Invasive Species Compendium

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Datasheet

Adiantum hispidulum
(rosy maidenhair fern)

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Datasheet

Adiantum hispidulum (rosy maidenhair fern)

Summary

  • Last modified
  • 13 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Adiantum hispidulum
  • Preferred Common Name
  • rosy maidenhair fern
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Pteridophyta
  •       Class: Filicopsida
  •         Family: Pteridaceae
  • Summary of Invasiveness
  • A. hispidulum, the rosy maidenhair fern, is known to be invasive and weedy in the main Hawaiian Islands where it has escaped from cultivation. In Hawaii it is a serious weed generally of mesic slopes and gulch...

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Pictures

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PictureTitleCaptionCopyright
Adiantum hispidulum (rosy or rough maidenhair fern); habit. Kula Botanical Garden, Maui, Hawaii, USA. March, 2011.
TitleHabit, showing fronds
CaptionAdiantum hispidulum (rosy or rough maidenhair fern); habit. Kula Botanical Garden, Maui, Hawaii, USA. March, 2011.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Adiantum hispidulum (rosy or rough maidenhair fern); habit. Kula Botanical Garden, Maui, Hawaii, USA. March, 2011.
Habit, showing frondsAdiantum hispidulum (rosy or rough maidenhair fern); habit. Kula Botanical Garden, Maui, Hawaii, USA. March, 2011.©Forest Starr & Kim Starr - CC BY 4.0
Adiantum hispidulum (rosy or rough maidenhair fern); habit. Pohakuokala Gulch, Maui, Hawaii, USA. August, 2005.
TitleHabit
CaptionAdiantum hispidulum (rosy or rough maidenhair fern); habit. Pohakuokala Gulch, Maui, Hawaii, USA. August, 2005.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Adiantum hispidulum (rosy or rough maidenhair fern); habit. Pohakuokala Gulch, Maui, Hawaii, USA. August, 2005.
HabitAdiantum hispidulum (rosy or rough maidenhair fern); habit. Pohakuokala Gulch, Maui, Hawaii, USA. August, 2005.©Forest Starr & Kim Starr - CC BY 4.0
Adiantum hispidulum (rosy or rough maidenhair fern); habit. Olinda, Maui, Hawaii, USA. November 15, 2009.
TitleHabit
CaptionAdiantum hispidulum (rosy or rough maidenhair fern); habit. Olinda, Maui, Hawaii, USA. November 15, 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Adiantum hispidulum (rosy or rough maidenhair fern); habit. Olinda, Maui, Hawaii, USA. November 15, 2009.
HabitAdiantum hispidulum (rosy or rough maidenhair fern); habit. Olinda, Maui, Hawaii, USA. November 15, 2009.©Forest Starr & Kim Starr - CC BY 4.0

Identity

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Preferred Scientific Name

  • Adiantum hispidulum Sw.

Preferred Common Name

  • rosy maidenhair fern

International Common Names

  • English: rough maidenhair fern

Local Common Names

  • Australia: five-fingered Jack
  • New Zealand: rosy maidenhair

EPPO code

  • ADIHI (Adiantum hispidulum)

Summary of Invasiveness

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A. hispidulum, the rosy maidenhair fern, is known to be invasive and weedy in the main Hawaiian Islands where it has escaped from cultivation. In Hawaii it is a serious weed generally of mesic slopes and gulch bottoms and is often abundant along intermittent and perennial streams. It is capable of invading intact plant communities and pristine areas. A. hispidulum usually begins to colonize in areas where there is some type of natural disturbance such as landslides, tree falls, disturbance by feral ungulates, or even a single dislodged rock. The dense clumps and rhizome mats of this fern prevent establishment of many native taxa including rare species. A. hispidulum can also overrun other ferns and herbs (Wilson, 1996; Palmer, 2003; H. Oppenheimer, Hawaii Plant Extinction Prevention Program (PEP), USA, personal communication, 2013). It is naturalized locally in a few other parts of the world including the southeastern USA and parts of its native range including eastern and southern Africa, Malaya, and Singapore.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Pteridophyta
  •             Class: Filicopsida
  •                 Family: Pteridaceae
  •                     Genus: Adiantum
  •                         Species: Adiantum hispidulum

Description

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Terrestrial ferns, plants stiff, erect. Rhizomes erect to decumbent, short-creeping, with dark brown scales. Fronds clustered at apex of rhizome, erect, (6-)15–60 cm tall, young fronds rosy pink; stipes dark brown, rough, up to 30 cm long, clothed with short dark fibrils and hairs; blades ± fan-shaped, deltate to ovate, dichotomously branched at 45º angle into 7–15 branches; pinnules asymmetrically oblong-rectangular to diamond shaped, 0.5–1.7 cm long, 3–8 mm wide, the upper and outer margins gently rounded, finely toothed, the lower straight and entire, softly pubescent to ± glabrate, veins mostly ending in marginal teeth; pinnule stalks 0.5–1 mm long. Sori 6–14 per segment, small, closely placed on upper and outer edges in notches between the lobes; indusium flaps 3–4(–5) mm wide, circular to broadly oblong or kidney-shaped, covered with numerous small, pointed brown hairs (Verdcourt, 2002; Palmer, 2003).

Distribution

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A. hispidulum is widespread and presumed to be native from Australia and Africa to Asia, Malesia and various Pacific Islands (Palau, Fiji, New Caledonia, Vanuatu), and eastward to Polynesia (Austral Islands (Rapa), Society Islands (Tahiti), Cook Islands (Raratonga), and New Zealand. However, it is sporadic throughout its native range, and in addition to native plant communities it often occurs in disturbed habitats. For example around villages, along trails, on roadside banks, rock walls, or in the understory of forestry plantings, suggesting a potential for invasiveness. Australia is the type locality and presumably the centre of diversity for A. hispidulum, as it occurs in all States and Territories. Bostock et al. (1998) recognize four taxonomic varieties in Australia. The widespread var. hispidulum is terrestrial or lithophytic in a variety of habitats, including open forest and as relictual populations in gorges in arid areas. The other three varieties are said to be more localized and endemic to Australia. In this treatment, however, a single variable species is recognized without infraspecific taxa (subspecies, varieties, or forms), following International Plant Names Index (IPNI) (2013) and Tropicos (2013).

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

IndonesiaPresentPresent based on regional distribution.
-JavaWidespreadNative Not invasive Holttum, 1954On steep earth banks in shady places
MalaysiaPresentPresent based on regional distribution.
-Peninsular MalaysiaLocalisedNative Not invasive Holttum, 1954Malaya, uncommon and never seen in the wild
PhilippinesPresentNative Not invasive Copeland, 1958Luzon
SingaporeLocalisedNative Not invasive Holttum, 1954On bare shaded earth banks around suburbs, uncommon, presumably native
Sri LankaWidespreadNative Not invasive Fraser-Jenkins et al., 2006Shady roadside banks, steep earth banks, 'jungle', tea plantations, 160-1675 m
TaiwanPresentNativeKnapp, 2011

Africa

ComorosPresentNative Not invasive Tardieu-Blot, 1958Mayotte, Anjouan, Grand Comore
EthiopiaPresentNative Not invasive Schelpe and Anthony, 1986
KenyaPresentNative Not invasive Schelpe and Anthony, 1986
MadagascarPresent, few occurrencesNative Not invasive Tardieu-Blot, 1958West Madagascar: Haut Bemarivo; Analamomby
MalawiLocalisedNative Not invasive Schelpe, 1970
MauritiusWidespreadNative Not invasive Lorence, 1976; Badre and Lorence, 2008Saxicolous, sometimes in soil, often in shade on talus, rocky walls, waterfalls, roadside banks, dry thickets and dry forest; widespread, to 1000 m
MayottePresentNative Not invasive Tardieu-Blot, 1958Mayotte, Anjouan, Grande Comore
MozambiqueLocalisedNative Not invasive Schelpe, 1970
RéunionWidespreadNative Not invasive Badre and Lorence, 2008Saxicolous: sometimes in soil, often in shade on talus or rocky walls, waterfalls, roadside banks, dry thickets and dry forest; widespread to 1000 m
Rodriguez IslandWidespreadNative Not invasive Lorence, 1976; Badre and Lorence, 2008Saxicolous: sometimes in soil, often in shade on talus or rocky walls, waterfalls, roadside banks, dry thickets and dry forest; widespread to 1000 m
South AfricaLocalisedNative Not invasive Schelpe and Anthony, 1986In Natal at 830 m where it may have escaped from cultivation. Also naturalised in a few localities on the Cape Peninsula
TanzaniaLocalisedNative Not invasive Schelpe and Anthony, 1986

North America

USAPresentPresent based on regional distribution.
-ConnecticutLocalisedIntroducedParis, 1993Introduced, sporadic escapes from cultivation, possibly naturalised locally. Banks and old walls; 0-100 m
-FloridaLocalisedIntroducedParis, 1993Introduced, sporadic escapes from cultivation, possibly naturalised locally. Banks and old walls; 0-100 m
-GeorgiaLocalisedIntroducedParis, 1993Introduced, sporadic escapes from cultivation, possibly naturalised locally. Banks and old walls; 0 - 100 m
-HawaiiWidespreadIntroduced Invasive Wilson, 1996; Palmer, 2003; Weller et al., 2010Widely naturalised on all main islands (not recorded from Niihau). First collected in wild on Oahu in 1923. Naturalised in native mesic forest on Kauai, forming dense mats
-LouisianaLocalisedIntroducedParis, 1993Introduced, sporadic escapes from cultivation, possibly naturalised locally. Banks and old walls; 0 - 100 m

Oceania

AustraliaWidespreadNative Not invasive Bostock et al., 1998All states and Territories of mainland Australia. Four varieties recognised. Var. hipsidulum is terrestrial or lithophytic in a variety of habitats, including open forest and relictual populations in gorges, arid areas. The other three endemic varieties are more localised
Cook IslandsPresent, few occurrencesNativeNational Tropical Botanical Garden, 2013; PIER, 2013Raratonga: locally common on shaded road cuts
FijiWidespreadNative Not invasive Brownlie, 1977; Kato et al., 2008Indigenous, common in forests of the dry and intermediate zones, also in very well-drained light forest of higher rainfall areas
French PolynesiaPresent, few occurrencesNative Not invasive PIER, 2013Rapa, lithophytic on basalt cliffs. Tahiti, on cliff faces
New CaledoniaWidespreadNative Not invasive Brownlie, 1969; Kato et al., 2008Abundant, especially on dry talus slopes and open sunny areas
PalauPresent, few occurrencesNativeHillman et al., 2008
Solomon IslandsPresentNativeJSTOR, 2013Bougainville
VanuatuPresent, few occurrencesNativeKato et al., 2008

History of Introduction and Spread

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A. hispidulum has become widely naturalized and weedy in Hawaii, where it grows in dry to mesic, sunny and rocky slopes, woods or trails on all of the main islands (not recorded from Niihau). Its date of introduction is unknown, but it was not recorded by Hillebrand (1888). It was first collected in the wild on Oahu in 1923 (Wilson, 1996).

In continental North America (USA) it is introduced and represented in the flora by sporadic escapes from cultivation and possibly naturalized locally (Paris, 1993) in Connecticut and Georgia. It is also reported from Florida and Louisiana, where it usually occurs on banks and old walls, 0-100 m elevation.

This species is sporadic throughout east tropical and temperate Africa. In East Africa it is confined to moist shaded habitats in South Africa (Cape Province, Natal, and Transvaal), Mozambique, Malawi, Kenya, Tanzania, and Ethiopia (Schelpe and Anthony, 1986). In Natal, it has been recorded at 830 m altitude where it may have escaped from cultivation, and it has also been found naturalized in a few localities on the Cape Peninsula.

It is uncommon in Malaya and not seen in the wild, or in primary forests, but only near villages (Holttum, 1954). In Java it is found throughout, usually in the lower hilly regions, on steep earth banks in shady places. In Singapore it is presumably native and occurs on bare but shaded earth banks around the suburbs of the city but it is not common (Holttum, 1954).

Risk of Introduction

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A. hispidulum is sporadic throughout its native range, but is known to escape from cultivation via spores. A. hispidulum may colonize native plant communities as well as disturbed habitats such as around villages, along trails, on roadside banks, rock walls, or in the understory of forestry plantings. This species is potentially invasive in tropical and subtropical moist to wet environments and most likely spreads from cultivated plants by spore dispersal.

Habitat

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A. hispidulum occurs in dry to mesic, sunny to shady, usually moist rocky slopes, cliffs, woods, gulch bottoms, trails, roadside banks, rock walls, and understory of forestry plantings and native forest.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
 
Terrestrial – ManagedManaged forests, plantations and orchards Present, no further details
Rail / roadsides Present, no further details
Terrestrial ‑ Natural / Semi-naturalNatural forests Present, no further details
Rocky areas / lava flows Present, no further details

Biology and Ecology

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Genetics

Based on chromosome counts of material from Australia and Ceylon, A. hispidulum is an apogamous hexaploid (has six sets of chromosomes), with a chromosome number of n = ca. 170-180 (Manton and Sledge, 1954). The base chromosome number in A. hispidulum appears to be 29 rather than 30, the usual base number for the genus (Brownlie, 1977).

Reproductive Biology

The apogamous (asexual) life cycle of certain ferns means that they produce 32, larger than normal diploid spores per sporangium, instead of the normal 64 haploid spores (Moran, 2004). Other apogamous taxa with higher polyploid chromosome numbers (triploid, tetraploid, hexaploid, etc.) cannot reproduce sexually either. When their spores germinate the resulting prothallus (gametophyte or gamete producing phase in a fern’s life cycle) does not produce sex organs but proliferates vegetatively without fertilization. The resulting sporophyte (spore producing) plantlet grows to maturity and in turn produces fronds with apogamous spores. Apogamous reproduction is advantageous in ferns that grow in dry habitats for two reasons: firstly apogamous ferns lack swimming sperm and there is no need for water in reproduction; and secondly their prothallus matures faster than those of sexually reproducing ferns (Moran, 2004). Apogamous ferns generally have wider geographic distributions than their sexually reproducing relatives. This appears to be the case for A. hispidulum, thus explaining its wide distribution, success in colonizing dry to mesic, disturbed habitats, and invasive tendencies.

Physiology and Phenology

Apogamy may explain the wide distribution, success in colonizing dry to mesic, disturbed habitats, and invasive tendencies of A. hispidulum. In addition to this the growth rate of all life stages of A. hispidulum is much greater than those of native fern species on Hawaiian islands.

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])
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
22 30

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall1800mm; lower/upper limits

Means of Movement and Dispersal

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Spores of A. hispidulum and other fern species are transported and dispersed primarily by air currents and wind. Spores may be transported in the soil on shoes or stuck to fur of animals but this method of spore dispersal is unlikely. The spores require a moist, shady, stable environment to germinate and grow into gameophytes and eventually into sporophytes or the spore producing phase. Many species of Adiantum, including A. hispidulum, are grown horticulturally and may spread unintentionally from cultivated plants.

Impact Summary

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CategoryImpact
Environment (generally) Negative

Environmental Impact

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Impact on Habitats

This species can cause habitat degradation by forming dense mats that outcompete or prevent the establishment and growth of native plants (Weller et al., 2010).

Impact on Biodiversity

The dense mats formed by this fern species can smother, exclude, or prevent establishment of native plant species, including at least five federally listed endangered species; Asplenium dielpallidum, Delissea kauaiensis, Euphorbia haeleeleana, Isodendrion laurifolium and Psychotria hobdyi (Weller et al., 2010). Other thretened and federally listed endangered species affected by A. hispidulum in Hawaii include Diellia erectaDiellia manniiCtenitis squamigera and Remya mauiensis (R. Aguraiuja, Tallin Botanical Garden, Estonia, personal communication, 2015).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Asplenium dielpallidumNo DetailsHawaiiCompetition - monopolizing resources; Competition - shading; Competition - smothering; Rapid growthWeller et al., 2010
Delissea kauaiensisUSA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resources; Competition - shading; Competition - smothering; Rapid growthWeller et al., 2010
Euphorbia haeleeleanaUSA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resources; Competition - shading; Competition - smothering; Rapid growthWeller et al., 2010
Isodendrion laurifoliumUSA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resources; Competition - shading; Competition - smothering; Rapid growthWeller et al., 2010
Psychotria hobdyi (Hobdy's wild-coffee)USA ESA listing as endangered species USA ESA listing as endangered species; USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resources; Competition - shading; Competition - smothering; Rapid growthWeller et al., 2010
Poa mannii (Mann's bluegrass)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resourcesUS Fish and Wildlife Service, 2010a
Pritchardia munroi (Kamalo pritchardia)No DetailsHawaiiCompetition - smotheringUS Fish and Wildlife Service, 2011a
Remya mauiensis (Maui remya)USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetitionUS Fish and Wildlife Service, 2009
Schiedea apokremnos (Kauai schiedea)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetitionUS Fish and Wildlife Service, 2010b
Schiedea hookeri (sprawling schiedea)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resources; Ecosystem change / habitat alterationUS Fish and Wildlife Service, 2011b

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
  • Benefits from human association (i.e. it is a human commensal)
  • Fast growing
  • Has high reproductive potential
  • Gregarious
  • Reproduces asexually
Impact outcomes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts cultural/traditional practices
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
  • Negatively impacts animal/plant collections
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - shading
  • Competition - smothering
  • Competition
  • Interaction with other invasive species
  • 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/costly to control

Detection and Inspection

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This species can be detected by visual identification, facilitated by using a reference book (Heath and Chinnock, 1974) or key (Palmer, 2003).

Similarities to Other Species/Conditions

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A. hispidulum may be distinguished from related maidenhair species (e.g. A. capillus-veneris, A. aethiopicum, A. raddianum) by it fronds that repeatedly fork at 45º angles, stipes clothed with short, dark fibrils and hairs, and pinnules with short, light brown hairs.

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.

Prevention
The cultivation and sale of A. hispidulum in the horticultural trade should be discouraged and the public should be made aware of its invasive tendencies.
 
Control
Manual eradication of A. hispidulum is possible and probably the only effective control measure if conducted repeatedly in accordance with their  growth and regeneration cycles. Pulling up plants destabilizes the soil, and new plants may become established at the same sites by germination of spores remaining in the soil.
 
Control of A. hispidulum is extremely difficult due to transport of spores by air currents and wind over long distances. In addition to this, A. hispidulum has high spore production, high germination and rapid growth rates.
 
Monitoring and Surveillance
A. hispidulum can be detected by visual identification and surveillance on the ground. 
 
Ecosystem restoration
A. hispidulum must be manually removed and eradicated to allow ecosystem recovery.

References

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Badré F, Lorence D, 2008. Adiantum. In: Flore des Mascareignes, Adiantacées [ed. by Bosser, J. \Badré, F. \Guého, J.]. Paris, France: Institut du recherche pour le dévelopment, Mauritius Sugar Industry Research Institute and Royal Botanic Gardens (Kew), 102-110.

Bostock PD, Farrant PA, Parris BS, 1998. Adiantum. In: Flora of Australia, Ferns, Gymnosperms and Allied Groups, 48 [ed. by McCarthy, P. M. \Orchard, A. E.]. Collingwood, Victoria, Australia: Australian Biological Resources Study/CISRO, 253-263.

Brownlie G, 1969. Adiantum. In: Flore de la Nouvelle-Calédonie et Dépendances. 3. Ptéridophytes, 3. Paris, France: Museum National D'Histoire Naturelle, 177-183.

Brownlie G, 1977. The pteridophyte flora of Fiji. Vaduz, Liechtenstein: J. Cramer, 189-193.

Copeland EB, 1958. Fern flora of the Philippines. Manila, Philippines: Bureau of Printing, 161.

Fraser-Jenkins CR, Verdcourt B, Walker TG, 2006. Pteridaceae. In: A Revised Handbook of the Flora of Ceylon XV, part B [ed. by Shaffer-Fehre, \M.]. Enfield, New Hampshire, USA: Science Publishers, 349-352.

Heath E, Chinnock RJ, 1974. Ferns and fern allies of New Zealand. Wellington, NZ: Reed AH and Reed, AW, 12-13.

Hillebrand W, 1888. Flora of the Hawaiian Islands. London, UK: Williams & Norgate, 633-634.

Hillman Kitalong AE, DeMeo R, Holm T, 2008. Native trees of Palau: a field guide. Koror, Palau: The Environment Inc, 220.

Holttum RE, 1954. A revised flora of Malaya, Singapore: Government Printing Office, 603-604.

International Plant Names Index (IPNI), 2013. International Plant Names Index. http://www.ipni.org/

JSTOR, 2013. Global Plants. https://plants.jstor

Kato M, Takehisa N, Matsumoto S, Ebihara A, 2008. Illustrated flora of ferns and fern-allies of South Pacific Islands. Tsukuba, Japan: National Museum of Nature and Science, 110 pp.

Knapp R, 2011. Ferns and fern allies of Taiwan, 352. Taipei, Taiwan: KBCC Press & Yuan-Liou Publishing, 433.

Lorence DH, 1976. The pteridophytes of Rodrigues Island. Botanical Journal of the Linnean Society, 72:269-283.

Lorence DH, 1978. The pteridophytes of Mauritius (Indian Ocean): ecology and distribution. Botanical Journal of the Linnean Society, 76:207-247.

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 Sciences, 238:16-180.

Moran RC, 2004. A natural history of ferns. Portland, USA: Timber Press, 38-42.

National Tropical Botanical Garden, 2013. Herbarium Database. http://ntbg.org/herbarium/

Palmer DD, 2003. Hawai`i's ferns and fern allies. Honolulu, Hawaii: University of Hawai`i Press, 42-45.

Paris CA, 1993. Adiantum. In: Flora of North America, 2. New York, USA: Oxford University Press, 125-130.

PIER, 2013. Pacific Islands Ecosystems at Risk. Honolulu, Hawaii, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Schelpe EACLE, 1970. Pteridophyta. In: Flora Zambesiaca [ed. by Exell, A. W. \ Launert, E.]. London, UK: Crown Agents for Oversea Governments and Administrations, 108-112.

Schelpe EACLE, Anthony NC, 1986. Pteridophyta. In: Flora of Southern Africa [ed. by Leistner, O. A.]. Pretoria, South Africa: Botanical Research Institute, Department of Agriculture and Water Supply, 100-101.

Tardieu-Blot ML, 1958. Flore de Madagascar et des Comores [ed. by Humbert, H.]. Paris, France: Museum National d'Histoire Naturelle, 131-132.

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

Tropicos, 2012. Tropicos. Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org

US Fish and Wildlife Service, 2009. In: Remya mauiensis (Maui remya). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 15 pp.

US Fish and Wildlife Service, 2010. In: 5-Year Review, Short Form Summary: Species Reviewed: Poa mannii (Mann's bluegrass). US Fish and Wildlife Service, 10 pp.

US Fish and Wildlife Service, 2010. In: Schiedea apokremnos (maolioli). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 16 pp.

US Fish and Wildlife Service, 2011. In: 5-Year Review, Short Form Summary: Species Reviewed: Pritchardia munroi (lo'ulu). US Fish and Wildlife Service, 11 pp.

US Fish and Wildlife Service, 2011. In: Schiedea hookeri (no common name). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 20 pp.

USDA-ARS, 2013. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

Verdcourt B, 2002. Flora of tropical East Africa. Rotterdam, The Netherlands: A.A. Balkema Publishers.

Weller SG, Cabin RJ, Lorence DH, Perlman S, Wood K, Flynn T, Sakai AK, 2010. Alien plant invasions, introduced ungulates, and alternative states in a mesic forest in Hawaii. Restoration Ecology, 19:671-680.

Wilson KA, 1996. Alien ferns in Hawaii. Pacific Science, 50(2):127-141.

Contributors

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21/12/2013 Original text by:

David H. Lorence, National Tropical Botanical Garden, Hawaii, USA

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