burhan et al., 1989) and 2,4-D (Rodgers, 1993).
Oxyfluorfen (Yasin et al., 1988) and paraquat (Khaw and Ho, 1988) proved to be effective against M. minuta in rice when applied as pre-emergence herbicides.
ReferencesTop of page
Achlderon JI; Hare CJ; Palis FV; Burhan H; Bhandhufalck A; Chong WC, 1987. Setoff - a new rice herbicide for S.E. Asia. Proceedings, 11th Asian Pacific Weed Science Society Conference Taipei, Taiwan; Asian Pacific Weed Science Society, No. 1:73-79
Aston HI, 1977. Aquatic plants of Australia. Melbourne, Australia: Melbourne University Press.
Braun A, 1871. Neuere Untersuchungen über die Gattungen Marsilea and Pilularia. Monatsberichte der königlichen Preussischen Akademie der Wissenschaften zu Berlin, 1870:652-753.
Braun A, 1873. NachtrSgliche Mitteilungen über die Gattungen Marsilea and Pilularia. Monatsberichte der königlichen Preussische Akademie der Wissenschaften zu Berlin, 1872:635-679.
Burhan H; Sozzi D; Zoschke A, 1989. Setoff for weed control in rice: practical experience from Indonesia. Proceedings, 12th Asian-Pacific Weed Science Society Conference Taipei, Taiwan; Asian-Pacific Weed Science Society, No. 1:127-131
Cook CDK; Gut BJ; Rix EM; Schneller J; Seitz M, 1974. Water Plants of the World: A Manual for the Identification of the Genera of Freshwater Macrophytes. The Hague, The Netherlands: Dr W Junk.
Holttum RE, 1954. A revised flora of Malaysia. Vol. II. Singapore: Government Printing Office.
Khaw CH; Ho NK, 1988. Some aspects on the use of paraquat in rice cultivation with special reference to the Muda area. Proceedings of the National Seminar and Workshop on Rice Field Weed Management, 153-167
Kostermans AJGH; Wirjahardja S; Dekker RJ, 1987. The weeds: description, ecology and control. Weeds of rice in Indonesia [edited by Soerjani, M.; Kostermans, A.J.G.H.; Tjitrosoepomo, G.] Jakarta, Indonesia; Balai Pustaka, 24-565
Launert E, 1968. A monographic study of the genus Marsilea. Linnaeus. I. The species of Africa and Madagascar. Senckenbergiana Biologica, 49:273-315.
Merlier H; Montagu J, 1982. Adventices Tropicales. ORSTOM-GERDAT-ENSH.
Pancho JV; Soerjani M, 1978. Aquatic weeds of Southeast Asia. A systematic account of common Southeast Asian aquatic weeds. Aquatic weeds of Southeast Asia. A systematic account of common Southeast Asian aquatic weeds., 130 pp.; [260 X 180 mm].
Peudpaichit S; Tongchairawewat P; Simagrai M, 1987. Londax + BAS 514..H: a broadspectrum weed control treatment in rice. Proceedings, 11th Asian Pacific Weed Science Society Conference Taipei, Taiwan; Asian Pacific Weed Science Society, No. 2:437-448
Rodgers JA, 1993. Aquatic four leaf clovers - Marsilea spp. Aquatics, 15:10-13.
Sculthorpe CD, 1967. The Biology of Aquatic Vascular Plants. London, UK: Edward Arnold Publications Limited.
Smith GM, 1938. Cryptogamic Botany II. Bryophytes and Pteridophytes. New York, USA: McGraw-Hill.
Waterhouse DF, 1993. The Major Arthropod Pests and Weeds of Agriculture in Southeast Asia. ACIAR Monograph No. 21. Canberra, Australia: Australian Centre for International Agricultural Research, 141 pp.
Distribution MapsTop of page
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