Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Paspalum distichum
(knotgrass)

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Datasheet

Paspalum distichum (knotgrass)

Summary

  • Last modified
  • 19 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Host Plant
  • Preferred Scientific Name
  • Paspalum distichum
  • Preferred Common Name
  • knotgrass
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Monocotyledonae
  • Summary of Invasiveness
  • P. distichum is a fast-growing rhizomatous grass of wet areas. It has become a major weed of rice and many other crops, as well as occurring in uncropped wetlands in both its native and introduced regions. Its...

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Pictures

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PictureTitleCaptionCopyright
Infestation of irrigation channel.
TitleP. distichum infestation
CaptionInfestation of irrigation channel.
Copyright©Chris Parker/Bristol, UK
Infestation of irrigation channel.
P. distichum infestationInfestation of irrigation channel.©Chris Parker/Bristol, UK
Vegetative stage.
TitleVegetative stage
CaptionVegetative stage.
Copyright©Chris Parker/Bristol, UK
Vegetative stage.
Vegetative stageVegetative stage.©Chris Parker/Bristol, UK
Inflorescences.
TitleInflorescences
CaptionInflorescences.
Copyright©Chris Parker/Bristol, UK
Inflorescences.
InflorescencesInflorescences.©Chris Parker/Bristol, UK
a, Ligule, ventral view; b, part of spike-like raceme (pseudo-spike); c1-2, spikelet, two views; d, lower glume (G1), dorsal view; e, upper glume (G2), dorsal view (opened); f, lower lemma (L1), ventral view; g, upper floret; h, upper lemma (L2), ventral view; i, upper palea (P2) ventral view; j, caryopsis.
TitleP. distichum - line drawing
Captiona, Ligule, ventral view; b, part of spike-like raceme (pseudo-spike); c1-2, spikelet, two views; d, lower glume (G1), dorsal view; e, upper glume (G2), dorsal view (opened); f, lower lemma (L1), ventral view; g, upper floret; h, upper lemma (L2), ventral view; i, upper palea (P2) ventral view; j, caryopsis.
CopyrightSEAMEO-BIOTROP
a, Ligule, ventral view; b, part of spike-like raceme (pseudo-spike); c1-2, spikelet, two views; d, lower glume (G1), dorsal view; e, upper glume (G2), dorsal view (opened); f, lower lemma (L1), ventral view; g, upper floret; h, upper lemma (L2), ventral view; i, upper palea (P2) ventral view; j, caryopsis.
P. distichum - line drawinga, Ligule, ventral view; b, part of spike-like raceme (pseudo-spike); c1-2, spikelet, two views; d, lower glume (G1), dorsal view; e, upper glume (G2), dorsal view (opened); f, lower lemma (L1), ventral view; g, upper floret; h, upper lemma (L2), ventral view; i, upper palea (P2) ventral view; j, caryopsis.SEAMEO-BIOTROP
P. distichum inflorescence.
TitleInflorescence
CaptionP. distichum inflorescence.
CopyrightTomas Marquez/DuPont-Spain
P. distichum inflorescence.
InflorescenceP. distichum inflorescence.Tomas Marquez/DuPont-Spain

Identity

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Preferred Scientific Name

  • Paspalum distichum L. 1759

Preferred Common Name

  • knotgrass

Other Scientific Names

  • Digitaria paspalodes Michx. (1803)
  • Panicum paspalodes Michx. (1803)
  • Paspalum paspalodes
  • Paspalum paspaloides (Michx.) Scribn. 1894
  • Paspalum paucispicatum Vasey (1893)

International Common Names

  • English: couch paspalum; dallisgrass; devil's grass; ditch-grass; eternity grass; ginger grass; jointgrass; mercer grass; seashore paspalum; seaside millet; silt grass; thompsongrass; victoria grass; water couch; wiregrass
  • Spanish: alcanache; camalote saladillo; capim-arame; chepica; grama colorado; grama de agua; grama-braba; gramilla blanca; gramilla dulce; pasto dulce
  • French: paspale a deux epis
  • Chinese: Hong-ban-gen-cao; liang er cao; shuang sui que bai
  • Portuguese: alcanache; graminhão

Local Common Names

  • Brazil: capim-pancuan; grama-de-forquilha; grama-doce; pancuam
  • Cuba: rapiente
  • Germany: Knotgras
  • Indonesia: lamhani
  • Iraq: shalhaw
  • Italy: gramignone d'acqua; panico aquatico; paspalo distico
  • Japan: Chikugo-suzumenohie; Kishu-suzume-no-hie
  • Nepal: ghunde banso
  • Portugal: grama de Joanopolis
  • South Africa: bankrotkweek; kweekpaspalum

EPPO code

  • PASDS (Paspalum distichum)

Summary of Invasiveness

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P. distichum is a fast-growing rhizomatous grass of wet areas. It has become a major weed of rice and many other crops, as well as occurring in uncropped wetlands in both its native and introduced regions. Its introduction to Europe, Asia and the Pacific is not well documented but apparently occurred many years ago. New records are reported in e.g. Indonesia, Spain and Croatia, suggesting that it continues to spread in countries to which it has been introduced.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Monocotyledonae
  •                     Order: Cyperales
  •                         Family: Poaceae
  •                             Genus: Paspalum
  •                                 Species: Paspalum distichum

Notes on Taxonomy and Nomenclature

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Both P. distichum and P. paspalodes are accepted names for this weed. The use of the Linnaean name ‘P. distichum’ was challenged by Renvoize and Clayton (1980) as they considered he had applied the name to a specimen of P. vaginatum on the same original herbarium sheet. This was in turn challenged by Guedes (1981) and after further deliberation by the International Association for Plant Taxonomy (IAPT) the original challenge was disallowed (Brummitt, 1983). Hence the original, more popular name P. distichum is retained for this data sheet, though P. paspalodes is still used by a number of authorities including Flora Europaea (Royal Botanic Garden Edinburgh, 2009).

There has been corresponding uncertainty over the spelling of the main synonym – ‘paspalodes’ often appearing as ‘paspaloides.’ In 1894, Scribner published the new combination ‘Paspalum paspaloides’, based on the earlier name Digitaria paspalodes Michx. 1803, but taking the liberty of ‘correcting’ the spelling from ‘paspalodes’ to ‘paspaloides’. However, this correction is not allowed under the rules of nomenclature. Hence, the use of the spelling ‘paspaloides’, although widespread, is not strictly correct (Henry Noltie, personal communication). FAO (2009) still uses ‘P. paspaloides’ for P. distichum as defined here (and ‘P. distichum’ for what is normally referred to as P. vaginatum).

A number of synonyms have been listed in the Identity section, but there are a great many more quoted by e.g. Bor (1960) and by Missouri Botanic Garden (2009).

The use of the name ‘P. distichum’ by Renvoize and Clayton for the closely related P. vaginatum, together with some understandable confusion in field identification means that there are inevitably some records of P. distichum in this data sheet which may more correctly be P. vaginatum.

A range of varieties have been named. In Japan and Korea, var. indutum is distinguished from the typical form and does show some differences in ecology etc. Most recently in France, P. distichum var. paucispicatum has been described, corresponding to Mexican material described originally as P. paucispicatum (Verloove and Reynders, 2007). A mutant type, ‘Flexi-green’, has been registered as a turf grass in Australia.

There is also plenty of confusion with regard to the popular names used; dallisgrass is applied to both P. distichum and P. dilalatum; knotgrass is the officially recognised common name in the USA, but in the UK this applies to the unrelated Polygonum aviculare.

Description

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The morphology of P. distichum has been described in detail by Chase (1929). It is a widely creeping perennial with slender rhizomes, extensively stoloniferous, often forming loose mats. The stolons are usually slender, subcompressed, sometimes as much as one metre long. On average, the sheaths are less loose than in Paspalum vaginatum, the blades are usually well developed; the branches are erect or ascending, most of them finally flowering, 6-50 cm tall, often sparingly branching. The culms are subcompressed, the nodes dark, often with a few ascending hairs; the sheaths are loose, keeled, and commonly pilose on the margins toward the summit. The ligule is membranaceous, about 0.5 mm long; the blades are flat, ascending, 3-12 cm long, and 2-6 mm wide at the rounded, ciliate base, tapering to an acuminate, sometimes involute apex; they are dull green, relatively soft in texture, and occasionally minutely pubescent on the upper surface. The peduncles are commonly short, often included; there are usually two racemes, rarely as many as four, from erect to reflexed, commonly incurved, 1.5-7 cm long, rarely longer. The rachis is slightly pedunculate in one, sometimes both racemes, usually with a few white hairs in the axil, 1-1.5 mm, rarely 2 mm wide, triangular, and minutely scabrous on the margin. The spikelets are solitary (rarely in pairs in the middle of the raceme), imbricate, 2.5-3 mm wide (size variation is sometimes found in the same plant), elliptic, abruptly acute, and pale green. The first glume is frequently developed; the second glume and sterile lemma are equal, with three to five veins, the midrib is relatively prominent. The glume is minutely appressed-pubescent, sometimes obscurely so. The fruit is 2.5-2.8 mm long, about 1.2 mm wide, and elliptic. Pollen morphology has been described by Ma GuoHua et al. (2001).

P. distichum is unusual in the variability in size of the lower glume: even within a single raceme it can vary from absent to conspicuous, up to half the length of the spikelet.

P. distichum exhibits considerable variability, thought to result from both the heteroploidy shown by the species, and the existence of genetic differences among individuals of the same cytotype (Echarte et al., 1992).

Plant Type

Top of page Aquatic
Grass / sedge
Herbaceous
Perennial
Seed propagated
Vegetatively propagated

Distribution

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P. distichum has almost world-wide distribution in tropical and subtropical regions where temperature and moisture conditions favour its C4 growth habit. Distribution is throughout North America, except Canada; Mexico, Central America, and all of South America; southern Europe; southern former USSR; the Middle East; the Indian subcontinent; South-East Asia; China; Japan; Korea Republic and Korea Democratic People's Republic; the Philippines; Australia; New Zealand; and the Pacific Islands. It occurs in southern and North Africa, but not in East, West or Central Africa.

It is far from clear just where P. paspalum is truly native. It is generally accepted that it is native in N. and S. America and introduced to Europe and most of Asia. PIER (2009) suggests that it is native in the Pacific area, though introduced to Hawaii, Australia and New Zealand. Other sources suggest it could also be native in southern Africa and in Australia.

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

BangladeshWidespreadIntroduced Invasive Pradhan and Chettri, 1987
BhutanWidespreadIntroduced Not invasive Pradhan and Chettri, 1987; Noltie, 2000
Brunei DarussalamPresentIntroducedWaterhouse, 1993
CambodiaPresentIntroducedJahn and Bunnarith, 2004
ChinaWidespreadIntroducedGong and Li, 1991; Waterhouse, 1993
-AnhuiPresentIntroducedMissouri Botanical Garden, 2009
-FujianPresentIntroducedMissouri Botanical Garden, 2009
-GuangdongWidespreadIntroducedLi, 1981
-GuangxiWidespreadIntroducedLi, 1981; Missouri Botanical Garden, 2009
-GuizhouPresentMissouri Botanical Garden, 2009
-HainanPresentIntroducedMissouri Botanical Garden, 2009
-HenanPresentIntroducedMissouri Botanical Garden, 2009
-Hong KongPresentIntroducedMissouri Botanical Garden, 2009
-HubeiWidespreadLi, 1981; Missouri Botanical Garden, 2009
-HunanPresentIntroducedWang, 1990; Missouri Botanical Garden, 2009
-JiangsuWidespreadIntroducedLi, 1981; Missouri Botanical Garden, 2009
-ShandongPresentIntroducedMissouri Botanical Garden, 2009
-SichuanPresentIntroducedMissouri Botanical Garden, 2009
-YunnanWidespreadLi, 1981; Missouri Botanical Garden, 2009
-ZhejiangPresentIntroducedMissouri Botanical Garden, 2009
Christmas Island (Indian Ocean)PresentIntroducedHäfliger and Scholz, 1980
Cocos IslandsPresentIntroducedHäfliger and Scholz, 1980
IndiaPresentIntroducedHäfliger and Scholz, 1980
-Andaman and Nicobar IslandsPresentIntroducedShukla, 1996
-AssamPresentIntroducedShukla, 1996
-BiharPresentIntroducedShukla, 1996
-DelhiPresentIntroducedShukla, 1996
-HaryanaPresentIntroducedAshok et al., 2007
-Himachal PradeshPresentIntroducedShukla, 1996
-Jammu and KashmirPresentIntroducedShukla, 1996
-Madhya PradeshPresentIntroducedShukla, 1996
-MaharashtraPresentIntroducedSrinivasan and Subbian, 1991; Shukla, 1996
-ManipurPresentIntroducedShukla, 1996
-MeghalayaPresentIntroducedShukla, 1996
-NagalandPresentIntroducedShukla, 1996
-OdishaPresentIntroducedMishra and Bhol, 1996
-RajasthanPresentIntroducedMiddleton, 1998
-SikkimPresentIntroducedSingh, 1992
-Tamil NaduPresentIntroducedBalasubramanian and Krishnarajan, 2001
-Uttar PradeshPresentIntroducedShukla, 1996
-West BengalPresentIntroducedShukla, 1996
IndonesiaWidespreadIntroducedWaterhouse, 1993
-JavaWidespreadIntroduced1970s Invasive Everaarts, 1981Only recently noticed on Java in 1979
IranPresentIntroduced Invasive Holm et al., 1979; Häfliger and Scholz, 1980
IraqWidespreadIntroducedTownsend et al., 1968
IsraelWidespreadIntroducedHolm et al., 1979
JapanPresentIntroduced Invasive Häfliger and Scholz, 1980; Shibayama, 1988
-KyushuWidespreadIntroduced Invasive Häfliger and Scholz, 1980; Le, 1991
-Ryukyu ArchipelagoPresentIntroducedHossain, 2005
Korea, Republic ofPresentIntroducedHäfliger and Scholz, 1980
LaosPresentIntroducedHäfliger and Scholz, 1980
MalaysiaPresentIntroducedHäfliger and Scholz, 1980; Waterhouse, 1993
MyanmarPresentIntroducedWaterhouse, 1993
NepalPresentIntroducedRanjit and Bhattarai, 1988
PakistanPresentIntroduced Invasive Holm et al., 1979; Häfliger and Scholz, 1980
PhilippinesWidespreadIntroduced Invasive Bernasor and de Datta, 1988; Fajardo and Moody, 1990; Waterhouse, 1993
Sri LankaPresentIntroducedHolm et al., 1979
TaiwanWidespreadIntroducedHäfliger and Scholz, 1980; Missouri Botanical Garden, 2009
ThailandPresentIntroducedFalvey and Hengmichai, 1978; Waterhouse, 1993
TurkeyPresentIntroducedTownsend et al., 1968; Royal Botanic Garden Edinburgh, 2009; EPPO, 2014
VietnamPresentIntroducedHäfliger and Scholz, 1980; Waterhouse, 1993

Africa

AlgeriaPresentIntroducedLe, 1991
EgyptPresentIntroducedTownsend et al., 1968
LibyaPresentIntroducedLe, 1991
MadagascarPresentIntroducedUSDA-ARS, 2009
MauritiusPresentIntroducedPIER, 2009
MoroccoPresentIntroducedHolm et al., 1979
SeychellesPresentIntroducedPIER, 2009Agalega Island (PIER, 2009)
South AfricaPresentIntroducedHolm et al., 1979
SwazilandPresentIntroduced Invasive Holm et al., 1979
ZimbabwePresentIntroducedClayton, 1989

North America

BermudaPresentNativeChase, 1929
MexicoPresentNativeHäfliger and Scholz, 1980
USAPresentNativeChase, 1929; Häfliger and Scholz, 1980; USDA-ARS, 2009
-AlabamaPresentNativeChase, 1929; USDA-ARS, 2009
-ArizonaPresentNativeChase, 1929; USDA-ARS, 2009
-ArkansasPresentNativeChase, 1929; USDA-ARS, 2009
-CaliforniaPresentNativeChase, 1929; Robbins et al., 1951; USDA-ARS, 2009
-FloridaPresentNativeChase, 1929; USDA-ARS, 2009
-GeorgiaPresentNativeChase, 1929; USDA-ARS, 2009
-HawaiiPresentIntroduced Invasive Häfliger and Scholz, 1980; PIER, 2009Kaua'l O'aha Islands (PIER, 2009)
-IdahoPresentNativeUSDA-ARS, 2009
-KansasPresentNativeUSDA-ARS, 2009
-KentuckyPresentNativeUSDA-ARS, 2009
-LouisianaPresentChase, 1929; Echarte et al., 1992
-MarylandPresentNativeUSDA-ARS, 2009
-MississippiPresentNativeChase, 1929; USDA-ARS, 2009
-MissouriPresentNativeUSDA-ARS, 2009
-NevadaPresentNativeChase, 1929; USDA-ARS, 2009
-New JerseyPresentChase, 1929
-New MexicoPresentChase, 1929
-North CarolinaPresentChase, 1929
-OklahomaPresentNativeChase, 1929; USDA-ARS, 2009
-OregonPresentNativeChase, 1929; USDA-ARS, 2009
-PennsylvaniaPresentNativeChase, 1929; USDA-ARS, 2009
-South CarolinaPresentNativeChase, 1929; USDA-ARS, 2009
-TennesseePresentNativeChase, 1929; USDA-ARS, 2009
-TexasPresentNativeChase, 1929; Gould, 1965; USDA-ARS, 2009
-UtahPresentNativeChase, 1929; Gould, 1965; USDA-ARS, 2009
-VirginiaPresentChase, 1929
-WashingtonPresentNativeUSDA-ARS, 2009

Central America and Caribbean

Antigua and BarbudaPresentNativeChase, 1929
BahamasPresentNativeChase, 1929
BelizePresentNativeChase, 1929; Häfliger and Scholz, 1980
Costa RicaPresentNativeChase, 1929; Häfliger and Scholz, 1980
CubaWidespreadNativeChase, 1929; Labrada, 1975
DominicaPresentNativeHäfliger and Scholz, 1980
Dominican RepublicPresentNativeChase, 1929; Häfliger and Scholz, 1980
El SalvadorPresentNativeHäfliger and Scholz, 1980; Häfliger and Scholz, 1980
GuatemalaPresentNativeChase, 1929
HaitiPresentNativeChase, 1929
HondurasPresentMissouri Botanical Garden, 2009
JamaicaPresentNativeChase, 1929
MartiniquePresentNativeChase, 1929
NicaraguaPresentNativeMissouri Botanical Garden, 2009
PanamaPresentNativeHäfliger and Scholz, 1980
Puerto RicoPresentNativeChase, 1929; Häfliger and Scholz, 1980; USDA-ARS, 2009
Saint LuciaPresentNativeChase, 1929
Saint Vincent and the GrenadinesPresentNativeChase, 1929
Trinidad and TobagoPresentNativeChase, 1929
United States Virgin IslandsPresentNativeChase, 1929; USDA-ARS, 2009

South America

ArgentinaPresentNativeChase, 1929; Häfliger and Scholz, 1980
BoliviaPresentNativeChase,1929; Häfliger and Scholz, 1980
BrazilPresentNativeChase, 1929; Häfliger and Scholz, 1980
-AlagoasPresentNativeLorenzi, 1982
-BahiaPresentNativeLorenzi, 1982
-Espirito SantoPresentNativeLorenzi, 1982
-GoiasPresentNativeLorenzi, 1982
-Mato Grosso do SulPresentNativeLorenzi, 1982
-Minas GeraisPresentNativeLorenzi, 1982
-ParanaPresentNativeLorenzi, 1982
-PernambucoPresentNativeLorenzi, 1982
-Rio de JaneiroPresentNativeLorenzi, 1982
-Rio Grande do SulPresentNativeLorenzi, 1982
-Santa CatarinaPresentNativeLorenzi, 1982
-Sao PauloPresentNativeLorenzi, 1982
-SergipePresentNativeLorenzi, 1982
ChilePresentNative Invasive Chase, 1929; Häfliger and Scholz, 1980
ColombiaPresentNative Invasive Chase, 1929; Häfliger and Scholz, 1980
EcuadorPresentNativeChase, 1929
French GuianaPresentMissouri Botanical Garden, 2009
GuyanaPresentNativeHolm et al., 1979
ParaguayPresentNativeChase, 1929; Häfliger and Scholz, 1980
PeruPresentNativeChase, 1929; Häfliger and Scholz, 1980
SurinamePresentNativeChase, 1929
UruguayPresentNativeChase, 1929; Häfliger and Scholz, 1980
VenezuelaPresentNativeHäfliger and Scholz, 1980

Europe

AlbaniaPresentRoyal Botanic Garden Edinburgh, 2009; EPPO, 2014
BulgariaPresentRoyal Botanic Garden Edinburgh, 2009; EPPO, 2014
CroatiaPresentIntroducedPandza and Tafra, 2008
FranceRestricted distributionIntroducedHäfliger and Scholz, 1980; Mesleard et al., 1993; Royal Botanic Garden Edinburgh, 2009; EPPO, 2014
GreecePresentIntroduced Invasive Häfliger and Scholz, 1980; Le, 1991; Royal Botanic Garden Edinburgh, 2009; EPPO, 2014
ItalyPresentIntroducedHäfliger and Scholz, 1980; Le, 1991; Royal Botanic Garden Edinburgh, 2009; EPPO, 2014
MaltaPresentEPPO, 2014
PortugalWidespreadIntroduced Invasive Häfliger and Scholz, 1980; Royal Botanic Garden Edinburgh, 2009; EPPO, 2014
-AzoresPresentRoyal Botanic Garden Edinburgh, 2009; EPPO, 2014
-MadeiraPresentIntroducedWeber, 2003
RomaniaPresentOprea et al., 2011
Russian FederationPresentHäfliger and Scholz, 1980
-Central RussiaPresentIntroducedHäfliger and Scholz, 1980; Tsvelev, 1984
-Russia (Europe)PresentHäfliger and Scholz, 1980; Tsvelev, 1984
-Russian Far EastPresentHäfliger and Scholz, 1980; Tsvelev, 1984
-Southern Russia Invasive Royal Botanic Garden Edinburgh, 2009
SloveniaRestricted distributionIntroducedHäfliger and Scholz, 1980
SpainPresentIntroducedHäfliger and Scholz, 1980; Le, 1991; Royal Botanic Garden Edinburgh, 2009; EPPO, 2014
UKLocalisedIntroduced Not invasive Stace, 1991
UkrainePresentEPPO, 2014

Oceania

AustraliaPresentHäfliger and Scholz, 1980; USDA-ARS, 2009
-Australian Northern TerritoryPresentIntroduced Not invasive PlantNet, 2009
-New South WalesWidespreadIntroduced Not invasive Häfliger and Scholz, 1980; PlantNet, 2009
-QueenslandPresentIntroduced Not invasive PlantNet, 2009
-South AustraliaPresentIntroduced Not invasive PlantNet, 2009
-TasmaniaPresentIntroduced Not invasive PlantNet, 2009
-VictoriaPresentIntroduced Not invasive PlantNet, 2009
-Western AustraliaPresentIntroduced Not invasive PlantNet, 2009
Cook IslandsPresentHäfliger and Scholz, 1980
FijiPresentHäfliger and Scholz, 1980; Häfliger and Scholz, 1980; PIER, 2009
French PolynesiaPresentHäfliger and Scholz, 1980; PIER, 2009
GuamPresentHäfliger and Scholz, 1980
Johnston IslandPresentHäfliger and Scholz, 1980
KiribatiPresentHäfliger and Scholz, 1980; PIER, 2009
Marshall IslandsPresentPIER, 2009Ailinglaplap, Jaluit, Ailuk, Arno and Likiep Islands (PIER, 2009)
Micronesia, Federated states ofPresent Invasive Häfliger and Scholz, 1980; PIER, 2009; USDA-ARS, 2009Chuuk, Kposrae, Pohnpei and Yap Islands (PIER, 2009)
NauruPresentHäfliger and Scholz, 1980
New ZealandLocalisedIntroduced Invasive Häfliger and Scholz, 1980; USDA-ARS, 2009
Northern Mariana IslandsPIER, 2009Anatahan, Pagan, Saipan and Tinian Islands (PIER, 2009)
PalauPresentPIER, 2009Angaur, Tobi, Babeldaob, Koror, Ngerkebesang and Peleliu Islands (PIER, 2009)
US Minor Outlying IslandsPresentHäfliger and Scholz, 1980
Wake IslandPresentPIER, 2009

History of Introduction and Spread

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There are few clear records of the introduction of P. distichum to new territories. There is a firm date for France of 1802 (Le Floc’h, 1991). In Portugal it was first reported in 1887 (Aguiar et al., 2005). In Java, Indonesia, Everaats (1981) indicates that it was relatively newly known at that time. Likewise in Iran in 2000 (Hamzeh'ee, 2000). In Spain it has only recently moved into the province of Léon (Egido Mazuelas et al., 2007). In Croatia it is also infesting new areas (Milovoic, 2001).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
France 1802 Forage (pathway cause) Yes Floc'h (1991); Floc'h E le (1991); Le (1991)
Java 1970s Yes Everaarts (1981)
Portugal 1887 Yes Aguiar et al. (2005)

Risk of Introduction

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There are highly significant risks of further spread of P. distichum into regions not already infested – especially northern sub-sahelian Africa, which has so far escaped invasion. The conditions, especially in West Africa are likely to be very favourable and the risks to the important rice growing areas there and in East Africa are high. Although spreading mainly vegatatively once established, P. distichum does produce abundant fertile seed which could be transferred as a contaminant of other crop seeds, or in hay or forage. As P. distichum has shown useful characteristics as an aid to soil conservation or as a livestock feed, there could be misguided deliberate introduction.

Habitat

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According to Häfliger and Scholz (1980)P. distichum is found in wasteland areas, rotation crops, perennial crops and aquatic habitats, but not too frequently on grassland.

P. distichum is a non-submerged aquatic plant commonly occurring in streams and alluvial flatlands in the tropics and subtropics, and throughout the world. It populates still or moving water to a depth of one metre or more; it may also be a problem in merely irrigated conditions. It is a C4 plant (Mesleard et al., 1993) adapted to semi-aquatic environments (Datta et al., 1979) and is not cold tolerant.

P. distichum has the capacity to bind soil in streams which are subject to erosion in the tropics, and provides grazing in flat coastal regions and stream beds. It is a valuable pasture grass on alluvial flats (Chase, 1929; Bor, 1960). Attempts have been made to select strains with high palatability and limited regenerative capacity (Ikeda et al., 1988). P. distichum has a capacity for explosive growth which restricts its use for binding soil in intensively cultivated areas (Li, 1981). The weed spreads readily by seed and vegetative reproduction, by its numerous stolons and rhizomes, to become a persistent weed (Smiley, 1922; Wilkinson and Jacques, 1979; Huang et al., 1987). The percentage of hemicryptophytes in the ruderal (riverbank) environments of North Africa is increasing and becoming a significant problem (Le-Floc'h, 1991).

P. distichum occurs in ditches and wet, but rarely brackish, areas as far north as Washington, USA, and as far south as Argentina and Chile; also on temperate coasts of the Eastern Hemisphere. Most references to this species occurring in saline coastal areas are attributable to confusion with P. vaginatum, but Leithead et al. (1971) describe both species and comment that P. distichum ‘grows primarily on fresh water marshes and occasionally on brackish marshes. Tolerates moderate salinity and standing water’. It is a serious weed in areas where irrigated crops such as rice or cotton are the main crops in rotation. Thus, it is an important weed in the natural floodplains and wetlands in India, and in the paddy fields of Japan, China and the Philippines. However, it can proliferate in drier conditions provided abundant water is present for at least part of the growth cycle. It can infest orchards, asparagus plantings, or vineyards which are irrigated. It is found in highland pastures in Thailand in years of heavy rainfall, but is considered as a tropical lowland pest in this region (Falvey and Hengmichai, 1978; Datta et al., 1979). The plant is predominantly lowland, rarely found at altitudes greater than 300 m (Robbins et al., 1951; Townsend et al., 1968). In Europe it has come to dominate riparian and wet alluvial plains in Portugal (Bernez et al., 2005) and in Greece (Stroh, 2006).

Habitat List

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CategoryHabitatPresenceStatus
Brackish
Estuaries Secondary/tolerated habitat Natural
Freshwater
Irrigation channels Principal habitat Harmful (pest or invasive)
Irrigation channels Principal habitat Natural
Lakes Secondary/tolerated habitat Natural
Ponds Secondary/tolerated habitat Harmful (pest or invasive)
Ponds Secondary/tolerated habitat Natural
Reservoirs Secondary/tolerated habitat Natural
Rivers / streams Secondary/tolerated habitat Harmful (pest or invasive)
Rivers / streams Secondary/tolerated habitat Natural
Littoral
Coastal areas Secondary/tolerated habitat Natural
Terrestrial-managed
Cultivated / agricultural land Principal habitat Harmful (pest or invasive)
Disturbed areas Secondary/tolerated habitat Natural
Managed forests, plantations and orchards Secondary/tolerated habitat Harmful (pest or invasive)
Managed grasslands (grazing systems) Secondary/tolerated habitat Productive/non-natural
Rail / roadsides Secondary/tolerated habitat Natural
Urban / peri-urban areas Secondary/tolerated habitat Natural
Terrestrial-natural/semi-natural
Natural grasslands Secondary/tolerated habitat Natural
Riverbanks Secondary/tolerated habitat Natural
Wetlands Principal habitat Natural

Host Plants and Other Plants Affected

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Growth Stages

Top of page Flowering stage, Fruiting stage, Pre-emergence, Seedling stage, Vegetative growing stage

Biology and Ecology

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Genetics

Hexaploid (2n=60) and tetraploid (2n=40) populations are common (Kadono, 1985; Shibayama, 1988; Echarte et al., 1992), and Echarte et al. (1992) have documented the existence of pentaploids (2n=50) and several hyperpentaploids (2n=52, 54, 57 and 58) in Argentina. Bor (1960) also records 2n=48.

P. distichum exhibits considerable variability, thought to result from both the heteroploidy shown by the species, and the existence of genetic differences among individuals of the same cytotype (Echarte et al., 1992).

Reproductive Biology

P. distichum has a high capacity for asexual reproduction. Under suitable conditions, almost every node of the aerial shoots (stolons) and rhizomes are capable of rooting and producing new plants. Under optimum conditions, P. distichum can elongate at a rate of 15-20 cm per week (Noda and Obayashi, 1971). Okuma et al. (1983) reported that within 2-3 years of planting overwintering stems of P. distichum on the bank of a stream, the entire surface of the stream was covered with a floating mat of the weed.

The weed also produces many seeds. Seed production by P. distichum has been estimated at 100 seeds per panicle and was about 100,000/m² in a river studied by Okuma and Chikura (1984). About 5-10% of the seeds produced were viable and the maximum, minimum and optimum temperatures for seed germination were 40, 20 and 30°C, respectively. Even at the optimum incubation temperature, 60% of viable seeds remained dormant. Exposure of seeds to 16-h daylength at optimum temperatures of 28-35°C increased germination from 14 to 40% (Huang and Hsiao, 1987). Pre-chilling seeds at 6°C and a 2-h heat treatment at 40°C had no effect on germination, and treatment with gibberellic acid (GA3) had little effect. Treatment of the seeds with oxidants including sodium hypochlorite, hydrogen peroxide and concentrated sulphuric acid indicated that seed dormancy was imposed mainly by seed coverings, including the hull and seed coat membranes. Seeds play an important role in the winter survival of P. distichum in regions where temperatures approach the critical limit for the species (Shibayama, 1988). 

In Argentina, Echarte et al. (1992) confirmed that P. distichum is an out-crossing species, though in China Ma GuoHua et al. (2003) concluded that the tetraploid P. distichum could be considered to be a facultative apomict reproducing by apospory. Quarín and Burson (1991) likewise concluded that it reproduces by aposporous apomixis and Srivastava and Purnima (1990) refer to it as an obligate asporous apomict with only aposporous embryo sacs. Seeds mature 20-30 days after flower emergence (Manuel et al., 1979). 

Physiology and Phenology

Single node segments of P. distichum sprout faster from stems than from rhizomes, and the rooting of shoots on contact with the soil stimulates new shoot production (Manuel and Mercado, 1977). Stolons root readily at the nodes, and elongation rates of up to 15-20 cm per week have been observed at high temperatures (Noda and Obayashi, 1971). On average, the maximum, optimum and minimum temperatures for sprouting and rooting of P. distichum segments are 40, 30-35 and 10°C, respectively (Huang et al., 1987). Shoots of P. distichum collected in different seasons differed in their sprouting and rooting responses at different incubation temperatures, providing a possible survival mechanism for the species. The rate of sprouting, rooting and the early growth of both single-node shoot and rhizome segments increased with incubation temperature up to 30°C, and then declined at 40°C. Growth was poor at 10°C. Single-node stem segments sprouted faster in a 16-h light regime than in darkness. Rooting was optimum in the dark at low temperatures; and in the light at high temperatures. Stems of P. distichum are kept in reserve by apical and bud dominance, to replace those destroyed under adverse conditions; a further survival mechanism for this species. Water, nutrient and light levels which favour photosynthesis and reduced transpiration are favourable to axillary bud growth (Liu et al., 1991).

P. distichum flourishes at high temperatures and is tolerant of high moisture conditions (Mesléard et al., 1993), as is characteristic of many C4 species.

Environmental Requirements

P. distichum is quite sensitive to shade. Plants kept at short daylengths (8 h) have smaller leaves and internodes than those kept at longer daylengths which tend to prolong maturation; plants in all treatments flower in 14-16 days. Deep (10-15 cm) water levels delay flowering. 

Response to climatic conditions in New Zealand has been studied by Campbell et al. (1999). As a C4 plant it showed less tolerance to cold conditions than C3 species.

P. distichum flourishes in wet soils but these are generally aerobic rather than anaerobic (Green and Brock, 1994). Tolerance of salinity is somewhat uncertain. There are many references to the occurrence of the weed in coastal areas, but possible confusion with the salt-tolerant P. vaginatum makes many of these references unreliable. Leithead et al. (1971) do however describe both species and comment that P. distichum ‘grows primarily on fresh water marshes and occasionally on brackish marshes. Tolerates moderate salinity and standing water’. 

 

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Tolerated > 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 Tolerated < 60mm precipitation driest month (in summer) and < (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
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Tolerated Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 5 25
Mean maximum temperature of hottest month (ºC) 35
Mean minimum temperature of coldest month (ºC) -3

Rainfall Regime

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Summer
Uniform

Soil Tolerances

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Soil drainage

  • free
  • impeded
  • seasonally waterlogged

Soil reaction

  • acid
  • alkaline
  • neutral

Soil texture

  • heavy
  • light
  • medium

Special soil tolerances

  • saline

Notes on Natural Enemies

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Relatively few natural enemies have been recorded for P. distichum and none have been proposed as potential biocontrol agents. The most significant natural enemy is the ergot fungus Claviceps paspali, infection with  which can lead to serious toxicity to livestock.

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

Natural dispersal must inevitably occur by movement of seeds in water.

Vector Transmission (Biotic)

Local movement may occur on or in livestock but is not documented.

Accidental Introduction

Accidental introduction is likely to occur in hay or as seed contamination of other crop seeds but is not documented.

Intentional Introduction

Most of the spread around the world has presumably occurred as a result of deliberate introduction of the species as a pasture grass, but there are few available records.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Animal production Yes Yes
Crop production Yes Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Plants or parts of plants Yes Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bulbs/Tubers/Corms/Rhizomes
True seeds (inc. grain)

Impact Summary

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CategoryImpact
Cultural/amenity Positive and negative
Economic/livelihood Negative
Environment (generally) Positive and negative

Economic Impact

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Economic impact occurs primarily in rice, where the weed can become dominant, causing loss of crop yield and/or increased costs of control, whether manually or by herbicides. As listed by Holm et al. (1979) it occurs as a ‘serious’ weed in Australia, Chile, Guyana, Iraq, Israel, Pakistan, Portugal, the former USSR, and Spain and as a ‘principal’ weed in Colombia, France, Iran, New Zealand and Swaziland. It is especially serious in Portugal, where surveys have shown the weed to be present in over 80% of rice fields (Vasconcelos et al., 1998). It is recorded as a major weed in Phaseolus vulgaris in Egypt (Hussein, 2003); in wheat in Bangladesh (Alam et al., 1997); and in asparagus in Chile and New Zealand.

P. distichum is a serious weed in irrigated crops, and irrigation ditches are particularly subject to invasion. Seasonal flooding in some locations, and enriched nitrogen along riverbeds favours its spread. Herbicides must be used with caution in paddy fields and irrigated areas. The weed is resistant to many herbicides, and its potential for vegetative reproduction makes it difficult to control even with intensive cultivation. No natural enemies suitable for biological control have yet been identified. The most effective control methods, involving intensive ground preparation and appropriate herbicide applications, are expensive. Low-till or no-till agricultural practices, which reduce labour costs, are not practical as crop losses can be as high as 75-80% in the presence of P. distichum. The major economic impact of P. distichum is the input of labour required for adequate control. 

Although P. distichum is often grown and/or exploited as feed for livestock, there are reports from South Africa, France, Albania and Japan of poisoning resulting from infection of the seedheads by Claviceps paspali, causing ergotism.

Environmental Impact

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In India, Keoladeo National Park has been seriously invaded by P. distichum, to the detriment of the native vegetation (Middleton, 1998). Invasion of natural wetlands by P. distichum is also serious in Portugal, where it is considered a threat for particular river flora and consequently for the integrity of the river system (Bernez et al., 2005), and also in Greece (Stroh, 2006). 

Dominance of P. distichum in natural wetlands or waterways can also result in reduced fish populations (Kumar and Mittal, 1993).

Social Impact

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Dense growth of P. distichum can greatly increase mosquito populations compared with more open vegetation, hence decreasing amenity and increasing risk of disease (Lawler et al., 2007). 

P. distichum is noted as a weed of ornamental turf in China (Zhou Qiang et al., 2004), and in France (Bourgoin and Guérin, 1999).

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
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Gregarious
  • 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 hydrology
  • Modification of nutrient regime
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts agriculture
  • Negatively impacts animal health
  • Negatively impacts livelihoods
  • Reduced amenity values
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Filtration
  • Poisoning
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field
  • Difficult/costly to control

Uses

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Economic Value

P. distichum is not always considered a weed. It is introduced into rice paddy fields as a forage crop by some Japanese farmers (Ikeda et al., 1983). Bor (1960) comments that P. distichum is considered in many parts of the world to be a valuable pasture grass and has been used extensively in areas subject to flooding. ‘P. paspalodes’ was cultivated in the Bordeaux region of France as early as 1802 for use as pasture. While P. distichum survives root flooding and submergence without difficulty, protracted deep flooding can reduce photosynthesis in existing leaves and can be detrimental to growth (Hsiao and Huang, 1989b).

In Japan, ‘roll bale silage’ from P. distichum grown in paddy fields with a large amount of nitrogen fertilizer show a potential in terms of feeding value (Asano et al., 2007). 

In Australia, Rosicky et al. (2006) comment that P. distichum is favoured as a highly palatable forage for stock. 

Environmental Services

In Australia, Rosicky et al. (2006) comment that P. distichum is highly favoured for its environmental benefits in providing thick mulch cover, and assisting in the revegetation of bare ground. Similarly it is favoured as a ground cover in Florida, USA (Jenkins et al., 2004), while in China, it is tolerant of heavy metals and suitable for revegetation of ground contaminated by lead, zinc and copper (Shu et al., 2002).

P. distichum has been used for purification of water, as in Korea, where it was considered valuable for extracting nitrogen and phosphorus from eutrophic water (Lee et al., 2004) and in Australia for removal of phosphorus (Wen et al., 2002).

Uses List

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Animal feed, fodder, forage

  • Fodder/animal feed
  • Forage

Environmental

  • Amenity
  • Erosion control or dune stabilization
  • Host of pest
  • Land reclamation
  • Revegetation
  • Soil conservation

Similarities to Other Species/Conditions

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P. distichum is sometimes confused with Paspalum vaginatum but is distinguishable by the slightly more turgid spikelets, a glume and sterile lemma which are not papery, a more pronounced midrib, and a glume which is at least obscurely pubescent (Chase, 1929). P. distichum is also distinguished by the unusually variable size of the lower glume. P. vaginatum is almost always associated with saline or brackish conditions, especially near coasts, but also in saline soils inland (e.g. Arabia), whereas P. distichum does not generally occur under these conditions.

Many other species of Paspalum can occur as weeds. One of the most widespread of these is a complex of closely-related taxa which include P. commersonii and P. orbiculare but which are usually known as P. scrobiculatum (qv). This is widespread in Asia and Africa. It is not likely to be confused with P. distichum as it has a tufted habit whether spreading or erect, annual or perennial, but without runners or rhizomes. The inflorescence also differs in having spikelets almost round in outline.

 

Prevention and Control

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SPS Measures

P. distichum is listed as an invasive alien plant in Europe (EPPO, 2009), requiring member countries to minimise the risks of introduction, and of spread within their countries.

Control

Cultural control and sanitary measures

Methods used to control P. distichum differ depending on whether the crops in which the weed is growing are produced under flooding or irrigation. Farmers in the Philippines consider P. distichum and Echinochloa spp. to be the most difficult weeds to control (Fajardo and Moody, 1990). Herbicides are expensive and are used sparingly, but acceptable levels of control can be achieved due to the high seeding rates, high fertilizer rates, and the use of integrated cultural practices. Flooding and deep burial inhibit the growth of P. distichum as oxygen is required for sprouting from stem cuttings centering around the joints. No sprouting was observed from severed creeping stems of P. distichum which were buried in puddled soil or submerged in water. Most sprouting occurs from overwintering stems lying above the water; almost none occurs from submerged overwintering stems. Flooding reduces the competitive advantage of P. distichum with respect to rice (Manuel and Mercado, 1977; Manuel et al., 1979). P. distichum is sensitive to shade (Manuel and Mercado, 1977; Manuel et al., 1979; Ito et al., 1987) and higher crop seeding rates, closer distances at transplanting, and the use of taller, more competitive cultivars are, therefore, potential control measures.

The effects of no-tillage systems on weed development in a variety of crops was investigated by Gong and Li (1991). The underground vegetative portions of P. distichum tend to be shallow, with aggressive shoot development. No-tillage systems are not feasible in areas heavily infested with P. distichum and minimum tillage is, at the very least, necessary for satisfactory yields of semi-dwarf rice (Datta et al., 1979; Bernasor and Datta, 1988).

In some regions nitrogen levels are increased by encouraging Azolla growth and P. distichum has emerged as a major weed under such regimes (Janiya and Moody, 1987; Krock et al., 1988).

In the Keoladeo National Park, India, P. distichum was apparently suppressed by the grazing of water buffalo. After removal of these animals, the weed expanded but was later brought under some degree of control by the grazing of wild pigs, geese and deer (Middleton, 1998). 

Physical/mechanical control

Thorough preparation of the land during the 30 day period prior to transplanting, augmented with handweeding, provides effective control of P. distichum in the Philippines (Rahman, 1992). Increasing tillage frequency from one to three harrowings substantially reduces stands of the weed (Bernasor and Datta, 1988). In fields infested by seeds, overwintering rhizomes and stems, cultivation or diskings which normally eradicate small seedlings, only help to propagate the weed by cutting the stems and rhizomes into small pieces. Ploughing often promotes the dominance of plants with bulbs, stolons or rhizomes (Hsiao and Huang, 1989a). However, deep ploughing is an effective method of control (Noda and Obayashi, 1971), as the depth of emergence is less (3.25 cm on average) than that of other perennial plants of the Cyperaceae found in similar environments (Okuma et al., 1983). Moisture conditions are important for the success of mechanical disturbance: under wet conditions control may depend on deep burial, whereas under seasonally dry conditions, shallow tillage may suffice.

Samantha et al. (1995) emphasise the need for two hand weedings to control P. distichum in rice in Bangladesh. 

Solarisation has been used with some success in Spain (Dalmau et al., 1993). 

Chemical control

P. distichum tolerates most of the popular pre- and post-emergent herbicides. Resistance to post-emergent herbicides is attributed to anatomical features such as: thickening of the cell membranes of the epidermis, cortex and central portion; poor development of vascular bundles, and no air capacity in the central portion; and the accumulation of starch granules in the cortex and central regions (Noda and Obayasi, 1971). The accumulation of starch is partly responsible for the regenerative ability of isolated segments of the stem or rhizome. Glyphosate is the most widely used herbicide for the control of P. distichum (Okuma and Chikura, 1985; Manuel et al., 1979). Thiobencarb (Shad, 1986; Singh, 1992), molinate (Srinivasan and Subbian, 1991), haloxyfop or quizalofop (Rahman and Sanders, 1991), pretilachlor (Llorente and Evangelista, 1990), butachlor (Pradhan and Chettri, 1987; Bajwa et al., 1985), fluazifop (Okuma and Chikura, 1985), sethoxydim (Parker, 1982), paraquat (Manuel et al., 1979) and fluroxypyr plus triclopyr (Schultz, 2005) can also provide effective control, depending on the crop. Metolachlor, oxyfluorfen, prodiamine, and pendimethalin were most effective in leeks (Gilreath et al., 2008). Nicosulfuron is moderately effective in maize (James and Rahman, 1997). 

Newer herbicides for rice include fenoxaprop (Reddy et al., 2000), bispyribac-sodium (Wang Qiang et al., 2000; He JinHao and Ma ZhaoJiang, 2000), pyanchor (pyribenzoxim) (Zhou XiaoJun et al., 2000), bensulfuron methyl and prosulfuron-ethyl (Sumiyoshi, 2000). 

Glufosinate is also effective in conjunction with transgenic glufosinate-tolerant rice varieties (Yu LiuQing et al., 2005). 

The use of pre-planting herbicides can reduce the need for extensive land preparation. The 30-day preparation period required to effectively control P. distichum can be reduced to 15 days when combined with glyphosate treatment. The combined use of low-tillage practices and herbicide treatment is increasing in popularity in areas where labour costs are high.

Biological Control

There are currently no biological control methods for P. distichum. The weed is an alternative host for many of the fungi, viruses, bacteria and nematodes that afflict crops in which the weed occurs.

References

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GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.

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12/08/2009 Updated by:

Chris Parker, Consultant, UK

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