Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

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Blechnum appendiculatum
(palm fern)

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Datasheet

Blechnum appendiculatum (palm fern)

Summary

  • Last modified
  • 27 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Blechnum appendiculatum
  • Preferred Common Name
  • palm fern
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Pteridophyta
  •       Class: Pteridopsida
  •         Family: Blechnaceae
  • Summary of Invasiveness
  • B. occidentale is a medium sized fern, native to the Americas, that has become naturalized in Hawaii following escape from cultivation. Preferring shady conditions in mesic forests, particularly on rocks or roc...

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Pictures

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PictureTitleCaptionCopyright
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
TitleHabit
CaptionBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
HabitBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Waikapu Valley, Maui, Hawaii, USA. April, 2012.
TitleHabit
CaptionBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Waikapu Valley, Maui, Hawaii, USA. April, 2012.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Waikapu Valley, Maui, Hawaii, USA. April, 2012.
HabitBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Waikapu Valley, Maui, Hawaii, USA. April, 2012.©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit and habitat. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
TitleHabit
CaptionBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit and habitat. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit and habitat. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
HabitBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit and habitat. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
TitleHabit
CaptionBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
HabitBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit, showing fronds. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
TitleFronds
CaptionBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit, showing fronds. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); habit, showing fronds. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.
FrondsBlechnum occidentale (palm fern, catalogued as B. appendiculatum); habit, showing fronds. Honokowai Ditch Trail, Maui, Hawaii, USA. June 2010.©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); sori. Kula Botanical Garden, Maui, Hawaii, USA. March 2011.
TitleSori
CaptionBlechnum occidentale (palm fern, catalogued as B. appendiculatum); sori. Kula Botanical Garden, Maui, Hawaii, USA. March 2011.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); sori. Kula Botanical Garden, Maui, Hawaii, USA. March 2011.
SoriBlechnum occidentale (palm fern, catalogued as B. appendiculatum); sori. Kula Botanical Garden, Maui, Hawaii, USA. March 2011.©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); sori. West Maui, Maui, Hawaii, USA. February 2009.
TitleSori
CaptionBlechnum occidentale (palm fern, catalogued as B. appendiculatum); sori. West Maui, Maui, Hawaii, USA. February 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Blechnum occidentale (palm fern, catalogued as B. appendiculatum); sori. West Maui, Maui, Hawaii, USA. February 2009.
SoriBlechnum occidentale (palm fern, catalogued as B. appendiculatum); sori. West Maui, Maui, Hawaii, USA. February 2009.©Forest Starr & Kim Starr - CC BY 4.0

Identity

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Preferred Scientific Name

  • Blechnum appendiculatum L.

Preferred Common Name

  • palm fern

Other Scientific Names

  • Blechnum appendiculatum Willd.
  • Blechnum extensum Fée
  • Blechnum glandulosum Kaulf. ex Link
  • Blechnum glandulosum var. distans (C. Presl) C. Chr.
  • Blechnum helveolum Fée
  • Blechnum meridionale C. Presl
  • Blechnum mucronatum Fée
  • Blechnum occidentale var. intermedium (Link) E. Fourn.
  • Blechnum occidentale var. lacerata Rosenstock
  • Blechnum occidentale var. pubirhachis Rosenstock

International Common Names

  • English: hammock fern

Local Common Names

  • USA: New World midsorus fern

Summary of Invasiveness

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B. occidentale is a medium sized fern, native to the Americas, that has become naturalized in Hawaii following escape from cultivation. Preferring shady conditions in mesic forests, particularly on rocks or rocky substrates, and being able to survive periods of drought, it produces sporophytes that can be long-lived and spread asexually by branching or budding. It can form large clonal colonies that suppress native plants, and is listed as one of the species causing habitat degradation and impacting endangered native plants in Hawaii. The large populations form mats effectively preventing the germination and growth of native species (Wilson, 1996). 

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Pteridophyta
  •             Class: Pteridopsida
  •                 Family: Blechnaceae
  •                     Genus: Blechnum
  •                         Species: Blechnum appendiculatum

Notes on Taxonomy and Nomenclature

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The Blechnaceae is a family of 24 genera and 265 species (Gasper et al., 2016a; PPG I, 2016).The family has centres of diversity and endemism in the Neotropics and Australasia/Oceania, with a predominance of species and genera in the southern hemisphere (Gasper et al., 2016a). The genus Blechnum s.s. includes 30 species (PPG I, 2016). Most species are neotropical, with a few in southern Africa (Gasper et al., 2016a).

Blechnum appendiculatum was described by Carl Ludwig von Willdenow in 1810. The species epithet appendiculatum comes from the Latin appendicular, diminutive of appendix, meaning small appendages or projections, and atus, indicating possession. This possibly refers to the short glandular hairs on the rachises (Palmer, 2003). B. appendiculatum has been treated both as a separate species, and as an infraspecific taxon within, B. occidentale. Mickel and Smith (2004) noted that morphological differences between B. appendiculatum and B. occidentale (sensu stricto) “appear correlated with differences in elevation and pubescence, and there are relatively few specimens that are difficult to place in one species or the other.” In the most recent treatment of the B. occidentale group, however, Dittrich et al. (2015) found “a nearly continuous gradation in specimens from glabrous to few hairs to many hairs, indicating that this can be treated as the same taxon without any discontinuity between these variations”. This datasheet follows Dittrich et al. and treats B. occidentale as the accepted name, with B. appendiculatum as a synonym.

Description

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The following description is from Dittrich et al. (2015): Plants terrestrial, rarely epipetric; rhizomes erect to decumbent, stoloniferous, the scales mostly bicolorous, linear-lanceolate or narrowly triangular, with a dark central stripe, 3.5–8.0 × 1.0–1.9 mm at base, margin entire; fronds monomorphic, 12.9–71.9 cm long; stipes 3.2–34.3 cm long, 1.5–2.1 mm diam., espcially at the base with tan, concolorous scales, the margin predominantly entire, with few denticles; blade 9.7–37.6 × 2.2–19.8 cm, ovallanceolate, ovate or deltoid, papyraceous to subcoriaceous, truncate at base, without vestigial pinnae, sometimes the pair of proximal pinnae slightly smaller than those immediately above, tapered toward apex, pinnate at base, pinatissect toward apex, glabrous or with multicellular trichomes abaxially on the veins; rachis glabrous or with multicellular, tan, hyaline trichomes; pinnae 10–27 pairs, 1.3–11.0 × 0.8–1.6 cm, reflexed or patent (basal ones), patent or ascending (median ones) or ascending (apical ones), sessile (basal ones), partially (median ones) or fully adnate (apical ones), falcate, apex acute, cuneate or mucronate, glabrous or rarely with trichomes on abaxial costa similar to those of abaxial rachis, basal ones acroscopically auriculate, the auricles usually overlapping the rachis, margins finely denticulate, but often slightly revolute and appearing entire; veins free, simple or 1–4(–5)-bifurcate, with clavate ends before the margin.

Distribution

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B. occidentale is currently found in the Americas, from Texas and Florida in the USA, down as far south as Argentina and Paraguay in South America, and including some islands in the Caribbean (Dittrich et al., 2015; USDA-ARS, 2015). It is introduced and invasive in Hawaii (PIER, 2015). While USDA-ARS (2015) lists it as native throughout its range in the Americas, Seigler and Lockwood (1975) suggested that the USA populations had probably arisen from spores blown by the wind from Mexico or the West Indies. Diggs and Lipsocmb (2015) list it as rare and with limited distribution in Texas.

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

North America

MexicoPresentNativeMissouri Botanical Garden, 2015; USDA-ARS, 2015Sal Luis Potosi, Tamaulipas, Chiapas, Colima, Federal District, Guerrero, Hidalgo, Jalisco, Mexico, Michoacan, Morelos, Nayarity, Oaxaca, Puebla, Queretaro, Tlaxcala, Veracruz
USAPresentPresent based on regional distribution.
-FloridaPresentNativeUSDA-ARS, 2015
-HawaiiWidespreadIntroduced Invasive Palmer, 2003
-TexasLocalisedIntroducedDiggs and Lipscomb, 2015; USDA-ARS, 2015Limited distribution, of conservation concern in the state

Central America and Caribbean

BelizePresentDittrich et al., 2015
Costa RicaPresentNativeUSDA-ARS, 2015
CubaPresentMissouri Botanical Garden, 2015
Dominican RepublicPresentMissouri Botanical Garden, 2015
El SalvadorPresentNativeUSDA-ARS, 2015
Greater AntillesPresentDittrich et al., 2015
GuatemalaPresentNativeUSDA-ARS, 2015
HaitiPresentMissouri Botanical Garden, 2015
HondurasPresentNativeUSDA-ARS, 2015
JamaicaPresentMissouri Botanical Garden, 2015
Lesser AntillesPresentDittrich et al., 2015
NicaraguaPresentNativeUSDA-ARS, 2015
PanamaPresentNativeUSDA-ARS, 2015
Trinidad and TobagoPresentNativeUSDA-ARS, 2015Trinidad
United States Virgin IslandsPresentMissouri Botanical Garden, 2015

South America

ArgentinaPresentNativeUSDA-ARS, 2015
BoliviaPresentNativeMissouri Botanical Garden, 2015; USDA-ARS, 2015
BrazilPresentNativeUSDA-ARS, 2015
-AmazonasPresentNativeDittrich et al., 2015
-BahiaPresentNativeDittrich et al., 2015
-CearaPresentNativeDittrich et al., 2015
-Espirito SantoPresentNativeDittrich et al., 2015
-GoiasPresentNativeDittrich et al., 2015
-Mato GrossoPresentNativeDittrich et al., 2015
-Mato Grosso do SulPresentNativeDittrich et al., 2015
-Minas GeraisPresentNativeDittrich et al., 2015
-ParaPresentNativeDittrich et al., 2015
-ParanaPresentNativeDittrich et al., 2015
-PernambucoPresentNativeDittrich et al., 2015
-Rio de JaneiroPresentNativeDittrich et al., 2015
-Rio Grande do SulPresentNativeDittrich et al., 2015
-RoraimaPresentNativeDittrich et al., 2015
-Santa CatarinaPresentNativeDittrich et al., 2015
-Sao PauloPresentNativeDittrich et al., 2015
ChilePresentNativeDittrich et al., 2015
ColombiaPresentNativeUSDA-ARS, 2015
EcuadorPresentNativeUSDA-ARS, 2015
French GuianaPresentNativeDittrich et al., 2015
GuyanaPresentNativeDittrich et al., 2015
ParaguayPresentNativeUSDA-ARS, 2015
PeruPresentNativeUSDA-ARS, 2015
SurinamePresentNativeDittrich et al., 2015
VenezuelaPresentNativeUSDA-ARS, 2015

History of Introduction and Spread

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According to Palmer (2003), B. occidentale was first collected in Hawaii in 1918 having escaped from gardens. It has spread extensively to other parts of Hawaii, and is a clonal naturalized species. 

Habitat

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In its native habitat, B. occidentale inhabits the interior of forests, most often growing along forest edges, trails, roads and rarely on rocks near streams (Dittrich et al., 2015). It occurs between 40 and 1600 m, and is present in seasonal deciduous and semideciduous forests, dense and mixed humid forests.

In Hawaii, large colonies are formed by clonal plants in closed-canopy mesic forests in all but the most extreme habitats at 30-1560 m in all islands (Palmer, 2003). Wilson (1996) reported it as weedy along trail sides, stream banks, forested slopes, and gulches on all islands, often growing in solid stands. It is common over rocks and rocky substrates, and in Texas it is found in granite crevices (Diggs and Lipscomb, 2015). 

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedDisturbed areas Present, no further details Natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Principal habitat Natural
Riverbanks Present, no further details Natural
Rocky areas / lava flows Present, no further details Natural

Biology and Ecology

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Adult ferns produce sporangia on the underside of the leaves and from these spores are expelled which, when they find a suitable habitat, develop into gametophytes. These form antheridia that produce many sperm and archegonia that produce a single ovum. Once maturation occurs, and in the presence of a thin film of water on the underside of the gametophyte, the sperm actively swim towards the ovum and fertilise it. Mechanisms prevent self-fertilization. The fertilized ovum now has two set of chromosomes and develops into a sporophyte. In Blechnum spp., the sporophytes can be long-lived and spread asexually by producing stolons that bud off new plants thereby forming large colonies (Palmer, 2003).

B. occidentale adapts to disturbance, is environmentally versatile, and thrives under drought conditions (Weller et al., 2011). 

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])

Means of Movement and Dispersal

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The spores can be dispersed by wind, and probably by water (PIER, 2015). Diggs and Lipscomb (2015) quote Seigler and Lockwood (1975) as suggesting that populations in the USA have probably arisen from spores blown by the wind from Mexico or the West Indies.

Impact Summary

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CategoryImpact
Environment (generally) Negative

Environmental Impact

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All the reports for negative environmental impact of this fern come from Hawaii, where it is an introduced species which competes against a number of native plants. B. occidentale forms thick mats that prevent establishment of seeds and spores of other plants (Wilson, 1996). Weller et al. (2011) report that B. occidentale increased in percentage cover during a drought period when native ferns decreased substantially. It competes with a number of native fern species, having particular impact on the rare genus Diellia (Palmer, 2003). Aguraiuja et al. (2004), for example, reported that on Kauai a site investigated for Diellia pallida was completely overgrown by Erigeron karvinskianus and B. occidentale. A review of the endangered Diellia erecta (US Fish and Wildlife Service, 2007) reports competition with invasive species including B. occidentale as a threat to the species. In IUCN assessments of the critically endangered fern Asplenium diellaciniatum (Kishida and Wood, 2015) and the endangered plant Cyrtandra heinrichii (Lorence and Edmonds, 2015), both found on Kauai Island in the Hawaiian Island chain, B. occidentale is listed as one of the alien species causing habitat degradation. The fern is also listed as degrading habitat for Nototrichium humile (US Fish and Wildlife Service, 2008a).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Asplenium diellaciniatumCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)HawaiiKishida and Wood, 2015
Asplenium dielpallidumNo DetailsHawaiiCompetitionClark, 2015
Cyrtandra heinrichiiEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)HawaiiCompetitionLorence and Edmonds, 2015
Diellia erectaNo DetailsHawaiiCompetitionUS Fish and Wildlife Service, 2007
Nototrichium humile (kaala rockwort)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resourcesNational Tropical Botanical Garden, 2007; US Fish and Wildlife Service, 2008a
Phyllostegia hirsuta (Molokai phyllostegia)NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetitionUS Fish and Wildlife Service, 2008b
Phyllostegia mollis (Waianae Range phyllostegia)USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetitionUS Fish and Wildlife Service, 2009b
Pittosporum napaliense (royal cheesewood)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resources; Competition - smotheringUS Fish and Wildlife Service, 2010b; US Fish and Wildlife Service, 2010d
Plantago princepsNatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resourcesUS Fish and Wildlife Service, 2010e
Platydesma rostrataCR (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; Competition - smotheringUS Fish and Wildlife Service, 2010d
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 hardyi (Makaleha pritchardia)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - smotheringUS Fish and Wildlife Service, 2010f
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 - smotheringUS Fish and Wildlife Service, 2010d
Remya kauaiensis (Kauai remya)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition (unspecified)US Fish and Wildlife Service, 2010c
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 speciesHawaiiCompetition (unspecified)US Fish and Wildlife Service, 2010g
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
Pteris lidgatei (Lidgate's brake)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, 2009a
Stenogyne purpurea (purplefruit stenogyne)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - smotheringUS Fish and Wildlife Service, 2010f
Bidens campylotheca subsp. pentamera (ko`oko`olau)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as threatened species USA ESA listing as threatened speciesHawaiiCompetition (unspecified)US Fish and Wildlife Service, 2013
Peucedanum sandwicense (makou)NatureServe NatureServe; USA ESA listing as threatened species USA ESA listing as threatened speciesHawaiiCompetition - shadingUS Fish and Wildlife Service, 2011a

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Is a habitat generalist
  • Tolerant of shade
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - shading
  • Competition - smothering
  • Competition

Similarities to Other Species/Conditions

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B. occidentale is close to B. austrobrasilianum and B. laevigatum. It is also similar to the hybrids B. × caudatum and B. × leopoldense. A key to identification is given in Dittrich et al. (2015).

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.

Aguilar-Dorantes et al. (2015) report that glyphosate inhibits spore germination of B. occidentale even at low concentrations. The herbicide also killed the fern at all green life stages at most concentrations tested. However, herbicides should be used with care as associated plant species would also be affected. Mechanical control is likely to be ineffective given vegetative reproduction via stolons (PIER, 2015).

References

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Aguilar-Dorantes K, Mehltreter K, Mata-Rosas M, Vibrans H, Esqueda-Esquivel V, 2015. Glyphosate susceptibility of different life stages of three fern species. American Fern Journal, 105(3):131-144. http://www.bioone.org/loi/amfj

Aguraiuja R, Moora M, Zobel M, 2004. Population stage structure of Hawaiian endemic fern taxa of Diellia (Aspleniaceae): implications for monitoring and regional dynamics. Canadian Journal of Botany, 82(10):1438-1445.

Andrews SB, 1990. Ferns of Queensland., Australia: Queensland Dept. of Primary Industries, 427 pp.

Clark M, 2015. Asplenium dielpallidum. The IUCN Red List of Threatened Species, 2015(e.T78774640A78774709). http://dx.doi.org/10.2305/IUCN.UK.2015-4.RLTS.T78774640A78774709.en

Diggs G, Lipscomb B, 2015. Ferns of Texas. http://ferns.brit.org/

Dittrich VAO, Salino A, Monteiro R, 2015. The Blechnum occidentale (Blechnaceae, Polypodiopsida) species group in southern and southeastern Brazil. Phytotaxa, 231(3):201-229.

Gasper AL, Almeida TE, Dittrich VAO, Smith AR, Salino A, 2016. Molecular phylogeny of the fern family Blechnaceae (Polypodiales) with a revised genus-level treatment. Cladistics. DOI: 10.1111/cla.12173

Gasper AL, Dittrich VAO, Smith AR, Salino A, 2016. A classification for Blechnaceae (Polypodiales: Polypodiopsida): new genera, resurrected names, and combinations. Phytotaxa, 275(3):191-227.

Kishida W, Wood K, 2015. Asplenium diellaciniatum. The IUCN Red List of Threatened Species, e.T80229055A80229060. http://dx.doi.org/10.2305/IUCN.UK.2015-4.RLTS.T80229055A80229060.en

Lorence DH, Edmonds M, 2015. Cyrtandra heinrichii. The IUCN Red List of Threatened Species, e.T80231759A80231775. http://dx.doi.org/10.2305/IUCN.UK.2015- 4.RLTS.T80231759A80231775.en

Mickel JT, Smith AR, 2004. The pteridophytes of Mexico. New York, USA: New York Botanical Garden.

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

National Tropical Botanical Garden, 2007. https://ntbg.org/

Palmer DD, 2003. Hawaii's Ferns and Fern Allies. Honolulu, Hawaii, USA: University of Hawaii Press, 324 pp.

PIER, 2015. Pacific Islands Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

PPGI, 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution, 54(6):563-603.

Seigler DS, Lockwood TE, 1975. Blechnum occidentale new to Texas. American Fern Journal, 65:96.

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

US Fish and Wildlife Service, 2007. 5-year review, Diellia erecta. 9 pp. http://www.fws.gov/ecos/ajax/docs/five_year_review/doc2466.pdf

US Fish and Wildlife Service, 2008. In: Kulu`i (Nototrichium humile). 5-Year Review. US Fish and Wildlife Service, 11 pp.

US Fish and Wildlife Service, 2008. In: Phyllostegia hirsuta (no common name). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 10 pp.

US Fish and Wildlife Service, 2009. In: 5-Year Review, Short Form Summary: Species Reviewed: Pteris lidgatei (no common name). US Fish and Wildlife Service, 7 pp.

US Fish and Wildlife Service, 2009. In: Phyllostegia mollis (no common name). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 13 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: 5-Year Review, Short Form Summary: Species Reviewed: Pritchardia napaliensis (loulu palm). US Fish and Wildlife Service, 10 pp.

US Fish and Wildlife Service, 2010. In: 5-Year Review, Short Form Summary: Species Reviewed: Remya kauaiensis (no common name). US Fish and Wildlife Service, 12 pp.

US Fish and Wildlife Service, 2010. In: Determination of Endangered Status for 48 Species on Kauai and designation of Critical Habitat: Final Rule. US Fish and Wildlife Service, i + 205 pp.

US Fish and Wildlife Service, 2010. In: Plantago princeps (laukahi kuahiwi). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 19 pp.

US Fish and Wildlife Service, 2010. In: Recovery Outline for the Kauai Ecosystem. US Fish and Wildlife Service, 38 pp. + 3 maps.

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: Endangered and Threatened Wildlife and Plants; Listing 23 Species on Oahu as Endangered and Designating Critical Habitat for 124 Species. 76(148) US Fish and Wildlife Service, 46362-46593. http://www.gpo.gov/fdsys/pkg/FR-2011-08-02/pdf/2011-17162.pdf

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.

US Fish and Wildlife Service, 2013. In: Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for 38 Species on Molokai, Lanai, and Maui; Final Rule. 78(102) US Fish and Wildlife Service, 32014-32065. https://www.gpo.gov/fdsys/pkg/FR-2013-05-28/pdf/2013-12105.pdf

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

Weller SG, Cabin RJ, Lorence DH, Perlman S, Wood K, Flynn T, Sakai AK, 2011. Alien plant invasions, introduced ungulates, and alternative states in a mesic forest in Hawaii. Restoration Ecology, 19(5):671-680. http://onlinelibrary.wiley.com/doi/10.1111/j.1526-100X.2009.00635.x/full

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

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29/02/16 Original text by:

Emma Thompson, Egham, CABI

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