Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Cyrtomium falcatum
(Japanese holly fern)

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Datasheet

Cyrtomium falcatum (Japanese holly fern)

Index

Summary

  • Last modified
  • 22 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Cyrtomium falcatum
  • Preferred Common Name
  • Japanese holly fern
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Pteridophyta
  •       Class: Filicopsida
  •         Family: Dryopteridaceae
  • Summary of Invasiveness

  • Cyrtomium falcatum, commonly known as Japanese holly fern, is native to Asia (including China and Japan). It has been introduced across the world as an ornamental plant and has spread throughout tropical and subtropical areas, and in temp...

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Pictures

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PictureTitleCaptionCopyright
Cyrtomium falcatum (Japanese holly fern); habit. Old Santee Canal Park, Berkeley County, South Carolina, USA. July, 2015.
TitleHabit
CaptionCyrtomium falcatum (Japanese holly fern); habit. Old Santee Canal Park, Berkeley County, South Carolina, USA. July, 2015.
Copyright©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); habit. Old Santee Canal Park, Berkeley County, South Carolina, USA. July, 2015.
HabitCyrtomium falcatum (Japanese holly fern); habit. Old Santee Canal Park, Berkeley County, South Carolina, USA. July, 2015.©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); fronds, upper surface. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
TitleFronds
CaptionCyrtomium falcatum (Japanese holly fern); fronds, upper surface. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
Copyright©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); fronds, upper surface. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
FrondsCyrtomium falcatum (Japanese holly fern); fronds, upper surface. South Carolina State House, Richland County, South Carolina, USA. May, 2015.©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); frond, upper surface. Riverbanks Botanical Garden, Lexington County, South Carolina, USA. May, 2015.
TitleFrond
CaptionCyrtomium falcatum (Japanese holly fern); frond, upper surface. Riverbanks Botanical Garden, Lexington County, South Carolina, USA. May, 2015.
Copyright©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); frond, upper surface. Riverbanks Botanical Garden, Lexington County, South Carolina, USA. May, 2015.
FrondCyrtomium falcatum (Japanese holly fern); frond, upper surface. Riverbanks Botanical Garden, Lexington County, South Carolina, USA. May, 2015.©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); frond, upper surface. Riverbanks Botanical Garden, Lexington County, South Carolina, USA. May, 2015.
TitleFrond
CaptionCyrtomium falcatum (Japanese holly fern); frond, upper surface. Riverbanks Botanical Garden, Lexington County, South Carolina, USA. May, 2015.
Copyright©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); frond, upper surface. Riverbanks Botanical Garden, Lexington County, South Carolina, USA. May, 2015.
FrondCyrtomium falcatum (Japanese holly fern); frond, upper surface. Riverbanks Botanical Garden, Lexington County, South Carolina, USA. May, 2015.©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); habit. Maliko, Maui, Hawaii, USA. May, 2010.
TitleHabit
CaptionCyrtomium falcatum (Japanese holly fern); habit. Maliko, Maui, Hawaii, USA. May, 2010.
Copyright©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); habit. Maliko, Maui, Hawaii, USA. May, 2010.
HabitCyrtomium falcatum (Japanese holly fern); habit. Maliko, Maui, Hawaii, USA. May, 2010.©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
TitleHabit
CaptionCyrtomium falcatum (Japanese holly fern); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
Copyright©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
HabitCyrtomium falcatum (Japanese holly fern); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); habit. Maliko, Maui, Hawaii, USA. May, 2010.
TitleHabit
CaptionCyrtomium falcatum (Japanese holly fern); habit. Maliko, Maui, Hawaii, USA. May, 2010.
Copyright©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); habit. Maliko, Maui, Hawaii, USA. May, 2010.
HabitCyrtomium falcatum (Japanese holly fern); habit. Maliko, Maui, Hawaii, USA. May, 2010.©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); frond, upper surface. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
TitleFrond
CaptionCyrtomium falcatum (Japanese holly fern); frond, upper surface. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
Copyright©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); frond, upper surface. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
FrondCyrtomium falcatum (Japanese holly fern); frond, upper surface. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); fronds, lower surface (note devloping sori) and smooth upper surface. Honokowai Ditch Trai, Maui, Hawaii, USAl. June, 2010.
TitleFronds
CaptionCyrtomium falcatum (Japanese holly fern); fronds, lower surface (note devloping sori) and smooth upper surface. Honokowai Ditch Trai, Maui, Hawaii, USAl. June, 2010.
Copyright©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); fronds, lower surface (note devloping sori) and smooth upper surface. Honokowai Ditch Trai, Maui, Hawaii, USAl. June, 2010.
FrondsCyrtomium falcatum (Japanese holly fern); fronds, lower surface (note devloping sori) and smooth upper surface. Honokowai Ditch Trai, Maui, Hawaii, USAl. June, 2010.©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
TitleInvasive habit
CaptionCyrtomium falcatum (Japanese holly fern); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
Copyright©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.
Invasive habitCyrtomium falcatum (Japanese holly fern); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June, 2010.©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); underside of a frond, showing sori. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
TitleFrond
CaptionCyrtomium falcatum (Japanese holly fern); underside of a frond, showing sori. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
Copyright©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); underside of a frond, showing sori. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
FrondCyrtomium falcatum (Japanese holly fern); underside of a frond, showing sori. South Carolina State House, Richland County, South Carolina, USA. May, 2015.©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); frond, lower surface showing sori. Maliko, Maui, Hawaii, USA. May, 2010
TitleFrond
CaptionCyrtomium falcatum (Japanese holly fern); frond, lower surface showing sori. Maliko, Maui, Hawaii, USA. May, 2010
Copyright©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); frond, lower surface showing sori. Maliko, Maui, Hawaii, USA. May, 2010
FrondCyrtomium falcatum (Japanese holly fern); frond, lower surface showing sori. Maliko, Maui, Hawaii, USA. May, 2010©Forest & Kim Starr-2010 - CC BY 3.0
Cyrtomium falcatum (Japanese holly fern); extreme close-up of frond undersurface, showing sori and sporangia. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
TitleFrond
CaptionCyrtomium falcatum (Japanese holly fern); extreme close-up of frond undersurface, showing sori and sporangia. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
Copyright©Keith A. Bradley-2013
Cyrtomium falcatum (Japanese holly fern); extreme close-up of frond undersurface, showing sori and sporangia. South Carolina State House, Richland County, South Carolina, USA. May, 2015.
FrondCyrtomium falcatum (Japanese holly fern); extreme close-up of frond undersurface, showing sori and sporangia. South Carolina State House, Richland County, South Carolina, USA. May, 2015.©Keith A. Bradley-2013

Identity

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Preferred Scientific Name

  • Cyrtomium falcatum (L. f.) C. Presl

Preferred Common Name

  • Japanese holly fern

Other Scientific Names

  • Aspidium falcatum (L. f.) Sw.
  • Cyrtomium acutidens H. Christ
  • Cyrtomium yiangshanense Ching & Y. C. Lan
  • Dryopteris falcata (L. f.) O. Kuntze
  • Phanerophlebia falcata (L. f.) Copel.
  • Polypodium falcatum L. f.
  • Polypodium japonicum Houttuyn
  • Polystichum falcatum (L. f.) Diels

International Common Names

  • English: Asian net-veined holly fern; house holly fern; Japanese netvein holly fern
  • Chinese: quan yuan guan zhong

Local Common Names

  • Korea, Republic of: Do-kkae-bi-go-bi; Do-kkae-bi-soe-go-bi
  • USA/Hawaii: ka'ape'ape

EPPO code

  • CWUFA (Cyrtomium falcatum)

Summary of Invasiveness

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Cyrtomium falcatum, commonly known as Japanese holly fern, is native to Asia (including China and Japan). It has been introduced across the world as an ornamental plant and has spread throughout tropical and subtropical areas, and in temperate regions with milder climates, and has spread rapidly across the USA. Colonization of new areas is facilitated by its ability to reproduce apogamously. It can be found in a broad range of habitats, typically on rock or masonry substrates, including coastal areas, waterfalls, karst, boulders, streams, as well as man-made structures. Where it occurs in natural habitats it can displace native species, including rare bryophytes and ferns. In the USA it has been recognized by the US National Park Service as invasive in much of its naturalized range.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Pteridophyta
  •             Class: Filicopsida
  •                 Family: Dryopteridaceae
  •                     Genus: Cyrtomium
  •                         Species: Cyrtomium falcatum

Notes on Taxonomy and Nomenclature

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C. falcatum is a fern in the family Dryopteridaceae. The species was originally described as Polypodium falcatum by Linnaeus the Younger, based on a specimen from Japan. In 1836 Presl described the new genus Cyrtomium and transferred the species (Presl, 1836). The circumscription of Cyrtomium and the similar genus Polystichum has been controversial (Christensen, 1930), and the species has sometimes been treated as a member of that genus (Engler and Prantl, 1899). Genetic studies by Little and Barrington (2003), Li et al. (2004), and Lu et al. (2005) have shown that Cyrtomium is nested within a polyphyletic Polystichum. This required that Cyrtomium be included in a more broadly defined Polystichum, or, as recommended by Lu et al. (2005), separation of the Polystichum clade Balansana into a new genus, and retention of Cyrtomium. The genus Cyrtomium is currently recognized to contain about 35 species (Flora of China Editorial Committee, 2015).

Several subspecies have sometimes been recognized based on cytotypes and geographic distribution (Matsumoto, 2003b). This subdivision has not been followed by recent works (Flora of China Editorial Committee, 2015; Flora of North America Editorial Committee, 2015). 

Description

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C. falcatum is a fern with an erect rhizome covered with brown lanceolate scales, 30-60 cm or rarely longer, stipe stramineous, base with brown scales, leaves pinnate, stiff and glossy, pinnae alternate, 4-15 pairs, falcate, medial pinnae 6-10 cm long, to 3 cm wide, margins undulate, entire to coarsely dentate, apex long acuminate to caudate, sori throughout abaxial side of pinnae, indusial brown. 

Plant Type

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Broadleaved
Herbaceous
Perennial
Seed propagated

Distribution

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C. falcatum is native to coastal provinces of China and Japan (Ohwi et al., 1965; Flora of China Editorial Committee, 2015). Reports in older literature from other regions in Asia, such as Hong Kong (Bentham, 1861) and India (Blatter and D’Almeida, 1922) refer to other Cyrtomium species.

C. falcatum has become naturalized in North America, Africa, Europe and Oceania and a few occurrences have been recorded in South America and the Caribbean. Some reports represent short-lived colonies, particularly those in colder climates. These have been difficult to identify because of a lack of published follow-up data.  

In North America, the species has become established primarily in southern states but other populations have been reported in colder climates including Oregon, Virginia, Ohio, New York, New Jersey and Connecticut (Flora of North America Editorial Committee, 2015; USDA-NRCS, 2015; Weakley, 2015). In Texas, it has been reported as showing little or no increase of abundance or range (Nesom, 2009).

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 Feb 2022
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

MadagascarPresentIntroduced2005Invasive
RéunionPresentIntroduced
South AfricaPresentIntroducedInvasiveFirst reported: 1909-1911

Asia

ChinaPresentNative
-FujianPresentNative
-GuangdongPresentNative
-HubeiPresentNative
-JiangsuPresentNative
-LiaoningPresentNative
-ShandongPresentNative
-ZhejiangPresentNative
Hong KongAbsent, Invalid presence record(s)
IndiaAbsent, Invalid presence record(s)
JapanPresentNative
-HonshuPresentNative
-KyushuPresentNative
-Ryukyu IslandsPresentNative
-ShikokuPresentNative
North KoreaPresentNative
South KoreaPresentNativeDokdo
TaiwanPresentNative
VietnamPresentNative

Europe

AustriaPresentIntroduced1951As: Polystichum falcatum
BelgiumPresentIntroduced2002
CroatiaPresentIntroduced1994
CzechiaPresent
IrelandPresentIntroduced1983West Cork
ItalyPresentIntroduced1987Emilia-Romagna
-SicilyPresent
NetherlandsPresentIntroduced1915
PortugalPresentIntroduced1987
-AzoresPresentIntroduced1932
-MadeiraPresentIntroduced1986
RomaniaPresentIntroduced1969
SpainPresentIntroduced1986
-Canary IslandsPresent1985Salvage Islands, Canaries, Madeira
SwitzerlandPresentIntroduced2014Canton Ticino
United KingdomPresentIntroduced1976

North America

BermudaPresentIntroducedInvasive
United StatesPresentPresent based on regional distribution.
-AlabamaPresentIntroduced1915
-ArkansasPresentIntroduced2002
-CaliforniaPresentIntroduced1940
-ConnecticutPresent
-FloridaPresentIntroduced1923St. Augustine, Duval
-GeorgiaPresentIntroduced
-HawaiiPresentIntroduced1928Maui, Oahu, Molokai, Kauai, Hawaii
-LouisianaPresentIntroduced
-MississippiPresentIntroduced
-New JerseyPresentIntroduced1907
-New YorkPresentIntroduced
-North CarolinaPresentIntroduced
-OhioPresentIntroduced1956
-OregonPresentIntroduced
-South CarolinaPresentIntroduced
-TexasPresentIntroduced1998
-VirginiaPresentIntroduced

Oceania

AustraliaPresentIntroduced1959
-Lord Howe IslandPresentIntroduced1981
-New South WalesPresentIntroduced1981
-QueenslandPresentIntroducedInvasive
-South AustraliaPresentIntroducedAdelaide
-TasmaniaPresentIntroduced
-VictoriaPresentIntroduced
-Western AustraliaPresentIntroduced
French PolynesiaPresentNative
New ZealandPresentIntroduced1988
Norfolk IslandPresentIntroduced1989On walls in garden

South America

ArgentinaPresentIntroduced1996

History of Introduction and Spread

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The earliest report of naturalization was from New Jersey, USA, in 1907, when C. falcatum was found to be establishing in a well near a greenhouse where it was being cultivated (Benedict, 1907). It was then found in Alabama around 1915 (Graves, 1919), and in Florida in 1923 (Christensen, 1930). It has since spread and is found in at least 17 US states, including Hawaii.

It was first reported in South Africa between 1909 and 1911, but it may not have become permanently established there until recent years (Roux, 2011). It was recorded in the Netherlands (Europe) in 1915 (Denters, 2003), New South Wales (Australia) in 1981 (Pickard, 1984), and Argentina (South America) in 1996 (Missouri Botanical Garden, 2015). The species continues to be recorded in new countries, such as Switzerland in 2014 (Schoenenberger et al., 2014).      

Risk of Introduction

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New introductions will continue because C. falcatum is a popular ornamental plant.

Climate change will influence the spread of this species into cooler climates. Denters (2003) reports range expansion due to warmer winters in Belgium and the Netherlands. 

Habitat

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Despite its thick leaves suggesting a preference for xeric habitats, C. falcatum favours mesophytic or coastal habitats (Meyer, 1927; Katsuyama et al., 2011; Flora of China Editorial Committee, 2015). In China, the species grows in lowland forests and coastal areas, to 500 m (Flora of China Editorial Committee, 2015). In Taiwan, it similarly grows in rocky coastal areas (Turner et al., 2001). In Japan, it is primarily a coastal species, growing in rocky areas (Ohwi et al., 1965; Katsuyama et al., 2011) but is occasionally found inland (Ohwi et al., 1965).

In its introduced range, C. falcatum can be found in a variety of habitats, usually associated  with rocks or masonry, both in coastal and inland areas. Throughout the introduced range it occurs on man-made structures including walls, well interiors, stairways and other masonry or brick structures where there is shade and protection from excessive cold (Diddell, 1941; Degener and Hawkes, 1951; Esler, 1988; Mele et al., 2006; Roux, 2011).  

In the eastern USA it is associated with stream banks, limestone outcrops, bluffs, drainage ditches, wooded ravines and waterfalls (Faircloth, 1975; Hill, 1992; Woods and Diamond, 2008; Peck, 2011; Weakley, 2015). Unlike its native range and some parts of its naturalized range, it has not been reported for coastal habitats in this region. In California, USA, it has been found associated with a waterfall in a canyon (Tracy, 1940). In Hawaii it has been found in “bare, windswept precipice and in the wooded gullies” (Degener and Hawkes, 1951).

In Italy, C. falcatum occupies streams, cliffs, walls, wells and brackish areas (Marchetti, 2014) and humid cliffs (Mele et al., 2006). In the Azores, it colonizes coastal areas, rock crevices, walls, slopes, and shady ravines (Schafer, 2001). In South Africa, it is known from watercourses, moist cliffs, rocky outcrops, stone walls and sometimes inland forests (Roux, 2011).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial ManagedDisturbed areas Present, no further details Harmful (pest or invasive)
Terrestrial ManagedDisturbed areas Present, no further details Natural
Terrestrial ManagedUrban / peri-urban areas Present, no further details Harmful (pest or invasive)
Terrestrial ManagedUrban / peri-urban areas Present, no further details Natural
Terrestrial ManagedBuildings Present, no further details Harmful (pest or invasive)
Terrestrial ManagedBuildings Present, no further details Natural
Terrestrial Natural / Semi-naturalNatural forests Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural forests Present, no further details Natural
Terrestrial Natural / Semi-naturalRiverbanks Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalRiverbanks Present, no further details Natural
Terrestrial Natural / Semi-naturalRocky areas / lava flows Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalRocky areas / lava flows Present, no further details Natural
LittoralCoastal areas Present, no further details Harmful (pest or invasive)
LittoralCoastal areas Present, no further details Natural

Biology and Ecology

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Genetics

C. falcatum has a chromosome number of 2n=82 (Lu et al., 2006), although naturalizing populations are reported to be apogamous triploids with a chromosome number of 2n=123 (Flora of North America Editorial Committee, 2015; Weakley, 2015). Three cytotypes have been identified in the species, including sexual diploid, apogamous triploid and sexual tetraploid (Matsumoto, 2003a; Lu et al., 2006; Flora of China Editorial Committee, 2015).

Reproductive Biology

C. falcatum is a homosporous fern (Chung and Chung, 2013). Populations that have naturalized are apogamous triploids. Being apogamous means that these plants can avoid the sexual process in the gametophyte stage so sporophytes arise more efficiently, allowing for easier colonization of new sites (Robinson, 2009).

Physiology and Phenology

C. falcatum produces spores from spring to autumn (Diggs and George, 2006).

It is moderately tolerant of arsenic (Sridokchan et al., 2005).       

Associations

In Korea, the roots of C. falcatum were reported to be colonized by a mycorrhizal fungus with constricted hyphae, similar to those of Rhizoctonia spp. that colonize orchid roots (Lee et al., 2001).

Environmental Requirements

C. falcatum generally requires moist or shaded conditions for establishment, such as stream banks, shaded walls, waterfalls and ravines, and is sensitive to colder temperatures. In studies of phenology and wintering capacity, Toshiyuki (1982) found gametophytes resistant to temperatures of -30°C, and sporophytes to -15°C. Gametophytes survived well in a region where temperatures of -30°C can occur. They also found that gametophytes could survive independently of sporophytes under some conditions. 

Climate

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ClimateStatusDescriptionRemark
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
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
Df - Continental climate, wet all year Preferred Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)
Dw - Continental climate with dry winter Preferred Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) -15

Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • alkaline
  • neutral

Soil texture

  • light
  • medium

Special soil tolerances

  • infertile

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aphelenchoides Herbivore not specific
Coccus hesperidum Herbivore not specific
Colletotrichum acutatum Pathogen not specific
Heliothrips haemorrhoidalis Herbivore not specific
Idiopterus nephrelepidis Herbivore not specific
Macromyzus woodwardiae Herbivore not specific
Pseudococcus longispinus Herbivore not specific
Pseudomonas Pathogen not specific
Pseudomonas asplenii Pathogen
Pythium Pathogen not specific
Rhizoctonia Pathogen not specific

Notes on Natural Enemies

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Little published data exists on natural enemies of C. falcatum. In its native range in Korea, an aphid, Macromyzus woodwardiae (Sternorrhyncha, Aphididae), has been documented from the undersides of new leaves (Lee, 2002).          

Several pests have been identified in its introduced range, primarily associated with threats to cultivated plants. Greenhouse thrips (Heliothrips haemorrhoidalis) have been found on cultivated C. falcatum in Bermuda (Nakahara and Hilburn, 1989). Fern anthracnose (Colletotrichum acutatum) has severely damaged C. falcatum in central Florida, USA, and has caused economic losses (Strandberg et al., 1997).

Means of Movement and Dispersal

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Natural Dispersal

C. falcatum, like all ferns, produces spores which are small and dispersed by wind (Chung and Chung, 2013).

Accidental Introduction

C. falcatum is grown outdoors and indoors as an ornamental plant. Where grown outside, windborne spores may be dispersed from mature plants.

Intentional Introduction

C. falcatum is valued as an ornamental fern and may be introduced to new areas for this purpose.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Cut flower trade Yes Yes
Escape from confinement or garden escape Yes Yes
Garden waste disposal Yes
Horticulture Yes Yes
Landscape improvement Yes Yes
Nursery trade Yes Yes
Ornamental purposes Yes Yes

Impact Summary

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CategoryImpact
Economic/livelihood Positive
Environment (generally) Negative

Environmental Impact

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Impact on Biodiversity

C. falcatum can colonize sparsely vegetated rocks, often in moist and humid conditions. It therefore has the potential to displace co-occurring rare ferns and bryophytes that often grow in such habitats at low population sizes. In Hawaii, USA, it has been reported to displace the endangered Stenogyne bifida, a herbaceous vine in the mint family (Lamiaceae) (US Fish and Wildlife Service, 2010). In Bermuda, it is reported to compete with the endangered Bermuda shield fern (Thelypteris bermudiana) (Copeland and Malcolm, 2014) and the endangered epiphyte, wild Bermuda pepper (Peperomia septentrionalis) (Bárrios et al., 2015).

In Macaronesia, C. falcatum is displacing the fern, sea spleenwort (Asplenium marinum) (Robinson, 2009). It has the potential in Florida to displace rare, native fern species such as Trichomanes punctatum subsp. floridanum, but it has not been observed to displace rare native ferns in the southeastern USA (Alan Cressler, Atlanta, GA, USA, personal communication, 2015; Jennifer Possley, Fairchild Tropical Botanic Garden, Florida, USA, personal communication, 2015).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Peperomia septentrionalis (wild Bermuda pepper)EN (IUCN red list: Endangered)BermudaBarrios et al. (2015)
Stenogyne bifida (twocleft stenogyne)CR (IUCN red list: Critically endangered); USA ESA listing as endangered speciesHawaiiCompetition - shading; Competition - smotheringUS Fish and Wildlife Service (2010)
Thelypteris bermudiana (Bermuda shield fern)EN (IUCN red list: Endangered)BermudaCompetition - smotheringCopeland and Malcolm (2014)

Risk and Impact Factors

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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
  • Pioneering in disturbed areas
  • Tolerant of shade
  • Benefits from human association (i.e. it is a human commensal)
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Reproduces asexually
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Infrastructure damage
  • Modification of successional patterns
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Causes allergic responses
  • Competition - monopolizing resources
  • Competition - shading
  • Competition - smothering
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately
  • Difficult to identify/detect as a commodity contaminant

Uses

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Economic Value

C. falcatum is cultivated globally as an ornamental plant (Degener and Hawkes, 1951). The fronds are also used in cut flower arrangements (Will and Burch, 1985). 

Uses List

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General

  • Botanical garden/zoo

Ornamental

  • Cut flower
  • Potted plant

Similarities to Other Species/Conditions

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C. falcatum can be confused with some larger forms of Cyrtomium fortunei in herbaria, although they are not likely to be confused when alive because of the glossy pinna surface of C. falcatum (Christensen, 1930). 

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.

Control

New colonies are often the result of spread from cultivated plants so efforts should be made to eliminate plants from adjacent areas and may require the cooperation of landowners nearby.

Physical/Mechanical Control

Control of C. falcatum is primarily accomplished by hand pulling. Pulled plants should be removed from the site to prevent the spread of spores.

Chemical Control

Where there are denser populations of C. falcatum, or where plants occur on sensitive substrates such as historical masonry structures, control can be achieved with foliar glyphosate application.

References

Top of page

Aplaca JL, 2010. The Non-Native Flora of Texas. Master of Science Thesis. San Marcos, Texas, USA: Texas State University.

AVH, 2015. Australia's Virtual Herbarium. Canberra, ACT, Australia: Council of Heads of Australasian Herbaria. http://avh.chah.org.au/

Barker RM, Jessop JP, Vonow HP, 2005. Census of South Australian vascular plants, supplement 1, fifth edition. Journal of the Adelaide Botanic Gardens.

Barrios S, Copeland A, Malcolm P, 2015. Peperomia septentrionalis. The IUCN Red List of Threatened Species. Version 2015.2. http://www.iucnredlist.org/details/68982125/0

Benedict RC, 1907. Notes on some ferns collected near Orange, New Jersey. Torreya, 7(7):136-138.

Bentham G, 1861. Flora Hongkongensis - a description of the flowering plants and ferns of the island of Hong Kong. London, UK: L Reeve.

Bibiloni J, 2011. Future is written in green - The beautiful pest that came from China. http://mundani-garden.blogspot.com/2011/04/beautiful-pest-that-came-from-china.html

Biological Records Centre, 2015. Online atlas of the British and Irish flora. Wallingford, UK: Biological Records Centre. http://www.brc.ac.uk/plantatlas/

Bishop Museum, 2015. Online database. Natural sciences collections. Honolulu, Hawaii, USA: Bishop Museum. http://nsdb.bishopmuseum.org/

Blatter E, D'Almeida JF, 1922. The Ferns of Bombay. Bombay, India: DB Taraporevala Sons And Co.

Bonafede F, Ferrari C, Vigarani A, 1993. Cyrtomium falcatum, new to the Italian flora. Flora Meditteranea, 3:261-264.

Casasayas T, Farras A, 1986. Short notes: Polystichum falcatum (L. fil.) Diels, adventitious to Catalonia. (Notes breus: Polystichum falcatum (L fil) Diels, adventicia a Catalunya). Collectanea Botanica, 16(2):425-426.

Chang C-S, Kim H, Chang K, 2014. Provisional checklist of vascular plants for the Korea peninsula flora (KPF). Seoul, Korea: Seoul National University.

Christ H, 1911. Filices Wilsonianae. Botanical Gazette, 51(5):345-359.

Christenhusz MJM, Uffelen GAvan, 2001. Naturalized Japanese plants in the Netherlands, introduced by Von Siebold. Gorteria, 27(5):97-108.

Christensen C, 1930. The genus Cyrtomium. American Fern Journal, 20(2):41-52.

Chung MY, Chung MG, 2013. Significant spatial aggregation and fine-scale genetic structure in the homosporous fern Cyrtomium falcatum (Dryopteridaceae). New Phytologist, 199(3):663-672.

Copeland A, Malcolm P, 2014. Thelypteris bermudiana. The IUCN Red List of Threatened Species. Version 2015.2. http://www.iucnredlist.org/details/56604509/0

Corkhill P, 1977. Cyrtomium falcatum naturalised on Rhum. Fern Gazette, 11(5):277.

Cullinane JP, Crowley C, 1985. Cyrtomium falcatum in Ireland. Fern Gazette, 13(1):42.

Degener O, Hawkes AD, 1951. The holly fern. American Fern Journal, 41(4):117-120.

Denters T, 2003. History and present occurrence of Cyrtomium falcatum (Lf) CB Presl in the Netherlands. Gorteria, 29(5):125-133.

Diamond Jr AR, Woods M, 2007. Pteridophytes of southeast Alabama. Journal of the Alabama Academy of Science, 78(1):21-28.

Diddell MW, 1941. New Stations for Florida pteridophytes. American Fern Journal, 31(2):48-52.

Diggs GM, George R, 2006. Illustrated flora of east Texas, volume 1. Fort Worth, Texas, USA: Botanical Research Institute of Texas.

Dobrescu CM, Soare LC, 2012. Researches on pteridophytes from Valsan Valley Protected Area (Arges County, Romania). Analele Stiintifice ale Universitatii, Biologie Vegetala, 58:89-94.

Engler A, Prantl K, 1899. The natural plant families (Die naturlichen Pflanzenfamilien). Leipzig: Wilhelm Engelmann.

Esler AE, 1988. The naturalisation of plants in urban Auckland, New Zealand. 5. Success of the alien species. New Zealand Journal of Botany, 26(4):565-584.

Faircloth WR, 1975. Ferns and other primitive vascular plants of central south Georgia. Castanea, 40(3):217-228.

Flora of China Editorial Committee, 2015. 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 North America Editorial Committee, 1993. Flora of North America: north of Mexico, volume 2. Oxford, UK: Oxford University Press.

Flora of North America Editorial Committee, 2015. Flora of North America North of Mexico. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=1

Gardner R, 2006. Some Norfolk Island plant records. Auckland Botanical Society, 61:108-112.

Goslin CR, 1958. The holly-fern, Cyrtomium falcatum, outdoors in Ohio. American Fern Journal, 48(2):84-85.

Graves EW, 1919. A new station for Cyrtomium falcatum and Pteris longifolia in Alabama. American Fern Journal, 9(4):119-120.

Hazelton E, Frey M, DuPrey A, 2012. Invasive plant alert - Japanese holly fern (Cyrtomium falcatum). Washington, DC, USA: National Park Service, National Capital Region Exotic Plant Management Team.

Heiss-Dunlop S, Fillery J, 2006. Vascular flora of Motuora Island, Hauraki Gulf. Auckland Botanical Society, 61. 113-120.

Hibernicarum S, 2008. The way that we went. Pteridologist, 5(1):42.

Hill SR, 1992. Calciphiles and calcareous habitats of South Carolina. Castanea, 57(1):25-33.

Hill SR, Horn CN, 1997. Additions to the flora of South Carolina. Castanea, 62(3):194-208.

Howell C, Sawyer JWD, 2006. New Zealand naturalised vascular plant checklist. Wellington, New Zealand: New Zealand Plant Conservation Network, 60 pp.

Hutchinson G, Thomas BA, 1992. Distribution of Pteridophyta in Wales. Watsonia, 19(1):1-19.

Jermy AC, British PP, 1978. Atlas of ferns of the British Isles. London, UK: Botanical Society of the British Isles, British Pteridological Society.

Jung S, Byun J, Park S, Oh S, Yang J, Jang J, Chang K, Lee Y, 2014. The study of distribution characteristics of vascular and naturalized plants in Dokdo, South Korea. Journal of Asia-Pacific Biodiversity, 7(2):e197-e205.

Katsuyama T, Hasekura C, Kokubo K, 2011. An annotated checklist of the vascular plants of Aogashima Island in the Izu Islands, Japan. Bulletin Kanagawa Prefectural Museum (Natural Sciences), 40:7-34.

Lee J-K, Eom A-H, Lee S-S, Lee CH, 2001. Mycorrhizal symbioses found in roots of fern and its relatives in Korea. Journal of Plant Biology, 44(2):81-86.

Lee S, 2002. New record of Macromyzus woodwardiae (Sternorhyncha, Aphididae) on Cyrtomium falcatum from Korea. Korean Journal of Systematic Zoology, 18(1):127-133.

Li C, Shugang L, Qun Y, 2004. Asian origin for Polystichum (Dryopteridaceae) based on rbcL sequences. Chinese Science Bulletin, 49(11):1-5.

Li J, Chiou W, 2006. Changes in the flora on islet Pengchiayu across 100 years. Taiwania, 51(3):195-209.

Little DP, Barrington DS, 2003. Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae). American Journal of Botany, 90(3):508-514.

Lohr MT, Keighery G, 2014. The status and distribution of alien plants on the islands of the south coast of Western Australia. Conservation Science Western Australia, 9(2):181-200.

Lu JM, Cheng X, Wu D, Li DZ, 2006. Chromosome study of the fern genus Cyrtomium (Dryopteridaceae). Botanical Journal of the Linnean Society, 150(2):221-228.

Lu JM, Li DZ, Gao LM, Cheng X, Wu D, 2005. Paraphyly of Cyrtomium (Dryopteridaceae) - evidence from rbcL and trnL-F sequence data. Journal of Plant Research, 118(2):129-135.

Lu S, Yang TYA, 2005. The checklist of Taiwanese pteridophytes following Ching's system. Taiwania, 50(2):137-165.

Lubin D, 2015. Ferns of Bermuda. Street Allston, Massachussetts, USA. http://nefern.info/SiteList/bermudalist.htm

Marchetti D, 2004. Pteridophytes of Italy. (Le Pteridofite D'Italia). Annals of the Museum of Civilisation Rovereto, 19:71-231.

Matsumoto S, 2003. Cytotaxonomic study of Cyrtomium falcatum complex (Dryopteridaceae) in Japanese Archipelago (Symposium open to the public - Chromosome studies in connection with phylogeny and taxonomy). Chromosome Science, 7(4):108.

Matsumoto S, 2003. Species ecological study on reproductive systems and speciation of Cyrtomium falcatum complex (Dryopteridaceae) in Japanese archipelago. Annals of Tsukuba Botanical Garden, 22:1-141.

Mele C, Medagli P, Accogli R, Beccarisi L, Albano A, Marchiori S, 2006. Flora of Salento (Apulia, southeastern Italy) - an annotated checklist. Flora Mediterranea, 16:193-245.

Meyer BS, 1927. Studies on the physical properties of leaves and leaf saps. Ohio Journal of Science, 27(6):263-288.

Missouri Botanical Garden, 2015. Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

Mountier CF, 2014. Rarity and commonness in New Zealand ferns. Bachelor of Science Dissertation. Christchurch, New Zealand: Lincoln University.

Nakahara S, Hilburn DJ, 1989. Annotated checklist of the Thysanoptera of Bermuda. Journal of the New York Entomological Society, 97(3):251-260

Negrean G, 2011. Addenda to flora Romaniae, volumes 1-12 - Newly published plants, nomenclature, taxonomy, chorology and commentaries (Part 1). Kanitzia, 8:89-194.

Nesom GL, 2009. Assessment of invasiveness and ecological impact in non-native plants of Texas. Journal of the Botanical Research Institute of Texas, 3(2):971-991.

Ohwi J, Meyer FG, Walker EH, 1965. Flora of Japan (in English). Smithsonian Institution.

Peck JH, 2003. Arkansas flora - additions, reinstatements, exclusions, and re-exclusions. Sida, 20(4):1737-1757.

Peck JH, 2011. New and noteworthy additions to the Arkansas fern flora. Phytoneuron, 30:1-33.

Pickard J, 1984. Exotic plants on Lord Howe Island: distribution in space and time, 1853-1981. Journal of Biogeography, 11:181-208.

PlantNET, 2015. New South Wales flora online. Sydney, New South Wales, Australia: National Herbarium of New South Wales. http://plantnet.rbgsyd.nsw.gov.au/floraonline.htm

Popenoe J, 1989. Key plants/key pests. Commercial horticulture - woody ornamentals. UF/IFAS Lake County Extension Service. http://mrec.ifas.ufl.edu/lso/SCOUT/key.pdf

Presl C, 1836. Tentamen Pteridographiae - or filicacearum types of mines, especially in the course of distribution and exposure (Tentamen Pteridographiae - seu genera filicacearum praesertim juxta venarum decursum et distributionem exposita). Prague, Czech Republic: Typis filiorum Theophili Haase.

Press JR, Biscoito M, Zino FJ, 1986. New plant records from the Salvage Islands. Bocagiana, 90:1-4.

Robinson RC, 2009. Invasive and problem ferns - a European perspective. International Urban Ecology Review, 4:83-91.

Roux JP, 2011. The genus Cyrtomium (Pteridophyta, Dryopteridaceae) in Africa and Madagascar. Botanical Journal of the Linnean Society, 167(4):449-465.

Schafer H, 2001. Distribution and status of the pteridophytes of Faial island, Azores (Portugal). Fern Gazette, 16(5):213-237.

Schoenenberger N, Rothlisberger J, Carraro G, 2014. The exotic flora of the Canton Ticino (Switzerland). (La flora esotica del Cantone Ticino (Svizzera). Bolletino della Societa ticinese di scienze naturali, 102:13-30.

Segarra-Moragues JG, 2001. Data on pteridoflora subspontaneous Iberica - Cyrtomium falcatum (Dryopteridaceae) and Nephrolepis cordifolia (Nephrolepidaceae). (Datos sobre la pteridoflora subespontanea Iberica - Cyrtomium falcatum (Dryopteridaceae) y Nephrolepis cordifolia (Nephrolepidaceae)). Acta Botanica Malacitana, 26:247-249.

SIBIS, 2015. South African Biodiversity Information Facility. South Africa: South African National Biodiversity Institute. http://sibis.sanbi.org/faces/Home.jsp?1=1

Sridokchan W, Markich S, Visoottiviseth P, 2005. Arsenic tolerance, accumulation and elemental distribution in twelve ferns - a screening study. Australasian Journal of Ecotoxicology, 11(2):101-110.

Strandberg JO, Stamps RH, Norman DJ, 1997. Fern anthracnose - A guide for effective disease management. Florida Agriculture Experiment Station Bulletin (Technical), 900.

The Plant List, 2015. The Plant List: a working list of all plant species. Version 1.1. London, UK: Royal Botanic Gardens, Kew. http://www.theplantlist.org

Toshiyuki S, 1982. Phenology and Wintering Capacity of Sporophytes and Gametophytes of Ferns Native to Northern Japan. Oecologia, 55(1):53-61.

Tracy HH, 1940. Cyrtomium in southern California. American Fern Journal, 30(2):52-56.

Trinajstic I, Panjol Z, 1994. Cyrtomium falcatum (L. fil.) C Presl. (Polystichaceae) a newcomer in the Croatian flora. Natura Croatica, 3(1):87-90.

Turner IM, Xing FW, Corlett RT, 2001. An annotated check-list of the vascular plants of the South China Sea and its shores. Raffles Bulletin of Zoology, 48:23-116.

US Fish and Wildlife Service, 2010. In: 5-Year Review, Short Form Summary: Species Reviewed: Stenogyne bifida (no common name). US Fish and Wildlife Service, 8 pp.

USDA-ARS, 2015. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

USDA-NRCS, 2015. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Verloove F, Lambinon J, 2014. The sixth edition of the Nouvelle Flore de la Belgique - nomenclatural and taxonomic remarks. Dumortiera, 104:7-40.

Weakley AS, 2015. Flora of the southern and mid-Atlantic States - Working draft of 2015. Chapel Hill, North Carolina, USA: University of North Carolina Herbarium (NCU).

Will AA Jr, Burch DG, 1985. Foliage for flower arrangements from your Florida garden. Proceedings of the Florida State Horticultural Society, 98:349-350.

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

Woods M, Diamond AR Jr, 2008. Pteridophytes of southeast Alabama - dichotomous keys, illustrations and distribution maps. Journal of the Alabama Academy of Science, 79:200-237.

Distribution References

Aplaca JL, 2010. The non-native flora of Texas. In: Master of Science thesis, San Marcos, Texas, USA: Texas State University.

AVH, 2015. Australia's Virtual Herbarium., Canberra, ACT, Australia: Council of Heads of Australasian Herbaria. http://avh.chah.org.au/

Barker B, Barker R, Jessop J, Vonow H, 2005. Census of South Australian vascular plants. Journal of the Adelaide Botanic Gardens. 396 pp.

Barrios S, Copeland A, Malcolm P, 2015. Peperomia septentrionalis. In: The IUCN Red List of Threatened Species. Version 2015, 2 http://www.iucnredlist.org/details/68982125/0

Benedict RC, 1907. Notes on some ferns collected near Orange, New Jersey. In: Torreya, 7 (7) 136-138.

Bentham G, 1861. Flora Hongkongensis : a description of the flowering plants and ferns of the island of Hongkong., London, UK: L. Reeve. 596 pp.

Bishop Museum, 2015. Online database. Natural sciences collections., Honolulu, Hawaii, USA: Bishop Museum. http://nsdb.bishopmuseum.org/

Blatter E, D'Almeida JF, 1922. The Ferns of Bombay., Bombay, India: DB Taraporevala Sons And Co.

Bonafede F, Ferrari C, Vigarani A, 1993. Cyrtomium falcatum, new to the Italian flora. In: Flora Meditteranea, 3 261-264.

CABI, Undated. Compendium record. Wallingford, UK: CABI

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

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

Casasayas T, Farras A, 1986. Short notes: Polystichum falcatum (L. fil.) Diels, adventitious to Catalonia. (Notes breus: Polystichum falcatum (L fil) Diels, adventicia a Catalunya). In: Collectanea Botanica, 16 (2) 425-426.

Chang CS, Kim H, Chang K, 2014. Provisional checklist of vascular plants for the Korea peninsula flora (KPF)., Seoul, Korea: Seoul National University.

Christ H, 1911. Filices Wilsonianae. In: Botanical Gazette, 51 (5) 345-359.

Christenhusz M J M, Uffelen G A van, 2001. Naturalized Japanese plants in the Netherlands, introduced by Von Siebold. (Verwilderde Japanse planten in Nederland, ingevoerd door Von Siebold.). Gorteria. 27 (5), 97-108.

Christensen C, 1930. The genus Cyrtomium. In: American Fern Journal, 20 (2) 41-52.

Copeland A, Malcolm P, 2014. Thelypteris bermudiana. In: The IUCN Red List of Threatened Species. Version 2015, http://www.iucnredlist.org/details/56604509/0

Corkhill P, 1977. Cyrtomium falcatum naturalised on Rhum. In: Fern Gazette, 11 (5) 277.

Cullinane JP, Crowley C, 1985. Cyrtomium falcatum in Ireland. In: Fern Gazette, 13 (1) 42.

Degener O, Hawkes AD, 1951. The holly fern. In: American Fern Journal, 41 (4) 117-120.

Denters T, 2003. History and present occurrence of Cyrtomium falcatum (L.f.) C.B. Presl. in the Netherlands. (Verleden en heden van IJzervaren (Cyrtomium falcatum (L. f.) C. B. Presl.) in Nederland.). Gorteria. 29 (5), 125-133.

Diamond Jr AR, Woods M, 2007. Pteridophytes of southeast Alabama. In: Journal of the Alabama Academy of Science, 78 (1) 21-28.

Diddell MW, 1941. New Stations for Florida pteridophytes. In: American Fern Journal, 31 (2) 48-52.

Diggs GM, George R, 2006. Illustrated flora of east Texas., 1 Fort Worth, Texas, USA: Botanical Research Institute of Texas.

Dobrescu CM, Soare LC, 2012. Researches on pteridophytes from Valsan Valley Protected Area (Arges County, Romania). In: Analele Stiintifice ale Universitatii, Biologie Vegetala, 58 89-94.

Esler A E, 1988. The naturalization of plants in urban Auckland, New Zealand. 5. Success of the alien species. New Zealand Journal of Botany. 565-584.

Flora of China Editorial Committee, 2015. 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

Gardner R, 2006. Some Norfolk Island plant records. In: Auckland Botanical Society, 61 108-112.

Goslin CR, 1958. The holly-fern, Cyrtomium falcatum, outdoors in Ohio. In: American Fern Journal, 48 (2) 84-85.

Grasso S, Pane A, Cacciola S O, 2000. First report of Armillaria mellea on a fern from Italy. Plant Disease. 84 (5), 592. DOI:10.1094/PDIS.2000.84.5.592C

Graves EW, 1919. A new station for Cyrtomium falcatum and Pteris longifolia in Alabama. In: American Fern Journal, 9 (4) 119-120.

Heiss-Dunlop S, Fillery J, 2006. Vascular flora of Motuora Island, Hauraki Gulf. In: Auckland Botanical Society, 61 113-120.

Hibernicarum S, 2008. The way that we went. In: Pteridologist, 5 (1) 42.

Hill S R, Horn C N, 1997. Additions to the flora of South Carolina. Castanea. 62 (3), 194-208.

Howell C, Sawyer JWD, 2006. New Zealand naturalised vascular plant checklist., Wellington, New Zealand: New Zealand Plant Conservation Network. 60 pp.

Hutchinson G, Thomas BA, 1992. Distribution of Pteridophyta in Wales. In: Watsonia, 19 (1) 1-19.

Jermy AC, British PP, 1978. Atlas of ferns of the British Isles., London, UK: Botanical Society of the British Isles, British Pteridological Society.

Jung S, Byun J, Park S, Oh S, Yang J, Jang J, Chang K, Lee Y, 2014. The study of distribution characteristics of vascular and naturalized plants in Dokdo, South Korea. In: Journal of Asia-Pacific Biodiversity, 7 (2) e197-e205.

Li J, Chiou W, 2006. Changes in the flora on islet Pengchiayu across 100 years. In: Taiwania, 51 (3) 195-209.

Lohr M T, Keighery G, 2014. The status and distribution of alien plants on the islands of the south coast of Western Australia. Conservation Science Western Australia. 9 (2), 181-200. http://www.dpaw.wa.gov.au/images/documents/conservation-management/forests/forest-produce/LOHR__KEIGHERY_FINAL_LOW.pdf

Lubin D, 2015. Ferns of Bermuda., Massachussetts, USA: Street Allston. http://nefern.info/SiteList/bermudalist.htm

Marchetti D, 2004. Pteridophytes of Italy. (Le Pteridofite D'Italia). In: Annals of the Museum of Civilisation Rovereto, 19 71-231.

Mertelik J, Mokra V, Gotzova B, Gabrielova S, 2002. Occurrence and identification of Impatiens necrotic spot tospovirus in the Czech Republic. Acta Horticulturae. 79-83.

Missouri Botanical Garden, 2015. Tropicos database., St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

Mountier CF, 2014. Rarity and commonness in New Zealand ferns. In: Bachelor of Science Dissertation, Christchurch, New Zealand: Lincoln University.

Negrean G, 2011. Addenda to flora Romaniae, volumes 1-12 - Newly published plants, nomenclature, taxonomy, chorology and commentaries (Part 1). In: Kanitzia, 8 89-194.

Ohwi J, 1965. Flora of Japan. [ed. by Meyer F G, Walker E H]. Washington DC, USA: Smithsonian Institution. ix + 1067 pp. http://biodiversitylibrary.org/page/30045955 \ https://archive.org/details/floraofjapaninen00oiji

Peck JH, 2003. Arkansas flora - additions, reinstatements, exclusions, and re-exclusions. In: Sida, 20 (4) 1737-1757.

Peck JH, 2011. New and noteworthy additions to the Arkansas fern flora. In: Phytoneuron, 30 1-33.

Pickard J, 1984. Exotic plants on Lord Howe Island: distribution in space and time, 1853-1981. Journal of Biogeography. 181-208.

PlantNET, 2015. New South Wales flora online., Sydney, New South Wales, Australia: National Herbarium of New South Wales. http://plantnet.rbgsyd.nsw.gov.au/floraonline.htm

Press JR, Biscoito M, Zino FJ, 1986. New plant records from the Salvage Islands. In: Bocagiana, 90 1-4.

Roux JP, 2011. The genus Cyrtomium (Pteridophyta, Dryopteridaceae) in Africa and Madagascar. In: Botanical Journal of the Linnean Society, 167 (4) 449-465.

Schafer H, 2001. Distribution and status of the pteridophytes of Faial island, Azores (Portugal). In: Fern Gazette, 16 (5) 213-237.

Schoenenberger N, Rothlisberger J, Carraro G, 2014. The exotic flora of the Canton Ticino (Switzerland). (La flora esotica del Cantone Ticino (Svizzera)). In: Bolletino della Societa ticinese di scienze naturali, 102 13-30.

Seebens H, Blackburn T M, Dyer E E, Genovesi P, Hulme P E, Jeschke J M, Pagad S, Pyšek P, Winter M, Arianoutsou M, Bacher S, Blasius B, Brundu G, Capinha C, Celesti-Grapow L, Dawson W, Dullinger S, Fuentes N, Jäger H, Kartesz J, Kenis M, Kreft H, Kühn I, Lenzner B, Liebhold A, Mosena A (et al), 2017. No saturation in the accumulation of alien species worldwide. Nature Communications. 8 (2), 14435. http://www.nature.com/articles/ncomms14435

Segarra-Moragues JG, 2001. Data on pteridoflora subspontaneous Iberica - Cyrtomium falcatum (Dryopteridaceae) and Nephrolepis cordifolia (Nephrolepidaceae). (Datos sobre la pteridoflora subespontanea Iberica - Cyrtomium falcatum (Dryopteridaceae) y Nephrolepis cordifolia (Nephrolepidaceae)). In: Acta Botanica Malacitana, 26 247-249.

SIBIS, 2015. South African Biodiversity Information Facility., South Africa: South African National Biodiversity Institute. http://sibis.sanbi.org/faces/Home.jsp?1=1

Tracy HH, 1940. Cyrtomium in southern California. In: American Fern Journal, 30 (2) 52-56.

Trinajstic I, Panjol Z, 1994. Cyrtomium falcatum (L. fil.) C Presl. (Polystichaceae) a newcomer in the Croatian flora. In: Natura Croatica, 3 (1) 87-90.

USDA-NRCS, 2015. The PLANTS Database. Greensboro, North Carolina, USA: National Plant Data Team. https://plants.sc.egov.usda.gov

Verloove F, Lambinon J, 2014. The sixth edition of the Nouvelle Flore de la Belgique - nomenclatural and taxonomic remarks. In: Dumortiera, 104 7-40.

Weakley AS, 2015. Flora of the southern and mid-Atlantic States - Working draft of 2015., Chapel Hill, North Carolina, USA: University of North Carolina Herbarium (NCU).

Wilson K A, 1996. Alien ferns in Hawai'i. Pacific Science. 50 (2), 127-141.

Contributors

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Keith Bradley, Consultant, South Carolina, USA

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