Invasive Species Compendium

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Datasheet

Lygodium japonicum
(Japanese climbing fern)

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Datasheet

Lygodium japonicum (Japanese climbing fern)

Summary

  • Last modified
  • 27 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Lygodium japonicum
  • Preferred Common Name
  • Japanese climbing fern
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Pteridophyta
  •       Class: Filicopsida
  •         Family: Schizaeaceae
  • Summary of Invasiveness
  • L. japonicum is a high-climbing vine that has become established outside of its native range of Asia. L. japonicum is widely cultivated around the world. In the 1900s it became established in the south...

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Pictures

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PictureTitleCaptionCopyright
Lygodium japonicum (Japanese climbing fern); habit, as an invasive plant on the Jones State Forest. Montgomery County, Texas, USA.
TitleHabit
CaptionLygodium japonicum (Japanese climbing fern); habit, as an invasive plant on the Jones State Forest. Montgomery County, Texas, USA.
Copyright©Ronald F. Billings/Texas A&M Forest Service/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); habit, as an invasive plant on the Jones State Forest. Montgomery County, Texas, USA.
HabitLygodium japonicum (Japanese climbing fern); habit, as an invasive plant on the Jones State Forest. Montgomery County, Texas, USA.©Ronald F. Billings/Texas A&M Forest Service/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); habit, showing foliage. An invasive plant in south-east Texas, USA.
TitleHabit
CaptionLygodium japonicum (Japanese climbing fern); habit, showing foliage. An invasive plant in south-east Texas, USA.
Copyright©Ronald F. Billings/Texas A&M Forest Service/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); habit, showing foliage. An invasive plant in south-east Texas, USA.
HabitLygodium japonicum (Japanese climbing fern); habit, showing foliage. An invasive plant in south-east Texas, USA.©Ronald F. Billings/Texas A&M Forest Service/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); infestation. Tift County, Georgia, USA. May, 2004.
TitleInfestation
CaptionLygodium japonicum (Japanese climbing fern); infestation. Tift County, Georgia, USA. May, 2004.
Copyright©Chris Evans/University of Illinois/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); infestation. Tift County, Georgia, USA. May, 2004.
InfestationLygodium japonicum (Japanese climbing fern); infestation. Tift County, Georgia, USA. May, 2004.©Chris Evans/University of Illinois/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); fronds. Onomea, Hawaii, Hawaii, USA. July, 2012.
TitleFronds
CaptionLygodium japonicum (Japanese climbing fern); fronds. Onomea, Hawaii, Hawaii, USA. July, 2012.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Lygodium japonicum (Japanese climbing fern); fronds. Onomea, Hawaii, Hawaii, USA. July, 2012.
FrondsLygodium japonicum (Japanese climbing fern); fronds. Onomea, Hawaii, Hawaii, USA. July, 2012.©Forest Starr & Kim Starr - CC BY 4.0
Lygodium japonicum (Japanese climbing fern); invasive habit, begining to climb tree. Georgia, USA. May, 2004.
TitleHabit
CaptionLygodium japonicum (Japanese climbing fern); invasive habit, begining to climb tree. Georgia, USA. May, 2004.
Copyright©Chris Evans/University of Illinois/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); invasive habit, begining to climb tree. Georgia, USA. May, 2004.
HabitLygodium japonicum (Japanese climbing fern); invasive habit, begining to climb tree. Georgia, USA. May, 2004.©Chris Evans/University of Illinois/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); foliage. Spore-producing fronds. Tift County, Georgia, USA. June, 2004.
TitleFoliage
CaptionLygodium japonicum (Japanese climbing fern); foliage. Spore-producing fronds. Tift County, Georgia, USA. June, 2004.
Copyright©David J. Moorhead/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); foliage. Spore-producing fronds. Tift County, Georgia, USA. June, 2004.
FoliageLygodium japonicum (Japanese climbing fern); foliage. Spore-producing fronds. Tift County, Georgia, USA. June, 2004.©David J. Moorhead/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); foliage. Unfertile frond (a) and fertile frond (b) showing fingerlike projections where spores are produced. Georgia, USA.
TitleFoliage
CaptionLygodium japonicum (Japanese climbing fern); foliage. Unfertile frond (a) and fertile frond (b) showing fingerlike projections where spores are produced. Georgia, USA.
Copyright©Karan A. Rawlins/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); foliage. Unfertile frond (a) and fertile frond (b) showing fingerlike projections where spores are produced. Georgia, USA.
FoliageLygodium japonicum (Japanese climbing fern); foliage. Unfertile frond (a) and fertile frond (b) showing fingerlike projections where spores are produced. Georgia, USA.©Karan A. Rawlins/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); small plant with sterile fronds. Torreya State Park, Bristol, Florida, USA. June, 2010.
TitleSmall plant
CaptionLygodium japonicum (Japanese climbing fern); small plant with sterile fronds. Torreya State Park, Bristol, Florida, USA. June, 2010.
Copyright©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); small plant with sterile fronds. Torreya State Park, Bristol, Florida, USA. June, 2010.
Small plantLygodium japonicum (Japanese climbing fern); small plant with sterile fronds. Torreya State Park, Bristol, Florida, USA. June, 2010.©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); young plants. Tift County, Georgia, USA. May, 2004.
TitleYoung plants
CaptionLygodium japonicum (Japanese climbing fern); young plants. Tift County, Georgia, USA. May, 2004.
Copyright©Chuck Bargeron/University of Georgia/Bugwood.org - CC BY 3.0 US
Lygodium japonicum (Japanese climbing fern); young plants. Tift County, Georgia, USA. May, 2004.
Young plantsLygodium japonicum (Japanese climbing fern); young plants. Tift County, Georgia, USA. May, 2004.©Chuck Bargeron/University of Georgia/Bugwood.org - CC BY 3.0 US
Lygodium japonicum (Japanese climbing fern); winter habit. USA. January.
TitleHabit
CaptionLygodium japonicum (Japanese climbing fern); winter habit. USA. January.
Copyright©James H. Miller/USDA Forest Service/Bugwood.org - CC BY 3.0 US
Lygodium japonicum (Japanese climbing fern); winter habit. USA. January.
HabitLygodium japonicum (Japanese climbing fern); winter habit. USA. January.©James H. Miller/USDA Forest Service/Bugwood.org - CC BY 3.0 US
Lygodium japonicum (Japanese climbing fern); prescribed fire as a control measure. USA.
TitleControl measure
CaptionLygodium japonicum (Japanese climbing fern); prescribed fire as a control measure. USA.
Copyright©Chuck Bargeron/University of Georgia/Bugwood.org - CC BY 3.0 US
Lygodium japonicum (Japanese climbing fern); prescribed fire as a control measure. USA.
Control measureLygodium japonicum (Japanese climbing fern); prescribed fire as a control measure. USA.©Chuck Bargeron/University of Georgia/Bugwood.org - CC BY 3.0 US
Lygodium japonicum (Japanese climbing fern); dispersal pathway. Movement of plant propagules on winch of off-road vehicle. USA. August, 2009.
TitleDispersal pathway
CaptionLygodium japonicum (Japanese climbing fern); dispersal pathway. Movement of plant propagules on winch of off-road vehicle. USA. August, 2009.
Copyright©Joseph LaForest/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Lygodium japonicum (Japanese climbing fern); dispersal pathway. Movement of plant propagules on winch of off-road vehicle. USA. August, 2009.
Dispersal pathwayLygodium japonicum (Japanese climbing fern); dispersal pathway. Movement of plant propagules on winch of off-road vehicle. USA. August, 2009.©Joseph LaForest/University of Georgia/Bugwood.org - CC BY-NC 3.0 US

Identity

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Preferred Scientific Name

  • Lygodium japonicum (Thunb.) Sw.

Preferred Common Name

  • Japanese climbing fern

Other Scientific Names

  • Hydroglossum japonicum (Desv.) Steud.
  • Lygodidum pubescens Kaulf.
  • Lygodium japonicum var. microstachyum (Desv.) C.Chr & Tardieu
  • Lygodium chaerophylloides Desv.
  • Lygodium cochinchinense (Poir.) Desv.
  • Lygodium dissectum Desv.
  • Lygodium japonicum f. elongata Alderw.
  • Lygodium mearnsii Copel.
  • Lygodium microstachyum Desv.
  • Lygodium microstachyum var. glabrescens Nakai
  • Lygodium tenue Blume
  • Ophioglossum japonicum Thunb.

International Common Names

  • Chinese: hai jin sha

Local Common Names

  • China: dewuli
  • Indonesia: hata kawat; madik silae; paka areuj; pakis kembang
  • Japan: kani-kusa; tsuru-shinobu
  • Korea, DPR: sil-go-sa-ri
  • Nepal: aankhle jhar; janai lahara; lute jhar; ukuse jhar
  • Pakistan: kharshi
  • Philippines: nito; nitong puti; nitoq; tat-awwa
  • Vietnam: bòng bong nhât

EPPO code

  • LYFJA (Lygodium japonicum)

Summary of Invasiveness

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L. japonicum is a high-climbing vine that has become established outside of its native range of Asia. L. japonicum is widely cultivated around the world. In the 1900s it became established in the south east USA and has since spread to at least nine states. L. japonicum occupies most of the southeastern coastal plain and parts of the Piedmont. Outside of its native range, L. japonicum has become an invasive species in a variety of habitats; floodplain forests, marshes and other wetlands, pine flatwoods, timber plantations and disturbed sites. It is a threat to natural areas where it outcompetes native species, alters fire behaviour and also poses an economic threat to the timber and pine straw industries. The distribution of the species continues to expand and could eventually occupy 39% of the USA; it has not yet reached the limit of distribution and abundance (USDA, 2009). Populations of L. japonicum have also become established on two Hawaiian Islands, Puerto Rico, Singapore and South Africa. Here, the species has not spread as aggressively as in the USA, but is of major concern. 

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Pteridophyta
  •             Class: Filicopsida
  •                 Family: Schizaeaceae
  •                     Genus: Lygodium
  •                         Species: Lygodium japonicum

Notes on Taxonomy and Nomenclature

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Lygodium is a genus of about 26 species distributed though the tropical regions of the world (Garrison Hanks, 1998; Flora of China Editorial Committee, 2014). The genus was traditionally placed in the Schizaceae, but recent molecular studies have supported the recognition of monospecific family the Lygodiaceae, containing only Lygodium (Garrison Hanks, 1998). It is distinctive in having fronds of unlimited growth, specialized pinnae and marginal sporangia.

Description

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L. japonicum is a rhizomatous vine, climbing to 30 m. Rhizomes creeping, with black to reddish-brown hairs. Stipes spaced to 1 cm apart on rhizome. Rachis grooved, pubescent to glabrous, 3-30 m. Pinnae with short stalks, 3-5 cm. Pinnae deltoid shaped, 2-3-pinnate, to ca. 12 cm long, 12 cm wide, usually with a long central lobe, margins lobulate. Costae with scattered hairs, veins and pinnae surfaces typically glabrous, rarely with short hairs. Fertile segments with 3-5 separate lobes, subpalmate, sporangia born on sorophores, each with 4-8 sporangia pairs. Spore diameters 64-80 µm, averaging 76 µm.

Distribution

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L. japonicum has a wide natural range in south, south-east and east Asia, including Bangladesh, Bhutan, China, India, Indonesia, Japan, Laos, Myanmar, Nepal, New Guinea, Pakistan, Philippines, North and South Korea, Sri Lanka, Taiwan, Thailand and Vietnam (Garrison Hanks, 1998; Chang et al., 2014; Ferns of Thailand, 2014; Flora of China Editorial Committee, 2014; USDA-ARS, 2015). In China it is primarily found south of the Yangtze River (Zheng et al., 2006).

Outside of its natural range L. japonicum has become naturalized in the USA in at least nine southeastern states (EDDMapS, 2015; USDA-NRCS, 2015) including Hawaii (Wilson, 2002). Records in Pennsylvania are suspect (B. Isaac, Carnegie Museum, USA, personal communication, 2014). It has become established in the Caribbean in Puerto Rico (Acevedo-Rodríguez, 2005), in South Africa (Henderson, 2007), Singapore (Chong et al., 2009) and Australia (Orchard and McCarthy, 1998).

L. japonicum has also been reported from Palau by Costion (2009) based on a specimen collected on Malakal Island in 1996. Whether this specimen is correctly identified is unknown. Three other species of Lygodium are known from Palau (Garrison Hanks, 1998). Costion (2009) notes that L. japonicum had previously been confused with L. salicifolium. Further, if the identification is correct it is unknown whether this is a natural population or a recent naturalization; Palau is 1300 and 1500 miles from New Guinea and the Philippines, respectively, the closest locations for this species.

There has been much confusion of the status of L. japonicum in Australia. The species was reported in the 1800s by von Mueller (1871) and Bentham (1878) and later without comment by Garrison Hanks (1998). However, Orchard and McCarthy (1998) considered previous reports to be of L. flexuosum, with L. japonicum being known as an escape from cultivation in the Northern Territory. For example, L. japonicum has been recorded in Western Australia but reports are from L. flexuosum. Hosking (2011) also reported L. japonicum to be naturalized in New South Wales. It is also in Queensland where it is considered to be an environmental weed (Queensland Government, 2014).

There are specimens in several herbaria labelled L. japonicum from localities other than those listed above, including Cuba, Honduras and Mexico (EDDMapS, 2015; Missouri Botanical Garden, 2015). These specimens are most likely to be of the closely related L. venustum. A report from Pennsylvania, USA (EDDMapS, 2015) is either from a cultivated plant or a waif occurrence (Bonnie Isaac, Carnegie Museum of Natural History, USA, personal communication, 2014). It was collected before 1909 by Hugo Andriessen in Beaver County without additional data. Edgar T. Wherry annotated the sheet as “escaped”. Andriessen was the proprietor of a combined Drug Store and Natural History museum (Applegate, 2015) and was actively trading fern specimens with other collectors (Andriessen, 1884). Some summaries of the species have noted its presence in California e.g. van Loan (2006a), but only specimens and reports from cultivation have been found in herbarium and literature searches.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

BangladeshPresentNativePrain, 1903; Uddin et al., 1997South, eastern and central Bangladesh
BhutanPresentNativeFlora of China Editorial Committee, 2014
CambodiaPresentNativeLindsay and Middleton, 2013; Missouri Botanical Garden, 20152012 specimen, O Chum District
ChinaPresentNativeFlora of China Editorial Committee, 2014
-AnhuiPresentNativeFlora of China Editorial Committee, 2014
-ChongqingPresentNativeFlora of China Editorial Committee, 2014
-FujianPresentNativeFlora of China Editorial Committee, 2014
-GansuPresentNativeFlora of China Editorial Committee, 2014
-GuangdongPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-GuangxiPresentNativeFlora of China Editorial Committee, 2014
-GuizhouPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-HainanPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-HenanPresentNativeFlora of China Editorial Committee, 2014
-HubeiPresentNativeFlora of China Editorial Committee, 2014
-HunanPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-JiangsuPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-JiangxiPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-ShaanxiPresentNativeFlora of China Editorial Committee, 2014
-ShanghaiPresentNativeFlora of China Editorial Committee, 2014
-SichuanPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-TibetPresentNativeFlora of China Editorial Committee, 2014
-YunnanPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-ZhejiangPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
IndiaPresentNativeBeddome, 1892; Garrison Hanks, 1998; Flora of China Editorial Committee, 2014North India, abundant, extending west to Kashmir, up to 5,000 feet elevation; South India, western mountains, rare
-Arunachal PradeshPresentNativeSingh et al., 2012
-AssamPresentNativeGarrison Hanks, 1998; Singh et al., 2012
-Indian PunjabPresentNativeGarrison Hanks, 1998
-Jammu and KashmirPresentNativeFlora of China Editorial Committee, 2014Kashmir
-MaharashtraPresent only in captivity/cultivationNativeBlatter and D'Almeida, 1922
-ManipurPresentNativeSingh et al., 2012
-MeghalayaPresentNativeSingh et al., 2012
-MizoramPresentNativeSingh et al., 2012
-NagalandPresentNativeSingh et al., 2012
IndonesiaPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014Java
-JavaPresentNativeHolttum, 1959; Maideen et al., 2004Central and east Java
-MoluccasPresentNativeHolttum, 1959; Maideen et al., 2004
-Nusa TenggaraPresentNativeMaideen et al., 2004; Wardani and Jaenudin, 2014Lombok
-SulawesiPresentNativeHolttum, 1959; Maideen et al., 2004
-SumatraPresentNativeMaideen et al., 2004Bangka Island (as “Banka”)
JapanPresentNativeOhwi, 1965; Garrison Hanks, 1998; Flora of China Editorial Committee, 2014
-HonshuPresentNativeOhwi, 1965; Flora of Japan, 2015South Tohoku district
-KyushuPresentNativeOhwi, 1965; Flora of Japan, 2015
-Ryukyu ArchipelagoPresentNativeFlora of Japan, 2015Ryukyu
-ShikokuPresentNativeOhwi, 1965; Flora of Japan, 2015
Korea, DPRPresentNativeFlora of China Editorial Committee, 2014
Korea, Republic ofPresentNativeGarrison Hanks, 1998; Chang et al., 2014; Flora of China Editorial Committee, 2014South Korea
LaosPresentNativeNewman et al., 2007; Lindsay and Middleton, 2013
MalaysiaPresentNativeHolttum, 1959
-SabahPresentNativeMaideen et al., 2004
MyanmarPresentNative
NepalPresentNativeGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
PakistanPresentNativeStewart, 1945; Khan et al., 2010
PhilippinesPresentNativeHolttum, 1959; Garrison Hanks, 1998; Maideen et al., 2004; Flora of China Editorial Committee, 2014
SingaporePresentIntroducedWee and Chua, 1988; Chong et al., 2009Naturalized for at least 5 years
Sri LankaPresentNativeHolttum, 1959; Garrison Hanks, 1998; Flora of China Editorial Committee, 2014
TaiwanPresentNativeLu and Yang, 2005; Flora of China Editorial Committee, 2014
ThailandPresentNativeGarrison Hanks, 1998; Lindsay and Middleton, 2013; Missouri Botanical Garden, 2015Northern, northeastern, and southwestern Thailand
VietnamPresentNativeThin, 1997; Garrison Hanks, 1998; List of Flora of Cat Tien National Park, 2015; Missouri Botanical Garden, 2015Cat Tien National Park and Cucphuong National Park

Africa

South AfricaPresentIntroducedMacdonald et al., 2003; Henderson, 2007Recorded as a casual alien

North America

USARestricted distributionEPPO, 2014
-AlabamaPresentIntroduced Invasive Underwood, 1906; Graves, 1920; Diamond and Woods, 2007; Woods and Diamond, 2008Mobile
-ArkansasPresentIntroduced1981 Invasive Sundell, 1986; Peck, 2011; Peck, 2011Peck (2011a) states this species was introduced in 1955. L. japonicum is recorded in 25 counties
-CaliforniaAbsent, unreliable recordLoan, 2006No records seen to corroborate
-FloridaPresentIntroduced1931 Invasive Small, 1938; Wunderlin et al., 2000; Hutchinson and Langeland, 2010Earliest herbarium record from 1931 (Polk County)
-GeorgiaPresentIntroduced1903 Invasive Clute, 1903; Underwood, 1906First report for USA, Thomasville, Georgia
-HawaiiPresentIntroduced1936 Invasive Fosberg, 1943; Wilson, 1996; Wilson, 2002; Wilson, 2003Hawaii and in 1998 Oahu
-KentuckyPresentEDDMapS, 2015McCreary County
-LouisianaPresent Invasive Reese and Thieret, 1966; Nauman, 1987Common and abundant
-MississippiPresentIntroduced Invasive Pullen et al., 1968; EDDMapS, 2015Recorded in 12 counties
-North CarolinaPresentIntroduced Invasive Wherry, 1964; Radford et al., 1968; EDDMapS, 2015Wilson and Lee counties. Lee County, recorded as a rare escape from cultivation
-OklahomaPresentIntroducedEDDMapS, 2015McCurtain County
-PennsylvaniaAbsent, unreliable recordB. Isaac, Carnegie Museum, personal communication, Dec. 4, 2014 Locality could be suspect
-South CarolinaPresentIntroduced Invasive Anderson, 1921; Matthews, 1940; Radford et al., 1968Dorchester and Sumter counties, recorded as a rare escape from cultivation
-TennesseePresentIntroducedEDDMapS, 2015Franklin County
-TexasPresentIntroduced1937 Invasive Soxman, 1939; Billings, 2005; Nesom, 2009

Central America and Caribbean

Puerto RicoPresentIntroducedGarrison Hanks, 1998; Liogier and Martorell, 2000; Acevedo-Rodríguez, 2005

Oceania

AustraliaPresentGarrison Hanks, 1998; Flora of China Editorial Committee, 2014
-Australian Northern TerritoryPresentIntroducedHosking et al., 2011; APNI, 2014
-New South WalesPresentIntroduced2000Hosking et al., 2011; Hosking et al., 2011; APNI, 2014
-QueenslandPresentIntroducedGarrison Hanks, 1998; Hosking et al., 2011; APNI, 2014
Papua New GuineaPresentNativeGarrison Hanks, 1998; Maideen et al., 2004

History of Introduction and Spread

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L. japonicum was first recorded outside of its natural range in the USA. It was first reported as an escape from cultivation in southern Georgia in 1903 (Clute, 1903). Reports of new records in additional states came quickly. Populations were found in Alabama in 1906 (Underwood, 1906), South Carolina in 1920 (Anderson, 1921), central Florida in 1931 (Hutchinson and Langeland, 2010), north Florida in 1932 (Small, 1938), Texas in 1937 (Soxman, 1939) and North Carolina by 1938 (Small, 1938). The species has spread throughout the southeastern USA and occurs in at least 9 states. There are recent reports from Oklahoma, Tennessee and Kentucky that need to be verified.

Fosberg (1943) reported L. japonicum as establishing in Hawaii in 1936 where it was collected at Hilo, around a garden. It has persisted and spread at this site and exists along road edges (Wilson, 2002; 2003). Wilson (1996) stated “the species does not seem to have spread substantially or to have been particularly damaging”. However, six years later Wilson (2002) reports that the status of the species had “changed significantly”; it was found at a new site on Hawaii and also in 1998 on Oahu (Wilson, 2003). Three new locations have since been reported on Oahu in 2012 and one more location on Hawaii (Lau and Frohlich, 2012).

The species is established in several localities in Puerto Rico (Acevedo-Rodríguez, 2005), the only location for it in the Caribbean. The first record is believed to be from a collection made by George Proctor in 1985 in San Juan on the grounds of the Estacion Experimental Agricola. Other collections and reports have since been made in Arecibo, Bayamón, Caguas and Guaynabo (Liogier and Martorell, 2000; Acevedo-Rodríguez, 2005).

L. japonicum is sparingly established as a “casual alien” in South Africa (Macdonald et al., 2003; Henderson, 2007), the only location for it on the African continent. There is a 1941 specimen, but it may have been from a cultivated plant (L. Henderson, SRC-Plant Protection Research Institute, South Africa, personal communication, 2014). According to Henderson the oldest confirmed naturalized record of L. japonicum is from 1985 (L. Henderson, SRC-Plant Protection Research Institute, South Africa, personal communication, 2014).

It is also considered to be introduced in Singapore where in 1983 it was found escaped around a house (Wee and Chua, 1988).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Australia 2000 Yes No Hosking et al. (2011)
Palau 1996 No No Costion (2009) Possibly native
Puerto Rico 1985 Yes No Garrison Hanks (1998)
Singapore ca. 1983 Yes No Wee and Chua (1988)
South Africa 1941 Yes No Macdonald et al. (2003) First established with certainty in 1985
USA 1903 Yes No Clute (1903)

Risk of Introduction

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The risk of L. japonicum being introduced to new areas is high due to the very small size of spores which are readily dispersed by the wind. Risk assessments have been conducted for the USA, Australia and the Pacific Islands (Koop et al., 2011; PIER, 2014). All found the species to be a high risk.

Habitat

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Within its natural range L. japonicum typically occupies moist habitats but favours regions with a defined dry season (Holttum, 1959; Garrison Hanks, 1998). It has weedy tendencies in its natural range and is primarily a pioneer and gap species. It is found from sea level to 2550 m (Holttum, 1959; Wardani and Jaenudin, 2014), primarily on southern slopes (Zheng et al., 2006). L. japonicum occupies secondary forests in China (Flora of China Editorial Committee, 2014), Indonesia (Wardani and Jaenudin, 2014) and Malaysia (Holttum, 1959) and forest edges and thickets in Japan (Ohwi et al., 1965) and the Philippines (Barcelona et al., 2006). In Bali, Indonesia L. japonicum was seen to occupy barren soil after a volcanic eruption (Dilmy, 1965).

Where introduced in the USA, L. japonicum occupies a broad range of natural and disturbed habitats. While L. japonicum has a strong preference for moist soils it can sometimes occupy xeric sites (van Loan, 2006b). It is invasive in floodplain forests, swamps, marshes, river and stream banks, pine flatwoods, hammocks, upland woodlands, lakeshores and low woods (Wunderlin et al., 2000; Diggs and George, 2006; van Loan, 2006b; Langeland et al., 2008; Miller et al., 2010a). It is a common invader of pine plantations (Ferriter, 2001; van Loan, 2006b; USDA, 2009; Miller et al., 2010a). In Puerto Rico it is known from disturbed areas and second growth forests (NYBG, 2015). In Hawaii it is known from a restoration site (Lau and Frohlich, 2012), road banks (Wilson, 1996), gardens and grassy hills (Wilson, 2002). In South Africa it has been found in moist forest, scrub and road edges (SIBIS, 2015) whereas in Australia it has been found in wet forests and riparian areas (Queensland Government, 2014).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Present, no further details Harmful (pest or invasive)
Managed forests, plantations and orchards Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Natural
Rail / roadsides Present, no further details
Terrestrial ‑ Natural / Semi-naturalNatural forests Present, no further details Harmful (pest or invasive)
Natural forests Present, no further details Natural
Riverbanks Present, no further details Harmful (pest or invasive)
Riverbanks Present, no further details Natural
Wetlands Present, no further details Harmful (pest or invasive)
Wetlands Present, no further details Natural
Rocky areas / lava flows Present, no further details Natural
Scrub / shrublands Present, no further details Harmful (pest or invasive)
Scrub / shrublands Present, no further details Natural
Freshwater
Irrigation channels Present, no further details Harmful (pest or invasive)
Irrigation channels Present, no further details Natural
Lakes Present, no further details Harmful (pest or invasive)
Lakes Present, no further details Natural
Reservoirs Present, no further details Harmful (pest or invasive)
Reservoirs Present, no further details Natural
Rivers / streams Present, no further details Harmful (pest or invasive)
Rivers / streams Present, no further details Natural
Ponds Present, no further details Harmful (pest or invasive)
Ponds Present, no further details Natural

Hosts/Species Affected

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In the Philippines L. japonicum has been listed as a weed of upland rice (Moody, 1989; Galinato et al., 1999).

In southeastern USA it is an invader of pine plantations (Pinus spp.) (Beasley and Pijut, 2010).

Biology and Ecology

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Genetics

L. japonicum is a tetraploid, with a chromosome number of 2n = 58 (Manton and Sledge, 1954; Roy and Holttum, 1965; Madeira et al., 2008). In a molecular study Madeira et al. (2008) showed that it belongs in a clade with L. polystachyum and L. venustum. According to Garrison Hanks (1998) hybrids with L. flexuosum probably occur where their ranges overlap.

Reproductive Biology

L. japonicum has a reproductive cycle typical of ferns, with alternating gametophyte and sporophyte generations. The sporophyte generation is the conspicuous generation and produces spores. Spores of L. japonicum are 64-80 mm (Murtaza et al., 2004). The species can self-fertilize (Lott et al., 2003; Srivastava and Uniyal, 2013) which is likely to be a major factor contributing to the naturalization of this species. Lott et al. (2003) found a high rate of selfing with 90% of isolated plants producing sporophytes. Sporophytes could reach sexual maturity within five weeks of germination (Lott et al., 2003). A study by Srivastava and Uniyal (2013) found that 20% of spores were abnormal and that sporophyte production was lower for plants that selfed compared to plants which outcrossed (58% vs. 30%).

Physiology and Phenology

L. japonicum is a perennial vine that is deciduous in colder climates and diesback in the winter when exposed to frosts (Garrison Hanks, 1998; van Loan, 2006a; Langeland et al., 2008; Miller et al., 2010a). New fronds arise from subterranean rhizomes from March through to May and continue to grow until frost causes front dieback (van Loan, 2006a). Fronds can climb to 30 m on other vegetation, including shrubs and trees. It will also climb dead, persistent rachises from fronds of previous years. Frond growth is indeterminate. If the frond apex is damaged or cut, new growth will initiate from arrested growing points from bases of pinnae (Mueller, 1983). According to Mueller (1983) this provides an advantage in shaded conditions, allowing the plant to grow from sunnier positions in canopies, instead of having to regrow from the ground under existing vegetation. Vines can grow up to 6.5 cm/day (Mueller, 1983). The subterranean rhizomes have starch grains in adventitious roots that serve as storage organs (Clarke, 1936). In frost free years in the southern part of its range plants will remain green through the winter. L. japonicum is adapted to cold temperatures; Hutchinson and Langeland (2014) found that freezing temperatures do not affect spores. Gametophytes were sensitive to freezing temperatures, but could survive when exposed for several hours. Spore production can take place all year round in warmer climates, but in colder areas it typically occurs from June-November and peaks in October (van Loan, 2006a).       

Carmichael (2012) found that fire and other disturbance events such as windstorms promoted conditions favourable to the invasion of L. japonicum by creating canopy gaps that could be colonized.

Environmental Requirements

L. japonicum may have a preference for soils with a circumneutral pH (Nauman, 1987; Diggs and George, 2006; Langeland et al., 2008). 

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Tolerated > 60mm precipitation per month
Am - Tropical monsoon climate Tolerated Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
Aw - Tropical wet and dry savanna climate Tolerated < 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
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 Tolerated Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Austromusotima camptozonale Herbivore Leaves to genus
Glomerella cingulata Pathogen Leaves
Milesia lygodii Pathogen Leaves to genus
Neomusotima conspurcatalis Herbivore Leaves to genus
Neomusotima fuscolinealis Herbivore Leaves to genus
Octothrips bhattii Herbivore Leaves to genus

Notes on Natural Enemies

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There have been few reports of natural enemies of L. japonicum (Ferriter, 2001). The South American fungal pathogen Puccinia lygodii [Milesia lygodii] was first reported in Florida by Rayachhetry et al. (2001). Van Loan (2006b) later observed this fungus from additional sites in Florida and Louisiana. It caused severe damage to leaflets and in some stands 95% of foliage was affected. Another fungal pathogen, Colletotrichum gloeosporioides, has infested plants in Florida (Jones et al., 2003).

Three moths have been found to feed on L. japonicum: Austromusotima camptozonale, Neomusotima conspurcatalis and Neomusotima fuscolinealis. These have been subject to testing as biological control agents (Madeira et al., 2008; Boughton et al., 2009; Langeland and Hutchinson, 2013). The thrip insect Octothrips lygodii [Octothrips bhattii] was discovered on L. japonicum in China (Mound, 2002) and Ferriter (2001) reports a flea beetle (Coleoptera: Chrysomelidae) on L. japonicum in Thailand.

Means of Movement and Dispersal

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Natural Dispersal

The ability to produce large numbers of very small, wind borne spores, combined with the propensity to self, has given L. japonicum the capability to disperse over long distances (Ferriter, 2001; Lott et al., 2003). Nauman (1987) suggested that in Florida the species appeared to follow rivers in its spread, a trait that may also apply within its natural range.

Accidental Introduction

Dust-like spores of L. japonicum can adhere to a wide variety of surfaces. It has been hypothesized that they can be transported on clothing, vehicles, animal fur and cut timber (Ferriter, 2001; van Loan, 2006a; Langeland et al., 2008). The pine straw industry has also been implicated in the movement of the species (van Loan, 2006b; Miller et al., 2010a). In much of the southeastern USA the fallen needles of Pinus species are harvested as a non-timber forest product for use as landscape mulch. Since L. japonicum grows in pine-dominated habitats and in pine plantations, collection and movement of pine straw serves to distribute the spores to new locations. Van Loan (2006a) hypothesized that spores on the bark of trees could be moved during forestry operations.

Intentional Introduction

Cultivation as an ornamental has been the primary vector for the introduction of L. japonicum to regions outside of its natural range. It has been cultivated in the southeastern USA since the late 1800s. It was probably this species that was sold by Royal Palm Nurseries in Florida starting in 1888-1889 and called L. microphyllum (Pemberton and Ferriter, 1998; Ferriter, 2001). Clarke (1936) purchased L. japonicum from Royal Palm nursery, sold to him incorrectly as L. scandens. Early reports of the species naturalizing in the USA clearly indicate that plants were cultivated nearby, including Clute (1903) near a greenhouse in Georgia and Anderson (1921), from a florist and gardens in South Carolina.

Impact Summary

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CategoryImpact
Economic/livelihood Negative
Environment (generally) Negative

Economic Impact

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L. japonicum invasions have had a negative impact in the USA because the species is an invader of commercial pine plantations and hardwood plantations (USDA, 2009; Beasley and Pijut, 2010). Owners of plantations must use chemical control to reduce population sizes. Van Loan (2006b) estimates that control costs are $45-$450/acre.

The species has also had a negative impact on the pine straw industry. Fallen needles of Pinus species are collected from pine plantations or other pine-dominated areas and used as landscaping mulch. In Florida alone this is a $79 million industry. Spores residing in pine straw are transported to new areas from infested collection sites. The USDA (2009) found L. japonicum to be a threat to wood production systems and was likely harming wildlife viewing and hunting.

Environmental Impact

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Impact on Habitats

L. japonicum invasions can range from very sparse, with small numbers of individuals, to dense monocultures. The fronds trellis into trees, creating a novel strata of biomass between the ground and as much as 30 m into tree canopies. This creates changes in light levels and fuel levels. A major concern is the influence of this fuel structure on fire behaviour. When large vines or populations burn, fires can easily spread into canopy trees from the ground. Fires may also be able to penetrate into wetland areas that would otherwise be barriers (Munger, 2005).

Impact on Biodiversity

L. japonicum outcompetes native plant species by altering penetration of light to the forest floor. In established populations existing herb and understory species can be reduced or eliminated due to overshading. For example, Leichty et al. (2011) found that L. japonicum is likely to compete with mesic pine savannah species in Louisiana. It has been demonstrated that herbaceous layers respond positively when L. japonicum is treated with herbicide (Minogue et al., 2010; Bohn et al., 2011).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Thalictrum cooleyiNo DetailsFlorida; North CarolinaCompetition - shadingUSDA, 2009

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Tolerant of shade
  • Highly mobile locally
  • Benefits from human association (i.e. it is a human commensal)
  • Fast growing
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Host damage
  • Modification of fire regime
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts agriculture
  • Negatively impacts forestry
  • Negatively impacts livelihoods
  • Negatively impacts tourism
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
  • Damages animal/plant products
Impact mechanisms
  • 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
  • Difficult/costly to control

Uses

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A wide variety of uses have been reported for L. japonicum in its native range, including uses as medicine, decoration, landscaping, crafts, pesticide and food.

The most common use of the species is as a folk medicine. In China it is used as a diuretic (Puri, 1970) and to treat colds, inflammation, kidney stones and renal ailments (Eisenberg et al., 2009). In India it is used as an expectorant and to treat snakebites (Reutter, 1923; Puri, 1970; Srivastava and Uniyal, 2013) and to treat diabetes, wounds and ulcers (Yumkham and Singh, 2011). In Nepal a paste is used to treat scabies, the juice is used to treat herpes and wounds (Manandhar, 1995) and its juice is applied for boils, wounds, whitlow and scabies (Mall et al., 2015). In Pakistan a powder is used on wounds to help healing and a root extract for reducing body aches and swelling (Khan et al., 2010). Spores are sold on the Internet for use in traditional Chinese medicine as “Spora Lygodii” (Ferriter, 2001).

Formal studies have been conducted to determine medicinal benefits of L. japonicum. Duan et al. (2012) showed that this species has compounds with strong antioxidant properties. Cho et al. (2014) found it to be useful as a preventive and therapeutic agent against the formation of oxalate kidney stones, supporting one of the primary traditional uses. It has also been studied with potentially positive results for the regrowth of hair (Matsuda et al., 2002).

Lygodium species, including L. japonicum, have been used in many parts of the world for weaving and binding (Reid and Madulid, 1972; Laarman et al., 1995). Srivastava and Uniyal (2013) noted its use in worship in India. It is used as a pesticide and insect repellent in China (Yang and Tang, 1988; Zhang et al., 2014) and occasionally cooked as a vegetable (Xu et al., 2004; Liu et al., 2012; Zhang et al., 2014). Many members of the genus Lygodium are used as an attractive, fast growing ornamental fern.

Detection and Inspection

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When mature, L. japonicum is easily detected on the ground due to its unique morphology and high-climbing habitat. However, because the species can disperse easily into remote areas via windborne spores new populations are easily overlooked because of difficulty of access.

Similarities to Other Species/Conditions

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In its natural range L. japonicum can intergrade with L. flexuosum, which usually has shorter pinna stalks, serrate to dentate margins (not lobulated) and segments entire to auriculate (Garrison Hanks, 1998).

In the USA L. japonicum has two congeners, one native (L. palmatum) and one introduced (L. microphyllum). L. palmatum overlaps the range of L. japonicum from Louisiana to North Carolina. It is unique in having palmately lobed pinnae. In peninsular Florida L. japonicum overlaps with L. microphyllum, which is also an invasive species. L. microphyllum pinnae are primarily 1-pinnate, while those of L. japonicum are typically 2-3-pinnate.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Prevention

Containment/Zoning

Attempts have been made in Alabama and Florida to limit the movement of L. japonicum through legislative means. In Alabama it is a quarantined species with transport into or within the state prohibited (Nation, undated). It has been listed by the State of Florida as a Noxious Weed. It is considered a controlled landscape species in Miami-Dade County, Florida where it cannot be planted within 500 feet of a native plant community (Hunsberger, 2001). It is also regulated in Collier and Okaloosa counties, Florida (van Loan, 2006b). It is listed as a Category I Invasive Species by the Florida Exotic Pest Plant Council (FLEPPC, 2013). In Texas, Nesom (2009) ranked it as “F1”, invasive in disturbed and natural habitats.

Control

Cultural Control and Sanitary Measures

Spores of L. japonicum can adhere to clothing and vehicles. It has therefore been recommended that land managers and researchers working in proximity to L. japonicum clean clothing and equipment before traveling to new areas (Hutchinson and Langeland et al., 2008; Miller et al., 2010b).

The use of fire as a potential control agent has been examined by Minogue et al. (2009) and Leichty et al. (2011), but it has been found that fire is not effective at controlling L. japonicum.

Movement Control

The commercial pine straw industry has been identified as a vector for the movement of L. japonicum spores into new areas and it has been recommended that where possible spore free sources are used (van Loan, 2006a; Miller et al., 2010a; Miller et al., 2010b).

Biological Control

The potential for biocontrol of Lygodium species has been a subject of active research due to the invasiveness of two species in the genus in the USA: L. japonicum and L. microphyllum (Ferriter, 2001). While biocontrol agents have been released for L. microphyllum, no agents have been released specifically for L. japonicum. Finding a biocontrol suitable for this species will be difficult due to the sympatric, native species L. palmatum.

In 2005 a moth, Austromusotima camptozonale, was released in Florida for the control of L. microphyllum but failed to establish (Langeland and Hutchinson, 2013). A second species of moth Neomusotima conspurcatalis (Lygodium defoliator moth) was released in peninsular Florida in 2008 and 2009 as a biocontrol of L. microphyllum and populations have become established (Langeland and Hutchinson, 2013). This moth is genus specific and also feeds on L. japonicum, however, it is sensitive to cold temperatures. It was approved for release because of this characteristic as it will not pose a threat to the temperate native L. palmatum. While there are populations of L. japonicum in frost-free areas that will be impacted by N. conspurcatalis, populations outside of Florida will not be in the range of the moth (Madeira et al., 2008; Boughton et al., 2009). A second species in the genus, N. fuscolinealis, was also screened however it was rejected as it posed a threat to L. palmatum (Bennett and Pemberton, 2008). 

Chemical Control

Chemical control of L. japonicum has been studied in a number of ecosystems in the southeastern USA. Lockhart (2005) recommends cutting or pulling vines and spraying piles of foliage to minimize non-target damage. Researchers and land managers have come to a consensus that treatments using glyphosate are effective. Recommended concentrations do vary, including 1.5% by Langeland and Stocker (2001), 2% by Bohn et al. (2011) and 4% by Miller et al. (2010b). Lower concentrations should be used when limiting non-target damage is desirable. Some use glyphosate with other herbicides, including metsulfuron-methyl and imazapyr. Minogue et al. (2009) had the best results using 4% glyphosate with metsulfuron-methyl. Lockhart (2005) warns that this combination, while effective, should not be used in close proximity to cabbage palm trees (Sabal palmetto) because they are very sensitive to metsulfuron-methyl. Formulations using imazapyr cause excessive non-target damage (Minogue et al., 2009). Applications of 2,4-D in Florida pine plantations have not been effective (Ferriter, 2001). Retreats are always necessary and should be started 6-12 months after initial treatment and continued at regular intervals (Lockhart, 2005).

References

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31/01/2015 Original text by:
 
Keith A. Bradley, South Carolina, USA

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