Lygodium japonicum - (Thunb.) Sw.
Japanese Climbing Fern
Other Common Names: Japanese climbing fern
Taxonomic Status: Accepted
Related ITIS Name(s): Lygodium japonicum (Thunb. ex Murr.) Sw. (TSN 17983)
Unique Identifier: ELEMENT_GLOBAL.2.146045
Element Code: PPSCH02010
Informal Taxonomy: Plants, Vascular - Ferns and relatives
 
Kingdom Phylum Class Order Family Genus
Plantae Filicinophyta Filicopsida Filicales Lygodiaceae Lygodium
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Concept Reference
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Concept Reference: Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
Concept Reference Code: B94KAR01HQUS
Name Used in Concept Reference: Lygodium japonicum
Conservation Status
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NatureServe Status

Global Status: GNR
Global Status Last Reviewed: 10Sep2002
Global Status Last Changed: 10Sep2002
Rounded Global Status: GNR - Not Yet Ranked
Reasons: Abundance unknown in its native Japan. (Escaped from cultivation into much of the southeastern U.S. and Texas.)
Nation: United States
National Status: NNA

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (SNA), Arkansas (SNA), Florida (SNA), Georgia (SNA), Hawaii (SNA), Kentucky (SNA), Louisiana (SNA), Mississippi (SNA), North Carolina (SNA), South Carolina (SNA), Texas (SNA)

Other Statuses

NatureServe Global Conservation Status Factors

Other NatureServe Conservation Status Information

Distribution
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U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
NOTE: The distribution shown may be incomplete, particularly for some rapidly spreading exotic species.

U.S. & Canada State/Province Distribution
United States ALexotic, ARexotic, FLexotic, GAexotic, HIexotic, KYexotic, LAexotic, MSexotic, NCexotic, SCexotic, TXexotic

Range Map
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Ecology & Life History
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Habitat Comments: Terrestrial in woods, marshes, and roadside ditches, and on riverbanks in circumneutral soil (Lellinger, 1985).
Economic Attributes Not yet assessed
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Management Summary Not yet assessed
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Population/Occurrence Delineation Not yet assessed
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Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank)
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Disclaimer: While I-Rank information is available over NatureServe Explorer, NatureServe is not actively developing or maintaining these data. Species with I-RANKs do not represent a random sample of species exotic in the United States; available assessments may be biased toward those species with higher-than-average impact.

I-Rank: High/Medium
Rounded I-Rank: High
I-Rank Reasons Summary: A climbing fern established in the southeastern United States (mostly on the coastal plain, NC to e. TX, south to FL and north to s. AR) and in Hawaii (big island and Oahu); invades a variety of habitats including mesic upland forests, mesic to wet flatwoods, bottomland forests, cypress swamps, hydric hammocks, and riparian communities. Most problematic along the Gulf coast and in Florida. Although generally scattered throughout the landscape, it sometimes occurs in dense infestations where it can form mats that smother native understory vegetation. In some of these dense infestations, mats ascend to the upper vegetation layers to smother tall shrubs and trees, which can eventually result in complete dominance of the habitat. High-climbing fronds can function as fire ladders, conducting fire into the canopy where it can kill native dominants. Spreads by rhizomes and spores; spores dispersed long distances by wind and contamination. Spreading south through Florida. Control can be achieved by pulling vines from the trees and treating them with a foliar herbicide; it is generally considered difficult. In general, Lygodium japonicum is not considered to be as serious of a threat to entire forests as its congener L. microphyllum; however, L. microphyllum currently has a much smaller U.S. range, established only in central and southern Florida.
Subrank I - Ecological Impact: High/Medium
Subrank II - Current Distribution/Abundance: Medium
Subrank III - Trend in Distribution/Abundance: High/Medium
Subrank IV - Management Difficulty: Medium
I-Rank Review Date: 26Nov2008
Evaluator: Gravuer, K.
Native anywhere in the U.S?
Native Range: Native to eastern Asia, including eastern and southern China, Japan (incl. western Honshu, Kyushu, Ryukyu Islands, Shikoku), Korea, Bhutan, India, Nepal, Sri Lanka, Cambodia, Laos, Thailand, Vietnam, Indonesia, Malaysia, Papua New Guinea, and the Philippines (USDA-ARS 2008).

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Screening Questions

S-1. Established outside cultivation as a non-native? YES
Comments: This species is a non-native that is established outside of cultivation (Kartesz 1999).

S-2. Present in conservation areas or other native species habitat? Yes
Comments: Can invade natural undisturbed areas (Miller 2003, Weber 2003, ISSG 2006 2006, FL-DEP 2008, Weakley 2008). Substantial numbers of infestations have been recorded within the following natural communities: mesic upland forests (hardwood, pine, and mixed), mesic flatlands (e.g. mesic and scrubby flatwoods), floodplain and bottomland forests/swamps, wet flatlands (e.g. wet flatwoods and hydric hammocks), and communities alongside rivers and streams (FNAI 2008). Apparently also grows on the edges of lakes and marshes (Langeland and Burks 1998, Byrd and Westbrooks 2007, Nation 2008) and in estuarine habitats (ISSG 2006 2006). Known from cypress swamps in Louisiana and beech forests in east Texas (Randall 1996).

Subrank I - Ecological Impact: High/Medium

1. Impact on Ecosystem Processes and System-wide Parameters:Moderate significance
Comments: Can alter local fire ecology; the high-climbing fronds can function as fire ladders (particularly in the winter when they are dead and dry), conducting fire from the ground up into the forest canopy (Randall 1996, Weber 2003, Nation 2008). Native plants in many of the habitats that this species invades are adapted to ground fires, but are killed or severely damaged by crown fires (Randall 1996).

2. Impact on Ecological Community Structure:High/Moderate significance
Comments: Generally scattered throughout the landscape; out of approximately 1800 infestations in the Florida Invasive Plants Geodatabase (FLInv), most were described as scattered plants or clumps; only about 60 were "dominant cover" and only 6 were "dense monoculture" (FNAI 2008). However, Miller (2003) notes that scattered infestations may later increase in cover to form dense mats. At sites with denser infestations, this species' dense mats frequently smother and eliminate native understory vegetation, such as ground cover and low-growing shrubs (Langeland and Burks 1998, Lott et al. 2003, Center for Aquatic and Invasive Plants 2008). Seedlings of overstory tree species can also be smothered, altering successional trajectories (Center for Aquatic and Invasive Plants 2008). Less frequently, but still in a significant number of cases, mats ascend to the upper vegetation layers to smother and shade out taller shrubs and canopy trees, weakening or killing them and eliminating the vegetation below (Randall 1996, Lott et al. 2003, Weber 2003, Center for Aquatic and Invasive Plants 2008, FL-DEP 2008, Nation 2008). Given sufficient time, this species can completely dominate a native habitat and cause the natural community to collapse (ISSG 2006); for example, it is reported to be forming sun-blocking walls of fern in tributary floodplains of the Apalachicola River (Langeland and Burks 1998). L. japonicum is believed to be hardier than many native species in low light environments, allowing it to thrive and expand where natives cannot (ISSG 2006).

3. Impact on Ecological Community Composition:High significance
Comments: Generally scattered throughout the landscape; out of approximately 1800 infestations in the Florida Invasive Plants Geodatabase (FLInv), most were described as scattered plants or clumps; only about 60 were "dominant cover" and only 6 were "dense monoculture" (FNAI 2008). However, Miller (2003) notes that scattered infestations may later increase in cover to form dense mats. At sites with denser infestations, this species' dense mats frequently smother and eliminate native understory vegetation, such as ground cover and low-growing shrubs (Langeland and Burks 1998, Lott et al. 2003, Center for Aquatic and Invasive Plants 2008). Seedlings of overstory tree species can also be smothered, altering successional trajectories (Center for Aquatic and Invasive Plants 2008). Less frequently, but still in a significant number of cases, mats ascend to the upper vegetation layers to smother and shade out taller shrubs and canopy trees, weakening or killing them and eliminating the vegetation below (Randall 1996, Lott et al. 2003, Weber 2003, Center for Aquatic and Invasive Plants 2008, FL-DEP 2008, Nation 2008). Given sufficient time, this species can completely dominate a native habitat and cause the natural community to collapse (ISSG 2006); for example, it is reported to be forming sun-blocking walls of fern in tributary floodplains of the Apalachicola River (Langeland and Burks 1998). L. japonicum is believed to be hardier than many native species in low light environments, allowing it to thrive and expand where natives cannot (ISSG 2006).

4. Impact on Individual Native Plant or Animal Species:Insignificant
Comments: A congeneric native species, Hartford-Fern (Lygodium palmatum Sw.), occurs within the invaded range; however, no reports of hybridization or disproportionate competition impacts were found.

5. Conservation Significance of the Communities and Native Species Threatened:High/Moderate significance
Comments: In Florida, it is believed to threaten the rare plant ray fern (Actinostachys pennula), as well as the endangered Georgia bully (Sideroxylon thornei), common dutchmanspipe (Aristolochia tomentosa), and branched tearthumb (Polygonum meisnerianum) (ISSG 2006). Infests public conservation lands in north and western Florida (FL-DEP 2008); the Florida Invasive Plants Geodatabase (FLInv) documents infestation of approximately 100 managed areas (FNAI 2008). Reported to occur in relatively remote natural areas, at least some of which are likely to contain high-quality communities.

Subrank II. Current Distribution and Abundance: Medium

6. Current Range Size in Nation:Moderate significance
Comments: Occurs in the southeastern United States and in Hawaii. In the southeast, it occurs mostly on the coastal plain, ranging from North Carolina to eastern Texas, south into Florida and north to southern Arkansas. In Florida, it is most densely established in the northern and western parts of the states, although it does range down into central Florida and as far south as Broward and Collier counties (Center for Aquatic and Invasive Plants 2008). On Hawaii, it occurs on the Big Island (Hawaii) and Oahu (Wilson 2001). In total, the generalized range occupies approximately 14% of the United States.

7. Proportion of Current Range Where the Species is Negatively Impacting Biodiversity:Moderate significance
Comments: Seems to be problematic across the southern half of the Gulf States, especially near the Gulf coast (Randall 1996, Byrd and Westbrooks 2007, Nation 2008), and in Florida, where it is particularly densely established in the north and west, but also problematic into the central part of the state (Langeland and Burks 1998). Further inland and away from the Gulf, it is described as rare (Weakley 2008) and found at scattered locations (Byrd and Westbrooks 2007). Does not appear to be particularly problematic in Hawaii; Wilson (2001) notes that it occasionally escapes from cultivation and persists, but that it does not seem to have spread significantly or to have been particularly damaging.

8. Proportion of Nation's Biogeographic Units Invaded:Low significance
Comments: Approximately 12 ecoregions are invaded, based on visual comparison of the generalized range and ecoregions map (The Nature Conservancy 2001).

9. Diversity of Habitats or Ecological Systems Invaded in Nation:High significance
Comments: Grows in both sunny and shady habitats, usually on damp soils (Langeland and Burks 1998, ISSG 2006 2006, Byrd and Westbrooks 2007, Center for Aquatic and Invasive Plants 2008, FL-DEP 2008, Nation 2008); thought to prefer soils of circumneutral pH (Langeland and Burks 1998). Often invades disturbed areas (e.g. roadsides, under and around bridges, yards, ditches, newly cleared or timbered land, pastures, fencerows), but can also invade natural undisturbed areas (Miller 2003, Weber 2003, ISSG 2006 2006, FL-DEP 2008, Weakley 2008). The Florida Invasive Plants Geodatabase (FLInv) (FNAI 2008) has recorded substantial numbers of infestations within the following natural communities: mesic upland forests (hardwood, pine, and mixed), mesic flatlands (e.g. mesic and scrubby flatwoods), floodplain and bottomland forests/swamps, wet flatlands (e.g. wet flatwoods and hydric hammocks), and communities alongside rivers and streams. Apparently also grows on the edges of lakes and marshes (Langeland and Burks 1998, Byrd and Westbrooks 2007, Nation 2008) and in estuarine habitats (ISSG 2006 2006). Known from cypress swamps in Louisiana and beech forests in east Texas (Randall 1996).

Subrank III. Trend in Distribution and Abundance: High/Medium

10. Current Trend in Total Range within Nation:Moderate significance
Comments: Range is quickly expanding south down the Florida peninsula from the apparent initial introduction point in northern Florida (1932) for ornamental purposes; now found as far south as Collier and Broward counties (Lott et al. 2003, Center for Aquatic and Invasive Plants 2008, FL-DEP 2008). Range also expanding in the Hawaiian islands, where it was first collected in 1936; apparently, numerous new naturalized populations have recently been reported on both Hawaii (big island) and Oahu (Wilson 2001).

11. Proportion of Potential Range Currently Occupied:Low significance
Comments: Reported to grow in USDA hardiness zones 8a - 11 (Whitinger 2008). North of the frost line, even though leaflets are typically killed by winter frost, the stalks of leaves usually remain intact (providing a ladder for climbing stalks of new growth the following year) and the rhizomes persist (Langeland and Burks 1998, Center for Aquatic and Invasive Plants 2008). Will probably expand northward somewhat from its current stronghold in the southern portion of the Gulf states (Nation 2008), as well as southward through additional parts of central and southern Florida (Center for Aquatic and Invasive Plants 2008, FL-DEP 2008), and to additional Hawaiian islands. Portions of western California, Oregon, and Washington may also be suitable.

12. Long-distance Dispersal Potential within Nation:High significance
Comments: Introduced to Florida in 1932 as an ornamental and still being used as such by unsuspecting gardeners (Miller 2003); available for sale on the internet (Whitinger 2008). Spores, produced in large numbers, are routinely transported long distances by wind (Langeland and Burks1998, Miller 2003, Byrd and Westbrooks 2007, Nation 2008) and, sometimes, by water (Center for Aquatic and Invasive Plants 2008). This species' vine-like growth is believed to promote long-distance wind dispersal because spores are released at or above the tree canopy (Lott et al. 2003). The small spores can also achieve long-distance dispersal by adhering to various human implements, such as vehicles, equipment, tires, clothing, and shoes, and possibly by adhering to animals (Langeland and Burks1998, Byrd and Westbrooks 2007, Center for Aquatic and Invasive Plants 2008). Spores also contaminate pine straw on infested pine plantations (Center for Aquatic and Invasive Plants 2008), from which they are transported when the straw is used as landscape mulch (Nation 2008). This species' ability to reproduce through intragametophytic selfing assists it in establishing new populations following long-distance dispersal events (Lott et al. 2003).

13. Local Range Expansion or Change in Abundance:Moderate significance
Comments: Locally expanding within its stronghold of northern and western Florida (Lott et al. 2003, ISSG 2006, FL-DEP 2008). Plant densities are described as continually increasing (Nation 2008) in southern Alabama (already significantly infested); density may be similarly increasing in at least southern Mississippi and Louisiana as well.

14. Inherent Ability to Invade Conservation Areas and Other Native Species Habitats:High/Moderate significance
Comments: Several sources describe this species as capable of invading disturbed or undisturbed areas (Byrd and Westbrooks 2007, Center for Aquatic and Invasive Plants 2008). It is believed to be a generalist able to invade a wide range of habitats (Nation 2008).

15. Similar Habitats Invaded Elsewhere:Medium/Low significance
Comments: Some sources report as native to Australia, but Bostock and Spokes (1998) list it as "non-native; known only from Darwin, N.T.; probably a garden escape." Also naturalized in Puerto Rico and Mexico (ISSG 2006). Considered weedy in the Philippines and Taiwan (Langeland and Burks 1998, Byrd and Westbrooks 2007). Little information was found regarding habitats invaded in these other regions.

16. Reproductive Characteristics:High significance
Comments: Stems are typically found 1-3 cm below the soil surface as widely creeping rhizomes (ISSG 2006), but sometimes occur aboveground as runners, where they can form layered mats on the ground surface (FL-DEP 2008); these stems persist through the winter even where leaflets are killed by winter frost (Center for Aquatic and Invasive Plants 2008). If the fronds are cut, burned, or otherwise damaged, the stem will quickly sprout replacements (Nation 2008). Persists and colonizes by these spreading stems and also spreads rapidly by wind-dispersed spores (Miller 2003). Grows quickly to reproductive maturity; sexually mature gametophytes are produced within 5 weeks of germination, and, once sexual maturity is reached, sporophyte production continues rapidly through week 12 (Lott et al. 2003). This ability to reproduce quickly is believed advantageous in the wet-dry seasonal cycles of Florida habitats (Lott et al. 2003). Many thousands of tiny spores are released per plant (FL-DEP 2008). Spores of Lygodium species typically have very thick walls, giving them long environmental viability (ISSG 2006).

Subrank IV. General Management Difficulty: Medium

17. General Management Difficulty:Moderate significance
Comments: Several sources described control as difficult or extremely difficult (Byrd and Westbrooks 2007, Nation 2008). Small infestations can sometimes be controlled by repeated hand pulling or cutting (Weber 2003); this technique is most likely to be successful when it is possible to remove the stems (at or slightly below the soil surface) as well as the aerial foliage. Even in this best-case scenario, plants sometimes regrow following hand-pulling (Center for Aquatic and Invasive Plants 2008). For larger infestations, vines can be pulled from the trees and treated with a foliar herbicide (Randall 1996, Weber 2003); machinery can be used to aid in removing foliage mats, but can cause soil compaction that is detrimental to native species (Center for Aquatic and Invasive Plants 2008). Several sources listed foliar herbicides that have proven to be reasonably effective (Randall 1996, Miller 2003, Weber 2003, ISSG 2006), although one source felt that acceptable chemical control options were limited and that more research is needed (Byrd and Westbrooks 2007). For biological control, no agents have yet been released, although research is ongoing (Byrd and Westbrooks 2007). The rust Puccinia lygodii, a pest of Lygodium spp. in greenhouses, may eventually hold promise, but assessment of potential negative impacts to the native L. palmatum is needed before large-scale releases can be considered (ISSG 2006, Center for Aquatic and Invasive Plants 2008).

18. Minimum Time Commitment:Moderate significance
Comments: Small spores can easily be missed during plant removal (Byrd and Westbrooks 2007). Spores of the Lygodium genus have very thick walls, giving these propagules long environmental viability (ISSG 2006).

19. Impacts of Management on Native Species:Moderate significance
Comments: Pulling vines from trees prior to herbicide treatment may harm native species associated with tree branches and trunks, such as epiphytes. If heavy machinery is used to aid in removing foliage mats from the trees, soil compaction may result, which could be detrimental to native species (Center for Aquatic and Invasive Plants 2008). Usually, fire is ruled out as a control measure because of anticipated damage to native vegetation from the intense crown fire that would likely develop (Center for Aquatic and Invasive Plants 2008). Some herbicides that provide effective control can kill or injure native plants via root uptake (Miller 2003), although presumably an herbicide that does not cause these effects could be selected for use in natural areas.

20. Accessibility of Invaded Areas:Moderate significance
Comments: Out of approximately 1400 infestations documented in the Florida Invasive Plants Geodatabase (FLInv), about 10% were rated as "difficult" to access and another 17% as "medium" to access; the remainder were rated "easy" to access (FNAI 2008).
Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 06Mar1991
NatureServe Conservation Status Factors Author: Broaddus, Lynn

Botanical data developed by NatureServe and its network of natural heritage programs (see Local Programs), The North Carolina Botanical Garden, and other contributors and cooperators (see Sources).

References
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  • Bostock, P.D. and T.M.Spokes. 1998. Polypodiaceae. Flora of Australia Online. Australian Biological Resources Study, Canberra. Online. Available: http://www.environment.gov.au/biodiversity/abrs/online-resources/flora/main/index.html (Accessed 2008).

  • Byrd, J. D. and R. Westbrooks. 2007. Lygodium japonicum (Thunb. ex Murr.) Sw.: Japanese climbing fern. in Invasive Plant Atlas of the MidSouth, a project of the GeoResources Institute (GRI) of Mississippi State University. Online. Available: http://www.gri.msstate.edu/research/ipams/Species.php?SName=Lygodium+japonicum (Accessed 2008).

  • Center for Aquatic and Invasive Plants. 2008. Japanese climbing fern: Lygodium japonicum. University of Florida, Institute of Food and Agricultural Sciences (IFAS). Online. Available: http://aquat1.ifas.ufl.edu/node/639 (Accessed 2008).

  • Florida Department of Environmental Protection (FL-DEP). 2008. Weed Alert: Japanese climbing fern (Lygodium japonicum). Online. Avalable: http://www.dep.state.fl.us/lands/invaspec/2ndlevpgs/pdfs/JapaneseClimbingFern.pdf (Accessed 2008)

  • Florida Natural Areas Inventory (FNAI). 2008, October last update. Florida Invasive Plants GeoDatabase (FLInv). Online. Available: http://www.fnai.org/invasivespecies.cfm (Accessed 2008)

  • Invasive Species Specialist Group (ISSG). 2006. Global Invasive Species Database. Online. Available: http://www.issg.org/database (Accessed 2008).

  • Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.

  • Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T., and C.A. Meacham. Synthesis of the North American Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill, N.C.

  • Langeland, K.A. and K.C. Burks. 1998. Identification and Biology of Non-Native Plants in Florida's Natural Areas. University of Florida. 165 pp. [http://aquat1.ifas.ufl.edu/identif.html]

  • Lott, M. S., J. C. Volin, R. W. Pemberton, and D. F. Austin. 2003. The reproductive biology of the invasive ferns Lygodium microphyllum and L. japonicum (Schizaeaceae): implications for invasive potential. American Journal of Botany 90: 1144-1152.

  • Miller, J.H. 2003. Nonnative Invasive Plants of Southern Forests: A field guide for identification and control. Gen. Tech. Rep. SRS-62. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 93 pp.

  • Nation, F. 2008. Plants of Alabama: Japanese climbing fern. Alabama Forestry Commission: Alabama's Treasured Forests series. Online. Available: http://www.forestry.state.al.us/Publications/TF/TFSummer08/Plants_of_Alabama_Japanese_Climbing_Fern.pdf (Accessed 2008)

  • Randall, J.M. and J. Marinelli (eds.) 1996. Invasive plants: weeds of the global garden. Brooklyn Botanic Garden, New York.

  • The Nature Conservancy. 2001. Map: TNC Ecoregions of the United States. Modification of Bailey Ecoregions. Online . Accessed May 2003.

  • USDA, ARS, National Genetic Resources Program. 2008 last update. Germplasm Resources Information Network (GRIN). National Germplasm Resources Laboratory, Beltsville, MD. Online. Available: http://www.ars-grin.gov/cgi-bin/npgs/html/index.pl (Accessed 2008).

  • Weakley, A. S. 2008. Flora of the Carolinas, Virginia, Georgia, northern Florida, and surrounding areas. Working Draft of 7 April 2008. University of North Carolina Herbarium (NCU), North Carolina Botanical Garden, University of North Carolina at Chapel Hill. Online. Available: http://herbarium.unc.edu/flora.htm (Accessed 2008).

  • Weber, E. 2003. Invasive plant species of the world: a reference guide to environmental weeds. CABI Publishing, Cambridge, Massachusetts. 548 pp.

  • Whitinger, D. 2008. Dave's Garden: PlantFiles. Online. Available: http://davesgarden.com/pf/ (Accessed 2008)

  • Wilson, K. A. 2001. Alien ferns in Hawai'i. Originally published in Pacific Science vol. 50 no. 2, 1996 by University of Hawaii Press. Used with permission. Updated and modified 2001. The Natural History Museum of Los Angeles County Foundation, Los Angeles, CA. Online. Available: http://www.rsabg.org/herbarium/ferns/wilson/toc.html

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