Cyclosorus parasiticus (parasitic maiden fern)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Host Plants and Other Plants Affected
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Impact Summary
- Economic Impact
- Environmental Impact
- Threatened Species
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Cyclosorus parasiticus (L.) Farw.
Preferred Common Name
- parasitic maiden fern
Other Scientific Names
- Christella parasitica (L.) H. Lév.
- Dryopteris parasitica (L.) Kuntze
- Polypodium parasiticum L.
- Thelypteris fadenii Fosberg & Sachet
- Thelypteris parasitica (L.) Fosberg
Summary of InvasivenessTop of page
Cyclosorus parasiticus (=Christella parasitica) is a fern that is widespread throughout tropical Asia and some of the Polynesian islands, and also found in a few other parts of the tropics; its spores are likely to be spread long distances by the wind. It is not certain that it was introduced anywhere by humans, but this is thought to be probably the case in Hawaii (although the means of introduction is not known); it is regarded there as a threat to some native plants (sources differ as to the conservation significance of some of these). Throughout its range it can be a weed of plantations, but it is not a regulated pest anywhere. It is sometimes used medicinally.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Pteridophyta
- Class: Filicopsida
- Family: Thelypteridaceae
- Genus: Cyclosorus
- Species: Cyclosorus parasiticus
Notes on Taxonomy and NomenclatureTop of page
The genus Christella was subsumed into the large genus Cyclosorus along with several other genera in recent revisions (He and Zhang, 2012; Smith et al., 2006). Christella parasitica (L.) H. Lév. is a synonym of Cyclosorus parasiticus (L.) Farw. from the basionym Polypodium parasiticum L. (Evenhuis and Eldredge, 2011; Palmer, 2003). It appears that the generic name could still change as phylogenetic work on the group continues. Palmer (2003) indicates that derivation of the specific name parasitica/parasiticus ‘is obscure’.
DescriptionTop of page
An account of ferns in Thailand (Lindsay and Middleton, 2013) provides a succinct description of this species and some congeners:
Rhizome short to long creeping, about 8 mm diam.; scales narrow, up to 12 by 1 mm, pale to dark brown, with a few hairs at margin. Stipes 17-51 cm long, scaly at base, hairy throughout. Laminae oblong-lanceolate, acute at apex, 25-56 by 8-26 cm; 17-22 pairs of free pinnae, basal pinnae deflexed, lower pinnae patent or ascending, linear-lanceolate, sessile at acroscopic base, 4.3-16 by 0.9-2 cm, lobed more than half-way towards costa; segments oblong, oblique, rounded at apex, entire; thinly papyraceous, yellow-green to green, hairy on under surface; basal pairs of veins anastomosing, the other veinlets running to margin of lobes, glandular; glands sessile, rod-shaped, orange to red. Sori supramedial, usually not on lobes, i.e. on one or two basal veinlets; indusia persistent, hairy.
This species is easily confused with Christella dentata (=Cyclosorus dentatus) a species with which it hybridizes (Wagner, 1950). Living plants of Cyclosorus parasiticus are paler, with a duller green, and its proximal pinnae are narrower. Most importantly Cyclosorus parasiticus fronds are not dimorphic and they do not have the sharp break between sterile and fertile fronds that Christella dentata has. Christella dentata’ssterile fronds are broader, shorter, and more spreading than the others (and generally entirely without sori). A key to Chinese species of the subgenus Cyclosoriopsis (which includes both Cyclosorus parasiticus and Christella dentata) is provided by Li et al. (2013).
Plant TypeTop of page
DistributionTop of page
C. parasiticus is widespread throughout tropical Asia and some of the Polynesian islands. It is also recorded from Kenya, Uganda, Madagascar and St. Helena. The only place where it is regarded as probably (although not certainly) introduced is Hawaii, this being based on the record of collections which all occurred after 1926 (Holttum, 1976; Wagner, 1950; Whistler, 1998).
This species is more tropical in its range than its more widespread relative C. dentatus (=Thelypteris dentata or Christella dentata) which extends to subtropical and warm temperate areas free of frost. Andrew Darlow (Invasive Species Project Officer, St. Helena, personal communication, 2013) states that C. parasiticus is probably native in St. Helena, and common there above 500 m in altitude -- earlier reports had suggested it could be introduced there (Holttum, 1976)
Various sources (GBIF, 2012; Holttum, 1976; Lindsay and Middleton, 2013) indicate that the species is present in New Zealand (NZ), but the Flora of New Zealand and other New Zealand sources do not support this (Allan, 1961; Brownsey and Smith-Dodsworth, 1989; Howell and Sawyer, 2006).
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.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Saint Helena||Present, Widespread||Native||Probably native; Original citation: A. Darlow, Invasive Species Project officer, St. Helena, personal communication, 2013|
|China||Present||Present based on regional distribution.|
|India||Present, Widespread||Native||Invasive||NE and southern India. Weed of areca nut plantations and wrongly suspected of being toxic to cows|
|Indonesia||Present||Present based on regional distribution.|
|-Sumatra||Present, Localized||Native||Growing as understory in oil palm plantations|
|-Bonin Islands||Present, Widespread||Native||Common with other weedy plants|
|-Peninsular Malaysia||Present, Widespread||Native|
|Thailand||Present||Native||Reported as a weed in plantations|
|United States||Present||Present based on regional distribution.|
|-Hawaii||Present, Widespread||Introduced||1926||Invasive||Probably introduced. All islands -almost always co-occurring with C. dentatus [Christella dentata] with which it hybridizes|
|Australia||Present||Present based on regional distribution.|
|Federated States of Micronesia||Present||Native|
|Papua New Guinea||Present||Native|
|Pitcairn||Present, Localized||Native||Rare in understory of forests and roadsides. Sometimes grown in gardens|
History of Introduction and SpreadTop of page
Cyclosorus parasiticus is widespread throughout tropical Asia and some of the Polynesian islands, and also found in a few other parts of the tropics; it is likely to have reached many places by wind dispersal of spores. It is not certain that it was introduced to any location by humans, although the case for this having happened in Hawaii is relatively strong, being based on the record of collections which all occurred after 1926 (Holttum, 1976; Wagner, 1950; Whistler, 1998). The amount of early fern collecting in Hawaii is not described in available references, so this could be a reflection of collector effort; but scientists in Hawaii provided the best case for the species being an introduced invasive alien, though the pathway is not known – possibly it was introduced accidentally or as an ornamental.
Andrew Darlow (Invasive Species Project Officer, St. Helena, personal communication, 2013) states that C. parasiticus is probably native in St. Helena, and common there above 500 m in altitude -- earlier reports had suggested it could be introduced there (Holttum, 1976).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Hawaii||before 1926||Yes||No||Wagner (1950)||Probably introduced, based on herbarium collection records|
Risk of IntroductionTop of page
C. parasiticus is an effective disperser that has arrived on some oceanic islands via natural means, e.g. Samoa, Fiji, Guam, Norfolk Island and St. Helena. Only in Hawaii is the case made fairly well that it is not native. On balance the most likely human-mediated pathway for this plant’s introduction is accidental introduction via the horticultural trade. It is hard to rule out its possible introduction as an ornamental but it is not widely traded now as such. Once established in a region it will easily spread via wind-dispersed spores.
HabitatTop of page
C. parasiticus grows well in mesic forest edges, disturbed sites, gullies, humid tropical plantations and orchards (Wagner, 1950).
Habitat ListTop of page
|Terrestrial||Managed||Cultivated / agricultural land||Present, no further details||Natural|
|Terrestrial||Managed||Managed forests, plantations and orchards||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Managed||Managed forests, plantations and orchards||Present, no further details||Natural|
|Terrestrial||Managed||Disturbed areas||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Managed||Disturbed areas||Present, no further details||Natural|
|Terrestrial||Managed||Rail / roadsides||Present, no further details||Natural|
|Terrestrial||Natural / Semi-natural||Natural forests||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural forests||Present, no further details||Natural|
|Terrestrial||Natural / Semi-natural||Riverbanks||Present, no further details||Natural|
|Terrestrial||Natural / Semi-natural||Scrub / shrublands||Present, no further details||Natural|
Host Plants and Other Plants AffectedTop of page
|Areca catechu (betelnut palm)||Arecaceae||Main|
|Elaeis guineensis (African oil palm)||Arecaceae||Main|
|Phyllostegia mollis (Waianae Range phyllostegia)||Lamiaceae||Wild host|
|Phyllostegia parviflora (smallflower phyllostegia)||Lamiaceae||Wild host|
|Pteris lydgatei (Lidgate's brake)||Pteridaceae||Wild host|
|Schiedea hookeri (sprawling schiedea)||Caryophyllaceae||Wild host|
Biology and EcologyTop of page
There is only a limited amount of information available about the biology of this species.
The chromosome number of C. parasiticus is 72 and plants are tetraploid, according to Manton and Sledge (1954). The Missouri Botanical Garden (2014) indicates chromosome numbers of 36 and 72 for both gametophyte and sporophyte. According to Li et al. (2013), many different ploidy levels and hybrids have been detected. Other phylogenetic work has been done on the group to distinguish interspecific variation and generic differences in the family; the DNA data from that study is posted online as per current publication practices in the field (He and Zhang, 2012).
C. parasiticus has a typical fern life cycle. It shows intergametophytic mating behaviour and capacity for intragametophytic selfing, which was considered to contribute to its wide distribution in a variety of niches (Khare and Kaur, 1979). Isolated plants were just as good at producing sporophytes as paired plants, indicating that a single plant can produce viable offspring.
ClimateTop of page
|A - Tropical/Megathermal climate||Preferred||Average temp. of coolest month > 18°C, > 1500mm precipitation annually|
|Af - Tropical rainforest climate||Preferred||> 60mm precipitation per month|
|Am - Tropical monsoon climate||Preferred||Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))|
Natural enemiesTop of page
Notes on Natural EnemiesTop of page
One study identified a new fungal species Cercospora christellae growing on Christella parasitica (i.e. Cyclosorus parasiticus) in Thailand (To-anun et al., 2010) but the authors made no claims as to specificity, suggesting only that it could be a potential biocontrol for what they described as a weedy fern.
Means of Movement and DispersalTop of page
There is no clear evidence for how C. parasiticus has been introduced. The movement of horticultural products for the nursery trade is likely to spread it as a contaminant of the soil in which other plants are planted, as has been the case with some congeners (Murakami et al., 2007). No evidence could be found of it being sold as an ornamental in its own right, although it is an attractive fern and most likely occurs in Hawaiian gardens from time to time even if it is not intentionally planted. No evidence is available for its dispersal by people or their tools, vehicles or other vectors, although there is no reason to expect that this does not occur. It is a common plant along forested trails and roads throughout its range, which suggests a possible association as a result of habitat preference and/or dispersal.
Natural dispersal (non-biotic)
C. parasiticus has dispersed widely via natural means (most likely via wind dispersal of spores) in the tropics to Africa, Asia and oceanic islands in the Pacific and the Atlantic (Copeland, 1929; Holttum, 1976; Lindsay and Middleton, 2013; Whistler, 1998).
Vector transmission (biotic)
No evidence could be found of animal dispersal of C. parasiticus, though its tiny spores could easily hitchhike on animals. It is a common colonizer of disturbed sites in Hawaii and could easily be transported by pigs, e.g. in mud.
Pathway CausesTop of page
|Horticulture||Presumed cause. An attractive fern, but also likely to contaminate pots with other plants||Yes||Yes||Murakami et al. (2007)|
|Medicinal use||Presumed cause. Medicinal uses known -- see text||Yes||Yes|
|Nursery trade||Presumed cause. Congeners known to be transported between greenhouses as contaminants||Yes||Yes||Murakami et al. (2007)|
|People foraging||Presumed cause. Medicinal plant||Yes||Yes||Srivastava (2009)|
Pathway VectorsTop of page
|Clothing, footwear and possessions||Presumed vector. Often along trails||Yes||Yes||Lindsay and Middleton (2013); Wagner (1950)|
|Germplasm||Presumed vector. Congeners known to be transported between greenhouses||Yes||Yes||Murakami et al. (2007)|
|Land vehicles||Likely mode of spore dispersal||Yes||Yes|
|Soil, sand and gravel||Likely mode of spore dispersal||Yes||Yes|
|Water||Likely mode of spore dispersal||Yes|
|Wind||Arrived on oceanic islands via spore dispersal||Yes||Holttum (1976)|
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Growing medium accompanying plants|
Impact SummaryTop of page
Economic ImpactTop of page
It appears that C. parasiticus can be a significant component of forest understories, betelnut plantations (Areca catechu) and oil palm (Elaeis guineensis) plantations, citrus orchards and weedy disturbed gulches in wet to mesic habitats (Germer, 2003; Shridhar and Varadaraju, 2009; To-anun et al., 2010; Wagner, 1950). It was thought to be toxic to livestock but toxicity tests on rabbits suggest that is not the case; it must though occur in some pastures where it could compete with pasture plants (Shridhar and Varadaraju, 2009). It is mentioned as a weed much more rarely than its relative Christella dentata (=Cyclosorus dentatus) in the Global Compendium of Weeds by Randall (2012).
Environmental ImpactTop of page
In Hawaii, where C. parasticus is believed to have arrived prior to 1930 and spread to wet and mesic sites in open forests and disturbed areas (Wagner, 1950), it is regarded as a threat to biodiversity. It may be found in weedy gulches, along with other weedy ferns like Christella dentata (=Cyclosorus dentatus) and the hybrid that it forms with this species. Some government reports mention the fern along with a suite of competing non-native species that impact the endangered species Phyllostegia mollis Benth., Phyllostegia parviflora (Gaudich.) Benth., Schiedea hookeri A.Gray and Pteris lidgatei (Baker) H. Christ (U.S. Fish and Wildlife Service, 2008, 2009a, 2009b, 2011).
In that context this invader might be regarded as relatively benign, considering the relatively serious threats posed by woody weeds, grasses, and some larger ferns such as Cyathea cooperi (=Sphaeropteris cooperi) (Durand and Goldstein, 2001; Kraus and Duffy, 2010). C. parasiticus is more widespread and abundant than its relative Christella dentata (=Cyclosorus dentatus) in Hawaii (Hank Oppenheimer, Pacific Cooperative Studies Unit, University of Hawaii, Hilo, Hawaii, USA, personal copmmunication, 2013). Clearly it is one more element that may contribute to the degradation of rare plant habitat in Hawaii. Since it colonizes disturbed sites preferentially, it may be an early element in sites where introduced ungulates open up habitat and, together with other weedy colonizers, may exacerbate the effects of these ungulates (Smith, 1985). Also, it can probably establish and spread after large-scale disturbance by hurricanes.
Mascaro et al. (2008) found that the hybrid with Christella dentata (also introduced) was dominant in 2.66% of forest plots on the island of Hawaii, suggesting the hybrid is a stable and significant component of the plant community there. Considering the difficulty in distinguishing the species that make up this hybrid, it appears likely that all the threats attributed to any of the weedy Cyclosorus spp. in Hawaii could be due to Christella dentata, Cyclosorus parasiticus or the hybrid.
In an unpublished study on the island of Kauai (Hawaii), Ruth Aguraiuja (Tallinn Botanic Garden, Tallinn, Estonia, personal communication, 2013) found that the soil spore bank was highly dominated by non-native fern species, including Pityrogramma austroamericana [Pityrogramma calomelanos var. austroamericana], Adiantum hispidulum, Cyclosorus parasiticus, Cyclosorus dentatus (=Christella dentata), Cyclosorus intermedius, Blechnum appendiculatum and Nephrolepis multiflora. In many cases the distribution of native groundcover species varied from missing to very sparse and patchy, with dense patches (coverage 70-100%) of non-native species – ferns and others -- which quickly fill the gaps created by ungulates. The domination sequence of these species varied between sites, but these were the top species. Some replicates showed that the hybrid (Cyclosorus x intermedius) between Cyclosorus parasiticus and Christella dentata may have a tendency to dominate over its parent species. However, by their growth habit and life history characteristics, species like Adiantum hispidulum, Blechnum appendiculatum and Nephrolepis are the strongest competitors, capable of outcompeting not only groundcover species but also the juveniles of canopy species. Relatively dense patches of Cyclosorus were also found in the transitional zone from mesic to wet forest, including the areas above Kalalau Rim (a wet mountainous area on the island of Kauai).
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Phyllostegia mollis (Waianae Range phyllostegia)||USA ESA listing as endangered species||Hawaii||Competition; Competition - shading; Competition - smothering||US Fish and Wildlife Service (2009b)|
|Phyllostegia parviflora (smallflower phyllostegia)||NatureServe; USA ESA listing as endangered species||Hawaii||Competition - shading; Competition - smothering; Competition (unspecified); Ecosystem change / habitat alteration||US Fish and Wildlife Service (2008)|
|Schiedea hookeri (sprawling schiedea)||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - monopolizing resources; Competition - shading; Competition - smothering; Ecosystem change / habitat alteration||US Fish and Wildlife Service (2011)|
|Pteris lidgatei (Lidgate's brake)||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - monopolizing resources; Competition - shading; Competition - smothering||US Fish and Wildlife Service (2009a)|
|Platydesma rostrata||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - monopolizing resources; Ecosystem change / habitat alteration; Hybridization||US Fish and Wildlife Service (2010a)|
|Pritchardia hardyi (Makaleha pritchardia)||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - smothering||US Fish and Wildlife Service (2010b)|
|Pritchardia viscosa (stickybud pritchardia)||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - smothering||US Fish and Wildlife Service (1998); US Fish and Wildlife Service (2010b)|
|Stenogyne purpurea (purplefruit stenogyne)||CR (IUCN red list: Critically endangered); USA ESA listing as endangered species||Hawaii||Competition - smothering||US Fish and Wildlife Service (2010b)|
Risk and Impact FactorsTop of page
- Invasive in its native range
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Is a habitat generalist
- Pioneering in disturbed areas
- Tolerant of shade
- Long lived
- Fast growing
- Has high reproductive potential
- Ecosystem change/ habitat alteration
- Modification of successional patterns
- Monoculture formation
- Negatively impacts agriculture
- Negatively impacts livelihoods
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Competition - monopolizing resources
- Competition - shading
- Competition - smothering
- Competition (unspecified)
- Interaction with other invasive species
- Rapid growth
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
UsesTop of page
C. parasiticus is taken as an anthelmintic (Burkill, 1995), and has been investigated for antifungal and antibacterial properties (Ma et al., 2010; Paul et al., 2011; Srivastava, 2009) as well as other medicinal uses (Fang et al., 2008; Hunyadi et al., 2014). There are many other references to medicinal use, mainly from India and Asia.
Uses ListTop of page
- Source of medicine/pharmaceutical
Detection and InspectionTop of page
A useful key is due to be provided on a website describing ferns of Thailand (Lindsay and Middleton, 2013). Excellent descriptions and photos are already posted for the ferns of Thailand and include this species and several congeners. Wagner (1950) and Wagner et al. (1999) list both species; a useful key is provided and the article about naturalized ferns has detailed description of how to distinguish the species in the field and from herbarium specimens. See also the book about Hawaiian ferns (Palmer, 2003).
Similarities to Other Species/ConditionsTop of page
This species is easily confused with Christella dentata (=Cyclosorus dentatus), a species with which it hybridizes (Wagner, 1950). Living plants of Cyclosorus parasiticus are paler, with a duller green, and its proximal pinnae are narrower. Most importantly Cyclosorus parasiticus fronds are not dimorphic and they do not have the sharp break between sterile and fertile fronds that Christella dentata has. Christella dentata’s sterile fronds are broader, shorter, and more spreading than the others (and generally entirely without sori).
A key to Chinese species of the subgenus Cyclosoriopsis (including both Cyclosorus parasiticus and Christella dentata) is provided by Li et al. (2013).
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
It is not certain that this plant was introduced accidentally or intentionally to any location by humans, although the case for Hawaii seems relatively strong. If it was introduced, or were to be introduced elswehere, the most likely pathway for its spread to new areas is as a contaminant of horticultural plant products. Unregulated import of live plant material to isolated locations, sites with high value biodiversity or important agricultural industry is not recommended. Obviously public education and quarantine measures are warranted generally to prevent the introduction of unwanted plants in any jurisdiction (IUCN, 2000; Wittenberg and Cock, 2001). With long-distance dispersal (hundreds of kilometres) being possible via dispersal of its spores, as evidenced by its likely unassisted arrival on various oceanic Atlantic and Pacific islands, this is a species that may be difficult or impossible to contain once established in a region.
If control at a site is warranted e.g., to protect important biodiversity or agricultural resources, hand weeding or treatment with herbicide are known to work. The application of triclopyr applied at standard foliar rates, or at high concentrations with biodiesel to the rhizome, has been known to work (Jane Beachy, O'ahu Army Natural Resources Program, University of Hawaii, Hawaii, USA, personal communication, 2013). According to Ruth Aguraiuja (Tallinn Botanic Garden, Tallinn, Estonia, personal communication, 2013), the viability of Cyclosorus spores in the soil reduced remarkably within a year, and the species could be removed relatively efficiently from sites with endangered species if hand weeding were consistently carried out, as it is intolerant of disturbance to the rhizomes. Non-target impacts from chemical and manual control methods can be expected if the control method is inadvertently applied to co-occurring non-target species.
Gaps in Knowledge/Research NeedsTop of page
If more genetic work were done it might be possible to determine whether populations in Hawaii represent recent human introductions, and perhaps the source populations could be ascertained. Similarly the plant’s status as a native (or not) on other oceanic islands could potentially be ascertained. Also the evolutionary relationships could be clarified between widespread (arguably “weedy”) congeners that share a recent comment ancestor, especially Christella dentata/Cyclosorus dentatus, Christella hispidula/Cyclosorus hispidulus and Cyclosorus parasiticus (cf. He and Zhang, 2012).
ReferencesTop of page
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Khare PB, Kaur S, 1979. Mating behaviour in the homosporous fern, Cyclosorus parasiticus (Linn.) Tardieu. Proceedings of the Indian Academy of Sciences, 88:243-247. http://rd.springer.com/article/10.1007%2FBF03052180
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22/04/2013 Original text by:
Chris Buddenhagen, Florida State University, Tallahassee, Florida, USA
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