Nephrolepis hirsutula
(sword fern)

Pictures

Picture	Title	Caption	Copyright
Title Fronds Caption Nephrolepis hirsutula (sword fern); fronds, upper and lower surfaces. Note the small 'ear-lobe' at the base of each leaflet. Tonga. September 2007. Copyright ©Tauʻolunga/via wikipedia - CC BY-SA 3.0	Fronds	Nephrolepis hirsutula (sword fern); fronds, upper and lower surfaces. Note the small 'ear-lobe' at the base of each leaflet. Tonga. September 2007.	©Tauʻolunga/via wikipedia - CC BY-SA 3.0
Title Frond Caption Nephrolepis hirsutula (sword fern); close view of frond lower surface. Note the small 'ear-lobe' at the base of each leaflet. Tonga. September 2007. Copyright ©Tauʻolunga/via wikipedia - CC BY-SA 3.0	Frond	Nephrolepis hirsutula (sword fern); close view of frond lower surface. Note the small 'ear-lobe' at the base of each leaflet. Tonga. September 2007.	©Tauʻolunga/via wikipedia - CC BY-SA 3.0

Identity

Preferred Scientific Name

• Nephrolepis hirsutula (G. Forst.) C. Presl

Preferred Common Name

• sword fern

International Common Names

• English: Asian sword fern; scaly sword fern; scurfy sword fern

Local Common Names

• American Samoa: vao tuâniu
• Cook Islands: maire; tuartou; tu-rautou; turei’aua; turoutou
• Cuba: helecho; penquita; puntero
• New Zealand: lau maile kimoa; maile kimoa
• Philippines: an anam am; sipeng; tarotaro

Summary of Invasiveness

Nephrolepis hirsutula is a fern native to Asia and the Pacific, and is not recorded as having spread outside of this region. N. hirsutula is reported as an invasive species is in Cuba, however this is due to a misidentification of Nephrolepis brownii. N. brownii, which is considered an aggressive invasive species, is an Asian fern species that has been introduced in the New World. Within its range in Southeast Asia, Australia and the Pacific, N. hirsutula is not considered invasive.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Pteridophyta
•             Class: Filicopsida
•                 Family: Nephrolepidaceae
•                     Genus: Nephrolepis
•                         Species: Nephrolepis hirsutula

Notes on Taxonomy and Nomenclature

Ferns are the informal name given to a diverse group of vascular plants with over 10,500 species (Christenhusz and Chase 2014). Most living fern species are leptosporiangiate, defined as ferns having the sporangia developing from a single cell and producing a definite number of spores (Schuettpelz and Pruer, 2007).

Nephrolepis is a pantropical genus of approximately 19 species (Hovenkamp and Miyamoto 2005), and it is probably the most popular source of commercially used fern species (Yahaya et al., 2016). The name comes from the Greek ‘nephros’ (kidney) and ‘lepis’ (scale), referring to the shape of the indusia in some of the species (Bell, 2017).

The identification of some species has been confusing; being difficult in immature fronds and further complicated by the hybridisation reported occurring mostly in cultivation (Yahaya et al., 2016; Bell, 2017). The species name N. hirsutula has not been consistently applied across the literature, mostly due to incorrect use and confusion with N. brownii. N. hirsutula has been misidentified as N. brownii, or both have been treated as a single species (Hovenkamp and Miyamoto, 2005; Caluff and Regalado, 2011) Also, synonyms of other species have been wrongly applied to N. hirsutula (Missouri Botanical Garden, 2018; USDA-ARS, 2018). Hovenkamp and Miyamoto (2005) provide keys to the identification of the Nephrolepis species and also give detailed information of the characters used to distinguish between some of the species. Based on molecular studies, N. hirsutula and N. brownii are not closely related (Hennequin et al., 2010; Yahaya et al., 2016).

According to Christenhusz et al. (2011) and later Vernon and Ranker (2013) the family placement of Nephrolepis is an unresolved issue, suggesting a tentative use of Nephrolepidaceae. Christenhusz and Chase (2014) suggest placing the Nephrolepidaceae within the subfamily Lomariopsidoideae, and in the Polypodiaceae family. As this classification is not currently being widely accepted, the Nephrolepidaceae family is used in this compendium.

Description

The following description is from Hovenkamp and Miyamoto (2005):

Plants forming tufts of 4–8 fronds (or more). Runners often forming stilts supporting the erect rhizome, 1–2 mm thick, branching angle narrow. scales on runners sparse or dense, appressed or spreading. Tubers absent. Fronds 90–210 by 15–20 cm, stipe 32–80 cm long. Lamina base reduced, tapering over 15–30 cm (or more), basal pinnae 3.5–8 cm long, 2–5 cm distant, middle pinnae straight (sometimes recurved). Sterile pinnae 8–10 by 1.2–1.6 cm, herbaceous, thick, base strongly unequal, basiscopic base truncate or rounded, acroscopic base truncate, strongly auricled (with narrow auricle), margin in basal part entire, towards apex crenate, apex acute. Fertile pinnae 7.5–11 by 0.9–1.1 cm, more strongly crenate than the sterile pinnae. Indument basal scales peltate, appressed, 1.5–23.5 by 1 mm, central part dark brown or blackish, shining, hyaline margin wide, ciliate, marginal glands, absent apex obtuse, rachis scales very dense, with a well-developed protracted acumen, spreading or squarrose, rufous, acumen strongly dentate. Scales on lamina usually persistent. Hairs absent from lamina and costae. Sori submarginal to nearly medial, 24–33 pairs on fully fertile pinnae, round, not impressed. Indusium reniform, with open sinus, attached at sinus.

Plant Type

Distribution

According to Hovenkamp and Miyamoto (2005) N. hirsutula occurs in the southeast of Asia, throughout Malesia, to Australia and the Pacific, and distribution records elsewhere are due to a misidentification of N. brownii. The species has been wrongly reported as occurring in North America, Central America, South America and the Caribbean (Oviedo Prieto et al., 2012; García-Lahera, 2016; Encyclopedia of Life, 2018; Missouri Botanical Garden, 2018; USDA-ARS, 2018).There is some debate about the extent of the native region of the species at some of the Pacific region, being regarded as both native and introduced in Hawaii (Moomaw and Takahashi, 1960; Vernon and Ranker, 2013)

The wrongful attribution of synonyms is also adding to the confusion in the distribution reported for N. hirsutula. For example, in Missouri Botanical Garden (2018) some of the synonyms listed for N. hirsutula are synonyms of N. exaltata or N. brownii. In USDA-ARS (2018) GRIN online database N. multiflora is used as a synonym for N. hirsutula.

Nephrolepis hirsutula is recorded as as an invasive species in Cuba by Oviedo Prieto et al. (2012), but this is due to a misidentification of N. brownii, which is considered an aggressive invasive species and has been introduced in the Caribbean (Kairo et al., 2003). N. brownii is not listed as invasive by Oviedo Prieto et al. (2012). The authors cite Florida Exotic Pest Plant Council (2009) for N. hirsutula as being invasive in Florida, USA. Neither the FLEPPC invasive plant species list cited or the most recent one for 2019 (Florida Exotic Pest Plant Council, 2019) list N. hirsutula as an invasive species for Florida. The Nephrolepis species appearing in the list are N.brownii (= N. multiflora) and N. cordifolia. In an invasive report for N. hirsutula in Cuba, Caluff and Regalado (2011) report the synonyms of the species as being N. multiflora and Davallia multiflora, which are synonyms of N. brownii (Hovenkamp and Miyamoto, 2005; The Plant List, 2013). 

Distribution Table

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.

History of Introduction and Spread

According to Hovenkamp and Miramoto (2005) the species is found only in its native range and it is not spreading into other regions. There are spore records from 10,000 years ago from what is believed to be N. hirsutula from Sumatra, Indonesia (Morley, 1982).

Risk of Introduction

Without further details about the biology, the environmental requirements and the possible effects over non-native habitats, it is difficult to assess the risk of introduction. However, with the use of other Nephrolepis species as ornamentals, their spread facilitated by its stoloniferous vegetative reproduction and the easiness of spore dissemination it is highly possible that N. hirsutula could spread from its native area and slowly progress its way into suitable areas.

Habitat

N. hirsutula is reported as growing in grasslands, forest edges, open forests, ridges, scrublands, rainforests, and low elevations (Ridley, 1926; Kingston and Waldren, 2003; Banaticla and Buot, 2005; Bell, 2017). It is also reported at margins of freshwater lakes in Papua New Guinea (Chambers, 1987).

Habitat List

Biology and Ecology

Genetics

Molecular work shows that N. hirsutula is more closely related to N. biserrata than to N. brownii (Yahaya et al., 2016). The authors also report the species as being a tetraploid (2n= 82, 164), although Manton and Sledge (1954) report the species as being diploid (n=41).

Nephrolepis hirsutula has been suggested as one of the parent species of the hybrid Nephrolepis exaltata var. bostoniensis (Encyclopedia of Life, 2018). However molecular studies by Yahaya et al. (2016) found that N. hirsutula did not contribute to this hybrid species.

Reproductive Biology

Although no details are given for the reproduction of N. hirsutula, it is highly likely this species reproduces vegetatively through stolons or by spores, as most of the other Nephrolepis species (Yahaya et al. 2016).

Environmental Requirements

There is little detailed information available for the environmental requirements of N. hirsutula. The species grows on clay banks near water or in crevices of rock outcrops in Australia (Bell, 2017; 2018). In Papua New Guinea it is reported as growing in lowlands, in infertile, serpentine soils and in organic alluvium; with a mean annual rainfall of 3000 – 4000 mm (Takeuchi, 2003).

In Guangdong, China, it is reported in red soils in subtropical areas with an annual average temperature of 22°C and an annual rainfall of 2380 mm (Wang et al., 2006). In Hawaii, the species is reported as growing in nutrient poor soils, gravelly silty clay and volcanic soils with high infiltration rate; in an average temperature of 23°C, with a range between 13°-90°C; and in an annual precipitation of 1250 to 3810 mm with a relative high humidity (Moomaw and Takahashi, 1960).

Climate

Latitude/Altitude Ranges

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free

Soil texture

• heavy
• medium

Special soil tolerances

• infertile

Natural enemies

Means of Movement and Dispersal

Natural Dispersal

As other Nephrolepis species, N. hirsutula dispersal is likely through its stolons or by its lightweight spores.

Pathway Causes

Pathway Vectors

Impact Summary

Risk and Impact Factors

• Has a broad native range
• Has high reproductive potential
• Gregarious
• Reproduces asexually

• Highly likely to be transported internationally accidentally
• Difficult to identify/detect as a commodity contaminant

Uses

Social Benefit

Godwin and Harvey (1944) report N. hirsutula is used as food, but give no further details. The scales from the midrib are used in treating the scab of male circumcisions (Whistler 1990). The fronds are sometimes used for leis, house decorations, and mulch (Whistler, 1988). The fronds are worn on the head as protection from sun or rain and used by women as carrying-pads (Bodner and Gereau, 1988).

Environmental Services

The fronds of N. hirsutula are reported as habitat for mites, without causing any damage (Huang, 1991). It is eaten by feral goats (Capra hircus) in Raoul Island, New Zealand (Parkes, 1984). N. hirsutula is reported as being capable of arsenic accumulation (Wang et al., 2006).

Uses List

Environmental

• Amenity
• Wildlife habitat

General

• Sociocultural value

Human food and beverage

• Vegetable

Materials

• Mulches

Medicinal, pharmaceutical

• Traditional/folklore

Similarities to Other Species/Conditions

Hovenkamp and Miyamoto (2005) provide keys for the identification of the Nephrolepis species. N. hirsutula is mostly confused with N. brownii. N. brownii has the sori close to the margin while in N. hirsutula the sori are less close to the margin to nearly medial. N. hirsutula has dense, rufous rachis scales with a very strongly dentate acumen. N. brownii has usually sparser, paler rachis scales, with a longer, nearly always entire narrow apex. The upper costae in N. hirsutula is glabrous.

N. hirsutula can be distinguished from N. biserrata as the latter is not densely rusty wooly, but scaly. N. cordifolia and N. exaltata pinnules are oblong to oblong-lanceolate and acute to round at the apex, while N. hirsutula pinnules are linear-lanceolate, and attenuate toward the apex (Fosberg, 1943).

Gaps in Knowledge/Research Needs

Molecular studies are needed to assess generic and family delimitations in ferns (Smith et al., 2006; Christenhusz et al., 2011). More information on the biology and the environmental requirements for N. hirsutula is needed. Herbaria collections should be verified, and field work is needed to better assess the distribution of the species.

References

Banaticla, M. C. N., Buot, I. E., Jr., 2005. Altitudinal zonation of pteridophytes on Mt. Banahaw de Lucban, Luzon Island, Philippines. Plant Ecology, 180(2), 135-151. http://springerlink.metapress.com/link.asp?id=100328 doi: 10.1007/s11258-004-2494-7

Bell GH, 2017. Nephrolepis hirsutula. In: Flora of Australia, Canberra, Australia: Australian Biological Resources Study, Department of the Environment and Energy.https://profiles.ala.org.au/opus/foa/profile/Nephrolepis%20hirsutula

Bodner, C. C., Gereau, R. E., 1988. A contribution to Bontoc ethnobotany. Economic Botany, 42(3), 307-369. doi: 10.1007/BF02860159

Brownsey PJ, 2001. New Zealand’s pteridophyte flora - plants of ancient lineage but recent arrival?. Brittonia, 53(2), 307-369.

Caluff MG, Regalado L, 2011. Plantas invasoras Vol. 16: N. hirsutula. Cuba: GEF/UNDP.http://repositorio.geotech.cu/jspui/bitstream/1234/1653/16/Folleto%20informativo%2016_Nephrolepis%20hirsutula.pdf

Chambers MR, 1987. The freshwater lakes of Papua New Guinea: an inventory and limnological review. Journal of Tropical Ecology, 3(1), 1-23.

Christenhusz MJM, Chase MW, 2014. Trends and concepts in fern classification. Annals of Botany, 113(4), 571-594.

Christenhusz MJM, Zhang X-C, Schneider H, 2011. A linear sequence of extant lycophytes and ferns. (Phytotaxa). 19, 7-54.

Encyclopedia of Life, 2018. Encyclopedia of Life. In: Encyclopedia of Life . http://www.eol.org

Florida Exotic Pest Plant Council, 2009. Florida EPPC's 2009 list of invasive plant species. USA: Center for Invasive Species and Ecosystem Health.https://www.fleppc.org/list/2009/List-WW-F09-final.pdf

Florida Exotic Pest Plant Council, 2019. Florida EPPC's 2019 list of invasive plant species. In: Florida EPPC's 2019 list of invasive plant species Tifton, Georgia, USA: Center for Invasive Species and Ecosystem Health.https://www.fleppc.org/list/list.htm

Fosberg FR, 1943. Notes on plants of the Pacific Islands-III, Bulletin of the Torrey Botanical Club 70(4), 386-397.

García-Lahera JP, 2016. (Flora ruderal sobre las edificaciones del centro histórico en las ciudades de Trinidad y Sancti Spíritus, Cuba central). Revista del Jardín Botánico Nacional, 37, 103-113.

Godwin H, Harvey LA, 1944. Annual meeting in the University Department of Botany, Oxford, 5-6 January 1944. Journal of Ecology, 32(1), 131-134.

Hennequin, S., Hovenkamp, P., Christenhusz, M. J. M., Schneider, H., 2010. Phylogenetics and biogeography of Nephrolepis - a tale of old settlers and young tramps. Botanical Journal of the Linnean Society, 164(2), 113-127. http://onlinelibrary.wiley.com/doi/10.1111/j.1095-8339.2010.01076.x/full doi: 10.1111/j.1095-8339.2010.01076.x

Hovenkamp, P. H., Miyamoto, F., 2005. A conspectus of the native and naturalized species of Nephrolepis (Nephrolepidaceae) in the world. Blumea, 50(2), 279-322.

Huang, K. W., 1991. Three new eriophyoid mites recovered from ferns in Taiwan (Acarina: Eriophyoidea). Chinese Journal of Entomology, 11(4), 324-329.

Iwatsuki K, 1973. Phytogeography of the pteridophytes in Peninsular Thailand. American Fern Journal, 63(3), 129-134.

Kairo, M., Ali, B., Cheesman, O., Haysom, K., Murphy, S., 2003. Invasive species threats in the Caribbean region. Report to the Nature Conservancy. In: Invasive species threats in the Caribbean region. Report to the Nature Conservancy . Curepe, Trinidad and Tobago: CAB International.132 pp. http://www.issg.org/database/species/reference_files/Kairo%20et%20al,%202003.pdf

Kerr AM, 2000. Defoliation of an island (Guam, Mariana Archipelago, Western Pacific Ocean) following a saltspray-laden ‘dry’ typhoon. Journal of Tropical Ecology, 16(6), 895-901.

Kingston, N., Waldren, S., 2003. The plant communities and environmental gradients of Pitcairn Island: the significance of invasive species and the need for conservation management. Annals of Botany, 92(1), 31-40. doi: 10.1093/aob/mcg106

Maciel S, Pietrobom MR, 2010. (Dryopteridaceae e Lomariopsidaceae (Polypodiopsida) do Campo Experimental da Embrapa Amazônia Oriental, Moju, Pará, Brasil). Rodriguésia, 61(3), 405-414.

Manton I, Sledge WA, 1954. Observations on the cytology and taxonomy of the pteridophyte flora of Ceylon. Philosophical Transactions of the Royal Society of London, Series B, Biological Series, 238(654), 127-185.

Missouri Botanical Garden, 2018. Tropicos database. In: Tropicos database St. Louis, Missouri, USA: Missouri Botanical Garden.http://www.tropicos.org/

MOOMAW, J. C., TAKAHASHI, M., 1960. Vegetation on gibbsitic soils in Hawaii. Journal of the Arnold Arboretum, 41(3), 391-411.

Morley RJ, 1982. A palaeoecological interpretation of a 10,000 year pollen record from Danau Padang, Central Sumatra, Indonesia. Journal of Biogeography, 9(2), 151-190.

Morton CV, Lellinger DB, 1967. Notes on the ferns of Dominica and St. Vincent. American Fern Journal, 57(2), 66-77.

Oviedo Prieto, R., Herrera Oliver, P., Caluff, M. G., et al., 2012. National list of invasive and potentially invasive plants in the Republic of Cuba - 2011. (Lista nacional de especies de plantas invasoras y potencialmente invasoras en la República de Cuba - 2011). Bissea: Boletín sobre Conservación de Plantas del Jardín Botánico Nacional de Cuba, 6(Special Issue No. 1), 22-96.

Parkes JP, 1984. Feral goats on Raoul Island II. Diet and notes on the flora. New Zealand Ecological Society, 7, 95-101.

Ramírez-Barahona S, Torres-Miranda A, Palacios-Ríos M, Luna-Vega I, 2009. Historical biogeography of the Yucatán Peninsula, Mexico: a perspective from ferns (Monilophyta) and lycopods (Lycophyta). Biological Journal of the Linnean Society, 98(4), 775-786.

Ridley HN, 1926. The ferns of the Malay Peninsula. Journal of the Malayan Branch of the Royal Asiatic Society, 4(1), 1-121.

Schuettpelz E, Pryer K, 2007. Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon, 56(4), 1037-1050.

Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG, 2006. A classification for extant ferns. (Taxon). 55, 705-731.

Sykes WR, 1981. The vegetation of Late, Tonga. Allertonia, 2(6), 323-353.

Takeuchi W, 2003. Botanical summary of a lowland ultrabasic flora in Papua New Guinea. SIDA contributions to botany, 20(4), 1491-1559.

The Plant List, 2013. The Plant List: a working list of all plant species. Version 1.1. In: The Plant List: a working list of all plant species. Version 1.1 Richmond, London, UK: Royal Botanic Gardens, Kew.http://www.theplantlist.org

USDA-ARS, 2018. Germplasm Resources Information Network (GRIN). Online Database. In: Germplasm Resources Information Network (GRIN). Online Database Beltsville, Maryland, USA: National Germplasm Resources Laboratory.https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

Vernon AL, Ranker TA, 2013. Current status of the ferns and lycophytes of the Hawaiian Islands. American Fern Journal, 103(2), 59-111.

Wang, H. B., Ye, Z. H., Shu, W. S., Li, W. C., Wong, M. H., Lan, C. Y., 2006. Arsenic uptake and accumulation in fern species growing at arsenic-contaminated sites of southern China: field surveys. International Journal of Phytoremediation, 8(1), 1-11. doi: 10.1080/16226510500214517

Whistler WA, 1990. Ethnobotany of the Cook Islands: the plants, their Maori names, and their uses. Allertonia, 5(4), 347-424.

Whistler, W. A., 1984. Annotated list of Samoan plant names. Economic Botany, 38(4), 464-489. doi: 10.1007/BF02859089

Whistler, W. A., 1988. Ethnobotany of Tokelau: the plants, their Tokelau names, and their uses. Economic Botany, 42(2), 155-176. doi: 10.1007/BF02858917

Yahaya, N. H., Stech, M., Zonneveld, B. J. M., Hovenkamp, P. H., 2016. What is Nephrolepis 'bostoniensis'?: unravelling the origin of Nephrolepis hybrids and cultivars with molecular data. Scientia Horticulturae, 204, 153-160. http://www.sciencedirect.com/science/journal/03044238

Distribution References

Banaticla M C N, Buot I E Jr, 2005. Altitudinal zonation of pteridophytes on Mt. Banahaw de Lucban, Luzon Island, Philippines. Plant Ecology. 180 (2), 135-151. http://springerlink.metapress.com/link.asp?id=100328 DOI:10.1007/s11258-004-2494-7

Bell GH, 2017. Nephrolepis hirsutula. Canberra, Australia: Australian Biological Resources Study, Department of the Environment and Energy. https://profiles.ala.org.au/opus/foa/profile/Nephrolepis%20hirsutula

Brownsey PJ, 2001. New Zealand’s pteridophyte flora - plants of ancient lineage but recent arrival? Brittonia. 53 (2), 307-369.

CABI Data Mining, Undated. CAB Abstracts Data Mining.,

CABI, Undated. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Caluff MG, Regalado L, 2011. Plantas invasoras Vol. 16: N. hirsutula., Cuba: GEF/UNDP. http://repositorio.geotech.cu/jspui/bitstream/1234/1653/16/Folleto%20informativo%2016_Nephrolepis%20hirsutula.pdf

García-Lahera JP, 2016. (Flora ruderal sobre las edificaciones del centro histórico en las ciudades de Trinidad y Sancti Spíritus, Cuba central). Revista del Jardín Botánico Nacional. 103-113.

Huang K W, 1991. Three new eriophyoid mites recovered from ferns in Taiwan (Acarina: Eriophyoidea). Chinese Journal of Entomology. 11 (4), 324-329.

Iwatsuki K, 1973. Phytogeography of the pteridophytes in Peninsular Thailand. American Fern Journal. 63 (3), 129-134.

Kerr AM, 2000. Defoliation of an island (Guam, Mariana Archipelago, Western Pacific Ocean) following a saltspray-laden ‘dry’ typhoon. Journal of Tropical Ecology. 16 (6), 895-901.

Kingston N, Waldren S, 2003. The plant communities and environmental gradients of Pitcairn Island: the significance of invasive species and the need for conservation management. Annals of Botany. 92 (1), 31-40. DOI:10.1093/aob/mcg106

Maciel S, Pietrobom MR, 2010. (Dryopteridaceae e Lomariopsidaceae (Polypodiopsida) do Campo Experimental da Embrapa Amazônia Oriental, Moju, Pará, Brasil). Rodriguésia. 61 (3), 405-414.

Manton I, Sledge WA, 1954. Observations on the cytology and taxonomy of the pteridophyte flora of Ceylon. Philosophical Transactions of the Royal Society of London, Series B, Biological Series. 238 (654), 127-185.

Moomaw J C, Takahashi M, 1960. Vegetation on gibbsitic soils in Hawaii. Journal of the Arnold Arboretum. 41 (3), 391-411.

Morley RJ, 1982. A palaeoecological interpretation of a 10,000 year pollen record from Danau Padang, Central Sumatra, Indonesia. Journal of Biogeography. 9 (2), 151-190.

Morton CV, Lellinger DB, 1967. Notes on the ferns of Dominica and St. Vincent. American Fern Journal. 57 (2), 66-77.

Oviedo Prieto R, Herrera Oliver P, Caluff M G, et al, 2012. National list of invasive and potentially invasive plants in the Republic of Cuba - 2011. (Lista nacional de especies de plantas invasoras y potencialmente invasoras en la República de Cuba - 2011). Bissea: Boletín sobre Conservación de Plantas del Jardín Botánico Nacional de Cuba. 6 (Special Issue No. 1), 22-96.

Ramírez-Barahona S, Torres-Miranda A, Palacios-Ríos M, Luna-Vega I, 2009. Historical biogeography of the Yucatán Peninsula, Mexico: a perspective from ferns (Monilophyta) and lycopods (Lycophyta). Biological Journal of the Linnean Society. 98 (4), 775-786.

Ridley HN, 1926. The ferns of the Malay Peninsula. Journal of the Malayan Branch of the Royal Asiatic Society. 4 (1), 1-121.

Sykes WR, 1981. The vegetation of Late, Tonga. Allertonia. 2 (6), 323-353.

Takeuchi W, 2003. Botanical summary of a lowland ultrabasic flora in Papua New Guinea. SIDA contributions to botany. 20 (4), 1491-1559.

USDA-ARS, 2018. Germplasm Resources Information Network (GRIN). Online Database. In: Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

Vernon AL, Ranker TA, 2013. Current status of the ferns and lycophytes of the Hawaiian Islands. American Fern Journal. 103 (2), 59-111.

Whistler W A, 1984. Annotated list of Samoan plant names. Economic Botany. 38 (4), 464-489. DOI:10.1007/BF02859089

Whistler W A, 1988. Ethnobotany of Tokelau: the plants, their Tokelau names, and their uses. Economic Botany. 42 (2), 155-176. DOI:10.1007/BF02858917

Links to Websites

Contributors

16/07/2018 Original text by:

Jeanine Vélez-Gavilán, UPR-RUM, Mayagüez, Puerto Rico

Distribution Maps