Invasive Species Compendium

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Datasheet

Thelypteris opulenta
(jewelled maiden fern)

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Datasheet

Thelypteris opulenta (jewelled maiden fern)

Summary

  • Last modified
  • 19 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Thelypteris opulenta
  • Preferred Common Name
  • jewelled maiden fern
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Pteridophyta
  •       Class: Filicopsida
  •         Family: Thelypteridaceae
  • Summary of Invasiveness
  • Thelypteris opulenta is a perennial fern, native to tropical Asia and Madagascar, that has been introduced as an ornamental to gardens and amenity areas. It produces numerous spores that are easily dispersed by...

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Identity

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Preferred Scientific Name

  • Thelypteris opulenta (Kaulf.) Fosberg

Preferred Common Name

  • jewelled maiden fern

Other Scientific Names

  • Amblovenatum opulentum J.P. Roux
  • Amphineuron opulentum (Kaulf.) Holttum
  • Aspidium extensum Blume
  • Aspidium extensum Fée
  • Aspidium opulentum Kaulf.
  • Cyclosorus extensus (Blume) H. Itô
  • Cyclosorus opulentus (Kaulf.) Nakaike
  • Dryopteris extensa (Blume) Kuntze
  • Thelypteris extensa (Blume) C.V. Morton

International Common Names

  • Spanish: helecho jade; helechos
  • Chinese: xian mai mao jue

Local Common Names

  • Mexico: helecho palmita

Summary of Invasiveness

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Thelypteris opulenta is a perennial fern, native to tropical Asia and Madagascar, that has been introduced as an ornamental to gardens and amenity areas. It produces numerous spores that are easily dispersed by wind and water. It also spreads via rhizomes, stolons and tubers, which are often dispersed to new areas in dumped garden waste. T. opulenta is known to have escaped from cultivation and grows as a weed in disturbed sites, secondary forests and along roadsides and trails. It is listed as a common weed in moist and wet habitats in Central and South America and has been listed as an invasive species in Peru, Costa Rica (Cocos Island) and Cuba.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Pteridophyta
  •             Class: Filicopsida
  •                 Family: Thelypteridaceae
  •                     Genus: Thelypteris
  •                         Species: Thelypteris opulenta

Notes on Taxonomy and Nomenclature

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Thelypteridaceae is a family of ferns with a cosmopolitan distribution and high species richness. Depending on its circumscription, the number of genera in this family varies from one to approximately 30 and the number of species from 900 to 1000, most of which occur in tropical and subtropical regions of the world (Smith and Cranfill, 2002). In some circumscriptions, Thelypteris constitutes the only genus in the family, containing about 900 species divided into approximately 30 subgenera and/or sections. However, other circumscriptions recognize these subgenera/sections as 30 genera in their own right (Holttum, 1982; Smith, 1992).

The taxonomic status of the species Thelypteris opulenta is still uncertain. For some authors, the species T. opulenta belongs to the genus Cyclosorus and the accepted name is listed as Cyclosorus opulentus. However, for other authors, this species is a synonym of Amblovenatum opulentum (Smith, 1992; Labiak and Prado, 2007; Lindsay and Middleton, 2012; Flora of China Editorial Committee, 2017).

Description

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The following description is from the Flora of China Editorial Committee (2017):

Plants 60-100 cm tall. Rhizomes shortly creeping, apices and bases of stipes with dark brown linear-lanceolate scales. Fronds approximate; stipes ca. 30 cm, brownish; laminae 30-60 x 20-30 cm, bases not narrowed or slightly so, apices caudate; pinnae 10-20 pairs, almost sessile; proximal pair of pinnae slightly shortened; middle pinnae linear-lanceolate, 15-25 x 1-2 cm, bases rounded-truncate (or slightly cuneate on proximal pinnae), lobed 1/2-2/3 toward costae, apices long acuminate; segments 25-40 pairs, middle ones subfalcate, 4-8 x 2-3 mm, entire, subacute to obtuse at apices; veinlets 8-10 pairs, proximal pair anastomosing or only connivent, sometimes next vein running to sinus membrane. Laminae herbaceous, yellowish green when dried, with several acicular hairs along veins on both surfaces, minute hairs and many minute yellow spherical glands along veins abaxially, also minute hairs between veins abaxially. Sori orbicular, submarginal, usually sterile on proximal one or two pairs of veins; indusia glabrous or glandular along margins. Sporangia bearing golden spherical glands on stalks. Spores cristate.

Plant Type

Top of page Herbaceous
Perennial

Distribution

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T. opulenta is native to parts of tropical Africa and Asia (Lindsay and Middleton, 2012; Flora of China Editorial Committee, 2017). It has been introduced and is naturalized in Central America, South America and the Caribbean (Fosberg and Sachet, 1972; Smith, 1992; Smith, 1995; Labiak and Prado, 2007).

Distribution Table

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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

Africa

KenyaPresentFosberg and Sachet (1972)
MadagascarPresentNativeFlora of Madagascar (2017)
MozambiquePresentHyde et al. (2017)Listed as Amblovenatum opulentum; extremely rare
SeychellesPresentFlora of Madagascar (2017)
TanzaniaPresentFosberg and Sachet (1972)

Asia

ChinaPresentNativeCABI (Undated)Present based on regional distribution
-HainanPresentNativeFlora of China Editorial Committee (2017)
IndiaPresentNativeSubhash Chandra et al. (2008); Flora of China Editorial Committee (2017)South India
-Andaman and Nicobar IslandsPresentNativeSubhash Chandra et al. (2008)
IndonesiaPresentNativeFlora of China Editorial Committee (2017)
-JavaPresentFosberg and Sachet (1972)
MalaysiaPresentNativeFlora of China Editorial Committee (2017)
MyanmarPresentNativeFlora of China Editorial Committee (2017)
PhilippinesPresentNativeFlora of China Editorial Committee (2017)
SingaporePresentFosberg and Sachet (1972)
Sri LankaPresentNativeFlora of China Editorial Committee (2017)
ThailandPresentNativeFlora of China Editorial Committee (2017)

North America

BarbadosPresentIntroducedMoreno and Roubik (2017); Hassler (2017)Listed as Thelypteris extensa
Costa RicaPresentIntroducedInvasiveRojas-Alvarado (2011); Smith (1995)Invasive on Cocos Island
CubaPresentIntroducedInvasiveOviedo Prieto et al. (2012)
GuadeloupePresentIntroducedNaturalizedChristenhusz (2009)Naturalized
JamaicaPresentIntroducedNaturalizedSmith (1992)Naturalized
MartiniquePresentIntroducedNaturalizedSmith (1992)Naturalized
NicaraguaPresentIntroducedGómez and Arbeláez (2009)
PanamaPresentIntroducedNaturalizedSmith (1995)Naturalized
Puerto RicoPresentIntroducedLiogier and Martorell (2000)
Saint LuciaPresentIntroducedNaturalizedGraveson (2012)Naturalized
Trinidad and TobagoPresent, WidespreadIntroducedBaksh-Comeau (2000)
United StatesPresentIntroducedCABI (Undated)Present based on regional distribution
-FloridaPresentIntroducedWunderlin et al. (2017)

Oceania

AustraliaPresentLindsay and Middleton (2012)Listed as Cyclosorus opulentus
Federated States of MicronesiaPresentLindsay and Middleton (2012)Listed as Cyclosorus opulentus
French PolynesiaPresentLorence et al. (2011)Listed as Cyclosorus (subg. Amphineuron) opulentus
GuamPresentDonnegan et al. (2004)
Northern Mariana IslandsPresentIntroducedYoshioka (2008)

South America

BoliviaPresentIntroducedLabiak and Prado (2007)
BrazilPresentIntroducedNaturalizedFlora do Brasil (2018)Naturalized
-AcrePresentIntroducedNaturalizedFlora do Brasil (2018); Labiak and Prado (2007)Naturalized
-AmazonasPresentIntroducedNaturalizedFlora do Brasil (2018)Naturalized
-ParaPresentIntroducedNaturalizedFlora do Brasil (2018)Naturalized
ColombiaPresentIntroducedNaturalizedSmith (1995)Naturalized
EcuadorPresentIntroducedNaturalizedSmith (1995)Naturalized
French GuianaPresentIntroducedNaturalizedFunk et al. (2007)Naturalized
GuyanaPresentIntroducedNaturalizedFunk et al. (2007)Naturalized
PeruPresentIntroducedInvasiveAponte (2012); Smith (1992)Invasive in Pasco region
SurinamePresentIntroducedNaturalizedFunk et al. (2007)Naturalized
VenezuelaPresentIntroducedNaturalizedHokche et al. (2008)Naturalized

History of Introduction and Spread

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T. opulenta was intentionally introduced into new areas as an ornamental and as a houseplant. In Peru, where this species is naturalized and extremely common, the oldest herbarium collection is dated from 1956 (Smith, 1992). In Trinidad, T. opulenta appears in herbarium collections dated from 1966 and the species is currently spreading across the island (Baksh-Comeau, 2000). In Puerto Rico, it is part of a herbarium collection from Naguabo dated 1993 (US National Herbarium). By the year 2000, Liogier and Martorell (2000) listed this species as “recently collected” in the rain forest of Luquillo in Puerto Rico.

Habitat

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T. opulenta grows in lowland rain forests, secondary forests, gullies, old plantations and along riverbanks and streams. It also grows as a weed, primarily in disturbed sites, wastelands, urban shrublands and along fence lines, roadsides and trails (Smith, 1992; Baksh-Comeau, 2000; Flora of China Editorial Committee, 2017).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedDisturbed areas Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Natural
Rail / roadsides Present, no further details Harmful (pest or invasive)
Rail / roadsides Present, no further details Natural
Urban / peri-urban areas Present, no further details Harmful (pest or invasive)
Urban / peri-urban areas Present, no further details Natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Present, no further details Natural
Riverbanks Present, no further details Harmful (pest or invasive)
Riverbanks Present, no further details Natural

Biology and Ecology

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Genetics

The chromosome number for T. opulenta is not known, however, for other species within the genus Thelypteris, chromosome number varies from n = 27 to n = 36 (Smith, 1992).

Environmental Requirements

T. opulenta grows in moist and wet habitats at low elevations (100-900 m) and is adapted to grow in both sunny and deep shade sites. It has been recorded as thriving in limestone, volcanic and calcareous soils from pH 5.6 to 7.8 (Lindsay and Middleton, 2012; Hyde et al., 2017).

Climate

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ClimateStatusDescriptionRemark
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]))
As - Tropical savanna climate with dry summer Preferred < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Aw - Tropical wet and dry savanna climate Preferred < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
30 30

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 18 32

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall1200>2000mm; lower/upper limits

Rainfall Regime

Top of page Bimodal
Uniform

Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • acid
  • neutral

Soil texture

  • heavy
  • light
  • medium

Means of Movement and Dispersal

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T. opulenta spreads via spores as well as vegetatively via underground stems (i.e. rhizomes) and often via fleshy underground tubers. Spores are primarily dispersed by wind and water, while rhizomes and tubers are most often dispersed to new areas in dumped garden waste (Smith, 1992; Smith, 1995; Lindsay and Middleton, 2012; Flora of China Editorial Committee, 2017).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
DisturbanceWeed in disturbed sites and secondary forests Yes Yes Smith, 1995
Escape from confinement or garden escapeOften naturalized Yes Yes Smith, 1995
Garden waste disposalRhizomes, spores and tubers Yes Yes
HorticultureOrnamental Yes Yes
Intentional releaseOrnamental Yes Yes
Ornamental purposesCultivated in gardens, parks and as a houseplant Yes Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Debris and waste associated with human activitiesRhizomes, spores and tubers in dumped garden waste Yes Yes
WaterSpores Yes
WindSpores Yes

Impact Summary

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CategoryImpact
Cultural/amenity Positive
Economic/livelihood Positive and negative
Environment (generally) Positive and negative

Environmental Impact

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Under suitable conditions, T. opulenta forms dense stands with the potential to displace native plant species and alter successional patterns in disturbed areas (Smith, 1995; Aponte, 2012; Oviedo Prieto et al., 2012).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Tolerant of shade
  • Highly mobile locally
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Gregarious
  • Reproduces asexually
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Modification of successional patterns
  • Monoculture formation
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - smothering
  • Rapid growth
  • Rooting
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally deliberately
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field

Uses

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T. opulenta is commercialized and cultivated as an ornamental and as a houseplant (Smith, 1995; Flora of China Editorial Committee, 2017).

Uses List

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Environmental

  • Amenity

Ornamental

  • Potted plant

Prevention and Control

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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.

There is no information available for the control of T. opulenta. However, hand pulling for control of small infestations and foliar applications of glyphosate and metsulfuron are both recommended management strategies in Australia and the USA to control areas invaded by ferns e.g. species in the genus Nephrolepis (ISSG, 2017).

References

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Aponte H, 2012. Efectos de las actividades agropecuarias sobre la estructura poblacional de Astrocaryum perangustatum (Arecaceae) en Pozuzo (Pasco-Perú). Cientifica, 9:17-32

Baksh-Comeau YS, 2000. Checklist of the pteridophytes of Trinidad & Tobago. Fern Gazette, 16:111-122

Chandra S, Fraser-Jenkins CR, Kumari A, Srivastava A, 2008. A summary of the status of threatened Pteridophytes of India. Taiwania, 53(2), 170-209. http://tai2.ntu.edu.tw/taiwania

Christenhusz MJ, 2009. Index Pteridophytorum Guadalupensium or a revised checklist to the ferns and club mosses of Guadeloupe (French West Indies). Botanical Journal of the Linnean Society, 161(3):213-277

Donnegan JA, Butler SL, Grabowiecki W, Hiserote BA, Limtiaco D, 2004. Guam's forest resources, 2002. Resource Bulletin - Pacific Northwest Research Station, USDA Forest Service, (No.PNW-RB-243), ii + 32 pp.

Flora do Brasil, 2018. Brazilian Flora 2020 in construction. http://floradobrasil.jbrj.gov.br/reflora/listaBrasil/PrincipalUC/PrincipalUC.do#CondicaoTaxonCP

Flora of China Editorial Committee, 2017. Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

Flora of Madagascar, 2017. Catalogue of the Plants of Madagascar. St. Louis, Missouri, USA and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.tropicos.org/Project/Madagascar

Fosberg FR, Sachet MH, 1972. Three Indo-Pacific Thelypteris species reinterpreted and a new African species described. Smithsonian Contributions to Botany, 8:1-12

Funk V, Hollowell T, Berry P, Kelloff C, Alexander SN, 2007. Checklist of the plants of the Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana). Contributions from the United States National Herbarium, 55, 584 pp.

Gómez LD, Arbeláez AL, 2009. Flora de Nicaragua. Helechos (Tomo IV). In: Stevens WD, Montiel OM, Pool A, eds. Monographs in Systematic Botany from the Missouri Botanical Garden, St. Louis, USA: The Missouri Botanical Garden Press, 116:1-151

Graveson R, 2012. The Plants of Saint Lucia (in the Lesser Antilles of the Caribbean). http://www.saintlucianplants.com

Hassler M, 2017. World Ferns: Checklist of Ferns and Lycophytes of the World (version Nov 2017). In: Roskov Y, Abucay L, Orrell T, Nicolson D, Bailly N, Kirk PM, Bourgoin T, DeWalt RE, Decock W, De Wever A, Nieukerken E van, Zarucchi J, Penev L, eds. 2017. Species 2000 & ITIS Catalogue of Life. Leiden, The Netherlands: Naturalis

Hokche O, Berry PE, Huber O, 2008. Nuevo Catálogo de la Flora Vascular de Venezuela. Caracas, Venezuela: Fundación Instituto Botánico de Venezuela, 859 pp

Holttum RE, 1982. Thelypteridaceae. In: van Steenis CG, Holttum RE, eds. Flora Malesiana, Series II: Pteridophyta (Ferns and fern-Allies), 1(5), 289 pp

Hyde MA, Wursten BT, Ballings P, Coates Palgrave M, 2017. Flora of Mozambique. https://www.mozambiqueflora.com/index.php

ISSG, 2017. Global Invasive Species Database (GISD). Invasive Species Specialist Group of the IUCN Species Survival Commission. http://www.iucngisd.org

Labiak PH, Prado J, 2007. New records of Pteridophytes from Bolivia and Brazil. American Fern Journal, 97(2):113-123

Lindsay S, Middleton DJ, 2012. Ferns of Thailand, Laos and Cambodia. http://rbg-web2.rbge.org.uk/thaiferns/

Liogier AH, Martorell LF, 2000. Flora of Puerto Rico and adjacent islands: a systematic synopsis, 2nd edition revised. San Juan, Puerto Rico: La Editorial, University of Puerto Rico, 382 pp

Lorence DH, Wagner WL, Wood KR, Smith AR, 2011. New pteridophyte species and combinations from the Marquesas Islands, French Polynesia. PhytoKeys. 2011(4):5-51

Moreno E, Roubik D, 2017. Herbario Virtual Austral Americano. https://herbariovaa.org/

Oviedo Prieto R, Herrera Oliver P, Caluff MG, 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 1):22-96

Rojas-Alvarado AF, 2011. New species and new records of ferns (Pteridophyta: Polypodiales) from Cocos Island, Costa Rica. Brenesia, 75-76:7-15

Smith AR, 1992. Thelypteridaceae. In: Tryon RM, Stolze RG, eds. Pteridophyta of Peru Part III. 16. Thelypteridaceae. Chicago, USA: Fieldiana Botany, Field Museum of Natural History, 29:1-80

Smith AR, 1995. Thelypteridaceae. In: Davidse G, Sousa Sánchez M, Knapp S, eds. Flora Mesoamericana. México, D.F, México: Universidad Nacional Autónoma de México, 1:164-195

Smith AR, Cranfill RB, 2002. Intrafamilial relationships of the Thelypteroid ferns (Thelypteridaceae). American Fern Journal, 92(2):131-149

Wunderlin RP, Hansen BF, Franck AR, Essig FB, 2017. Atlas of Florida Plants. Tampa, USA: Institute for Systematic Botany, University of South Florida. http://florida.plantatlas.usf.edu/

Yoshioka JM, 2008. Botanical survey of the War in the Pacific National Historical Park Guam, Mariana Islands. Technical Report 161. Manoa, Hawaii: Pacific Cooperative Studies Unit University of Hawaii. https://scholarspace.manoa.hawaii.edu/bitstream/10125/26985/1/161.pdf

Distribution References

Aponte H, 2012. (Efectos de las actividades agropecuarias sobre la estructura poblacional de Astrocaryum perangustatum (Arecaceae) en Pozuzo (Pasco-Perú)). In: Cientifica, 9 17-32.

Baksh-Comeau YS, 2000. Checklist of the pteridophytes of Trinidad & Tobago. In: Fern Gazette, 16 111-122.

CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

Christenhusz MJ, 2009. Index Pteridophytorum Guadalupensium or a revised checklist to the ferns and club mosses of Guadeloupe (French West Indies). In: Botanical Journal of the Linnean Society, 161 (3) 213-277.

Donnegan J A, Butler S L, Grabowiecki W, Hiserote B A, Limtiaco D, 2004. Resource Bulletin - Pacific Northwest Research Station, USDA Forest Service, Portland, USA: Pacific Northwest Research Station, USDA Forest Service. ii + 32 pp.

Flora do Brasil, 2018. Brazilian Flora 2020 in construction., http://floradobrasil.jbrj.gov.br/reflora/listaBrasil/PrincipalUC/PrincipalUC.do#CondicaoTaxonCP

Flora of China Editorial Committee, 2017. Flora of China. In: Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

Flora of Madagascar, 2017. Catalogue of the Plants of Madagascar., St. Louis, Missouri; Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.tropicos.org/Project/Madagascar

Fosberg FR, Sachet MH, 1972. Three Indo-Pacific Thelypteris species reinterpreted and a new African species described. In: Smithsonian Contributions to Botany, 8 1-12.

Funk V, Hollowell T, Berry P, Kelloff C, Alexander S N, 2007. Contributions from the United States National Herbarium, Washington, USA: Department of Systematic Biology - Botany, National Museum of Natural History, Smithsonian Institution. 55, 584 pp.

Gómez LD, Arbeláez AL, 2009. (Flora de Nicaragua. Helechos (Tomo IV)). In: Monographs in Systematic Botany from the Missouri Botanical Garden, 116 [ed. by Stevens WD, Montiel OM, Pool A]. St. Louis, USA: The Missouri Botanical Garden Press. 1-151.

Graveson R, 2012. The Plants of Saint Lucia (in the Lesser Antilles of the Caribbean). In: The Plants of Saint Lucia (in the Lesser Antilles of the Caribbean), http://www.saintlucianplants.com

Hassler M, 2017. World Ferns: Checklist of Ferns and Lycophytes of the World (version Nov 2017). In: Species 2000 & ITIS Catalogue of Life, [ed. by Roskov Y, Abucay L, Orrell T, Nicolson D, Bailly N, Kirk PM, Bourgoin T, DeWalt RE, Decock W, De Wever A, Nieukerken E van, Zarucchi J, Penev L]. Leiden, The Netherlands: Naturalis.

Hokche O, Berry PE, Huber O, 2008. (Nuevo Catálogo de la Flora Vascular de Venezuela)., Caracas, Venezuela: Fundación Instituto Botánico de Venezuela. 859 pp.

Hyde MA, Wursten BT, Ballings P, Coates Palgrave M, 2017. Flora of Mozambique., https://www.mozambiqueflora.com/index.php

Labiak PH, Prado J, 2007. New records of Pteridophytes from Bolivia and Brazil. In: American Fern Journal, 97 (2) 113-123.

Lindsay S, Middleton DJ, 2012. Ferns of Thailand, Laos and Cambodia., http://rbg-web2.rbge.org.uk/thaiferns/

Liogier AH, Martorell LF, 2000. Flora of Puerto Rico and adjacent islands: a systematic synopsis., San Juan, Puerto Rico: La Editorial, University of Puerto Rico. 382 pp.

Lorence DH, Wagner WL, Wood KR, Smith AR, 2011. New pteridophyte species and combinations from the Marquesas Islands, French Polynesia. In: PhytoKeys, 2011 (4) 5-51.

Moreno E, Roubik D, 2017. Herbario Virtual Austral Americano., https://herbariovaa.org/

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.

Rojas-Alvarado AF, 2011. New species and new records of ferns (Pteridophyta: Polypodiales) from Cocos Island, Costa Rica. In: Brenesia, 75-76 7-15.

Smith AR, 1992. Thelypteridaceae. In: Pteridophyta of Peru Part III. 16. Thelypteridaceae, 29 [ed. by Tryon RM, Stolze RG]. Chicago, USA: Fieldiana Botany, Field Museum of Natural History. 1-80.

Smith AR, 1995. Thelypteridaceae. In: Flora Mesoamericana, 1 [ed. by Davidse G, Sousa Sánchez M, Knapp S]. México, D.F, México: Universidad Nacional Autónoma de México. 164-195.

Subhash Chandra, Fraser-Jenkins C R, Alka Kumari, Archana Srivastava, 2008. A summary of the status of threatened Pteridophytes of India. Taiwania. 53 (2), 170-209. http://tai2.ntu.edu.tw/taiwania

Wunderlin RP, Hansen BF, Franck AR, Essig FB, 2017. Atlas of Florida Plants., Tampa, USA: Institute for Systematic Botany, University of South Florida. http://florida.plantatlas.usf.edu/

Yoshioka JM, 2008. Botanical survey of the War in the Pacific National Historical Park Guam, Mariana Islands. Technical Report 161., Manoa, Hawaii, Pacific Cooperative Studies Unit University of Hawaii. https://scholarspace.manoa.hawaii.edu/bitstream/10125/26985/1/161.pdf

Links to Websites

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WebsiteURLComment
Ferns of the Worldwww.fernsoftheworld.com
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.

Contributors

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16/08/17 Original text by:

Julissa Rojas-Sandoval, Department of Botany-Smithsonian NMNH, Washington DC, USA

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