Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Urochloa mutica
(para grass)

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Datasheet

Urochloa mutica (para grass)

Summary

  • Last modified
  • 19 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Host Plant
  • Preferred Scientific Name
  • Urochloa mutica
  • Preferred Common Name
  • para grass
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Monocotyledonae
  • Summary of Invasiveness
  • U. mutica is a perennial grass widely naturalized in tropical and subtropical regions of the world where it was introduced as a fodder grass. Now, this species is considered one of the world’s worst weeds in the United States, Australia,...

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Pictures

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PictureTitleCaptionCopyright
Growth habit.
TitleHabit
CaptionGrowth habit.
Copyright©Sheldon Navie
Growth habit.
HabitGrowth habit.©Sheldon Navie
Growth habit at water's edge.
TitleHabit
CaptionGrowth habit at water's edge.
Copyright©Sheldon Navie
Growth habit at water's edge.
HabitGrowth habit at water's edge.©Sheldon Navie
Seed head.
TitleSeed head
CaptionSeed head.
Copyright©Sheldon Navie
Seed head.
Seed headSeed head.©Sheldon Navie
Leaf and stem, note profuse hairs.
TitleStem
CaptionLeaf and stem, note profuse hairs.
Copyright©Sheldon Navie
Leaf and stem, note profuse hairs.
StemLeaf and stem, note profuse hairs.©Sheldon Navie
Leaf and stem.
TitleStem
CaptionLeaf and stem.
Copyright©Sheldon Navie
Leaf and stem.
StemLeaf and stem.©Sheldon Navie

Identity

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Preferred Scientific Name

  • Urochloa mutica (Forssk.) T.Q.Nguyen

Preferred Common Name

  • para grass

Other Scientific Names

  • Brachiaria mutica (Forssk.) Stapf
  • Brachiaria numidiana (Lam.) Henrard
  • Brachiaria purpurascens (Raddi) Henr.
  • Panicum amphibium Steud.
  • Panicum barbinode Trin.
  • Panicum equinum Salzm. ex Steud.
  • Panicum limnaeum Steud.
  • Panicum muticum Forssk.
  • Panicum numidianum Lam.
  • Panicum pictigluma Steud.
  • Panicum punctulatum Arn. ex Steud.
  • Panicum purpurascens Raddi

International Common Names

  • English: buffalo grass; California grass; Carib grass; Mauritius grass; para grass; Scotch grass; tall panicum; water grass
  • Spanish: gramalote; hierba de para; hierba para; malojilla; papare; parana; pasto para; Zacate para
  • French: herbe borer; herbe de Para
  • Portuguese: capim-de-Angola

Local Common Names

  • Brazil: angolinha; bengo; braquiária; capim-angola; capim-bengo; capim-branco; capim-das-ilhas; capim-de-cavalo; capim-de-corte; capim-de-lastro; capim-de-pará; capim-de-planta; capim-do-Pará; capim-fino; capim-planta; egipto; erva-do-pará; vapim-fino
  • Germany: Paragras
  • Indonesia: rumput melela
  • Mexico: zacate para
  • Puerto Rico: malojillo; yerba pará

EPPO code

  • PANPU (Brachiaria mutica)

Summary of Invasiveness

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U. mutica is a perennial grass widely naturalized in tropical and subtropical regions of the world where it was introduced as a fodder grass. Now, this species is considered one of the world’s worst weeds in the United States, Australia, Mexico, and Central America (Holm et al., 1977; Chacon and Saborío, 2012; Hannan-Jones and Csurhes, 2012; Randall, 2012; USDA-NRCS, 2014). U. mutica has been intentionally introduced as a “ponded pasture grass” because its capability to establish on poorly drained (swampy or seasonally waterlogged) soils as well on free-draining soils in high rainfall environments (Cook et al., 2005). This grass species competes aggressively with other plants, with fast growth, high productivity, and allelopathic abilities that allow it to form dense stands (Holm et al., 1977; Langeland et al., 2008).

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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The grass species U. mutica was first described as Panicum muticum by Forsskal in 1775. It was moved to the genus Brachiaria in 1919 by Otto Stapf. The current name U. mutica was published in 1966. The genus Urochloa is paleotropical and includes 12 species native mainly to the African savannas (Torres-Gonzalez and Morton, 2005).

The weaknesses of the characters used to separate Brachiaria from Urochloa (i.e., spikelet orientation and presence or absence of an upper floret) have been discussed by several authors including Webster (1987, 1988) and Morrone and Zuloaga (1992, 1993). Consequently, floristic studies conducted in Australia (Webster, 1987), North America (Webster, 1988; Zuloaga and Morrone, 2003), South America, Mexico and Central America (Morrone and Zuloaga, 1992, 1993) have circumscribed species of Brachiaria into Urochloa. On the other hand, Sharp and Simon (2002) maintain the name Brachiaria for all species that occur in Australia and the annual species of Brachiaria are now included in the new genus Moorochloa (Veldkamp, 2004). The taxonomic positions of these genera still remain unclear.

Description

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U. mutica is a perennial; stoloniferous grass. Culms to 5 m long, long-decumbent and rooting at the lower nodes, vertical portion 90-200 (300) cm; nodes villous. Lower sheaths with papillose-based hairs, margins ciliate; collars pubescent; ligules 1-1.5 mm; blades 7.5-35 cm long, 4-20 mm wide, glabrous or sparsely pilose on both surfaces, margins scabrous. Panicles 10-25 cm long, 5-10 cm wide, pyramidal, with 10-30 spikelike branches in more than 2 ranks; primary branches 2.5-8 cm long, 0.4-0.9 mm wide, ascending to divergent, axes flat, glabrous or with a few papillose-based hairs, secondary branches present or absent; pedicels shorter than the spikelets, scabrous, sometimes with hairs. Spikelets 2.6-3.5 mm long, 1-1.4 mm wide, mostly in pairs, in 2-4 rows, appressed to the branches, purplish to green. Glumes scarcely separate, lower glumes 0.6-1.1 mm, 1/5-1/3 as long as the spikelets, glabrous, 0-1(3)-veined; upper glumes 2.6-3.5 mm, glabrous, 5-(7)-veined, without cross venation; lower florets staminate; lower lemmas 2.6-3.3 mm, glabrous, 5-veined, without cross venation; upper lemmas 2.3-2.8 mm long, 1-1.3 mm wide, apices rounded, mucronate; anthers 1-1.5 mm. Caryopses 1.8-2 mm long (Barkworth et al., 2003).

Plant Type

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Grass / sedge
Herbaceous
Perennial
Seed propagated
Vegetatively propagated

Distribution

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U. mutica is native to tropical areas of western and northern Africa (Parsons 1972) including areas from the Sahara to Angola, northern Africa to Syria, and the Southwestern Arabian Peninsula (Clayton et al., 2014). It is now also widely distributed in Australia, New Zealand, Asia, the southern United States, Mexico, Central America, South America and the West Indies (for details see Distribution Table).

Distribution Table

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

Last updated: 14 Dec 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AlgeriaPresentNative
AngolaPresentNative
BeninPresentNative
Burkina FasoPresentNative
CameroonPresentNative
ChadPresentNative
Congo, Democratic Republic of thePresentNative
Congo, Republic of thePresentNative
Côte d'IvoirePresentNative
EgyptPresentNative
GabonPresentNative
GambiaPresentNative
GhanaPresentNative
GuineaPresentNative
LiberiaPresentNative
MadagascarPresentIntroduced
MaliPresentNative
MauritaniaPresentNative
MauritiusPresentIntroducedInvasive
-RodriguesPresentIntroducedInvasive
MoroccoPresentNative
NigerPresentNative
NigeriaPresentNative
RéunionPresentIntroducedInvasive
SenegalPresentNative
SeychellesPresentIntroduced
Sierra LeonePresentNative
SomaliaPresentIntroduced
TanzaniaPresentIntroduced
TogoPresentNative
TunisiaPresentNative

Asia

BangladeshPresentIntroduced
CambodiaPresentIntroducedInvasive
ChinaPresentPresent based on regional distribution.
Hong KongPresentIntroduced
IndiaPresentPresent based on regional distribution.
-Andaman and Nicobar IslandsPresentIntroduced
-AssamPresentIntroduced
IndonesiaPresentIntroduced
-JavaPresentIntroduced
-SulawesiPresentIntroduced
-SumatraPresentIntroduced
IsraelPresentNative
JapanPresentIntroduced
LaosPresent
LebanonPresentNative
MalaysiaPresentIntroducedInvasive
MyanmarPresentIntroduced
NepalPresentIntroduced
PhilippinesPresentIntroducedInvasive
SingaporePresentIntroducedInvasive
Sri LankaPresentIntroduced
SyriaPresentNative
TaiwanPresentIntroducedInvasive
ThailandPresentIntroducedInvasive
VietnamPresentIntroducedInvasive
YemenPresentNative

Europe

PortugalPresentPresent based on regional distribution.
-AzoresPresentIntroduced
-MadeiraPresentIntroduced

North America

Antigua and BarbudaPresentIntroduced
ArubaPresentIntroduced
BahamasPresentIntroduced
BarbadosPresentIntroduced
BelizePresentIntroduced
BermudaPresentIntroduced
Cayman IslandsPresentIntroduced
Costa RicaPresentIntroducedInvasive
CubaPresentIntroducedInvasive
CuraçaoPresentIntroduced
DominicaPresentIntroduced
Dominican RepublicPresentIntroduced
El SalvadorPresentIntroduced
GrenadaPresentIntroduced
GuadeloupePresentIntroduced
GuatemalaPresentIntroduced
HaitiPresentIntroduced
HondurasPresentIntroduced
JamaicaPresentIntroduced
MartiniquePresentIntroduced
MexicoPresentIntroducedInvasive
MontserratPresentIntroduced
Netherlands AntillesPresentIntroduced
NicaraguaPresentIntroduced
PanamaPresentIntroduced
Puerto RicoPresentIntroducedInvasive
Saint Kitts and NevisPresentIntroduced
Saint LuciaPresentIntroducedVery common
Saint Vincent and the GrenadinesPresentIntroduced
Trinidad and TobagoPresentIntroduced
U.S. Virgin IslandsPresentIntroducedInvasive
United StatesPresentPresent based on regional distribution.
-AlabamaPresentIntroduced
-FloridaPresentIntroducedInvasive
-HawaiiPresentIntroducedInvasive
-MarylandPresentIntroduced
-OregonPresentIntroduced
-South CarolinaPresentIntroduced
-TexasPresentIntroduced

Oceania

American SamoaPresentIntroducedInvasive
AustraliaPresentPresent based on regional distribution.
-New South WalesPresentIntroducedInvasive
-Northern TerritoryPresentIntroducedInvasive
-QueenslandPresentIntroducedInvasive
-South AustraliaPresentIntroducedInvasive
-TasmaniaPresentIntroducedInvasive
-VictoriaPresentIntroducedInvasive
-Western AustraliaPresentIntroducedInvasive
Cook IslandsPresentIntroduced
FijiPresentIntroducedInvasive
French PolynesiaPresentIntroducedInvasive
GuamPresentIntroducedInvasive
Marshall IslandsPresentIntroduced
New CaledoniaPresentIntroducedInvasive
New ZealandPresentIntroducedInvasive
NiuePresentIntroducedInvasive
Northern Mariana IslandsPresentIntroduced
PalauPresentIntroducedInvasive
Papua New GuineaPresentIntroduced
SamoaPresentIntroducedInvasive
Solomon IslandsPresentIntroduced
TongaPresentIntroducedInvasive
U.S. Minor Outlying IslandsPresentIntroduced
VanuatuPresentIntroduced
Wallis and FutunaPresentIntroduced

South America

ArgentinaPresentIntroduced
BoliviaPresentIntroduced
BrazilPresentIntroducedDistrito Federal
-GoiasPresentIntroduced
-Mato GrossoPresentIntroduced
-Minas GeraisPresentIntroduced
-ParanaPresentIntroducedInvasive
-Sao PauloPresentIntroducedInvasive
ColombiaPresentIntroducedInvasive
EcuadorPresentIntroducedInvasiveInvasive on Galapagos Islands
French GuianaPresentIntroducedNaturalizedNaturalized
GuyanaPresentIntroducedNaturalizedNaturalized
ParaguayPresentIntroduced
PeruPresentIntroduced
SurinamePresentIntroducedNaturalizedNaturalized
VenezuelaPresentIntroduced

History of Introduction and Spread

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U. mutica was introduced into the Americas via Brazil in the early days of trading (Parsons, 1972; Smith, 1979). Consequently, in the 1800s, there was confusion about its origin, with suggestions that it was native to South America, and in 1823 it was described from Brazilian specimens as Panicum purpurascens, and as Panicum barbinode in 1829 (Stone, 1970). It was introduced into Florida in the 1870s and recommended as a forage plant by the Florida Agricultural Experiment Station in 1910 (Langeland et al., 2008; Chaudhari et al., 2012). In Australia, it was introduced into Queensland around 1880 to reduce soil erosion along the banks of waterways (Hannan-Jones and Csurhes, 2012). In the West Indies, it was first collected in 1883 in Puerto Rico (US Herbarium collection). By 1977, Holm and collaborators listed this species as a serious weed in Australia, Fiji and Thailand, a weed in Sri Lanka, Colombia, Hawaii, Jamaica, Malaysia, Peru, the Philippines, Puerto Rico, and Trinidad, and as a common weed in Borneo and Mauritius (Holm et al., 1977).

Risk of Introduction

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The risk of introduction of U. mutica is very high. It has been intentionally introduced repeatedly in tropical and subtropical regions to be used as a fodder, forage and silage crop (Cook et al., 2005). It has escaped from cultivation and rapidly naturalized into natural areas where it colonizes forming dense stands and displacing native vegetation (Holm et al., 1977; Langeland et al., 2008; Hannan-Jones and Csurhes, 2012). When growing under suitable environmental conditions (i.e., moist soils), U. mutica spreads rapidly (up to 5 metres in a year) through its long stolons and possibly through water-borne seed (Cook et al., 2005).

Habitat

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U. mutica can be found growing in poorly drained, swampy or seasonally waterlogged areas, along creeks, rivers, floodplains, wetlands and drainage channels, around lakes and dams, in roadside ditches and in other damp habitats (Holm et al., 1977; Cook et al., 2005; Hannan-Jones and Csurhes, 2012). In Florida, the species has been reported growing in coastal berms, hardwood hammocks, mesic and wet flatwoods, bottomland forests, floodplain forests, stream and spring shores, and ruderal communities. U. mutica also grows as a weed of summer crops, plantation crops such as sugarcane, sown pastures, rice plantations and orchards (Holm et al., 1977; Queensland Department of Primary Industries and Fisheries, 2011).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial ManagedCultivated / agricultural land Present, no further details Harmful (pest or invasive)
Terrestrial ManagedCultivated / agricultural land Present, no further details Natural
Terrestrial ManagedCultivated / agricultural land Present, no further details Productive/non-natural
Terrestrial ManagedManaged forests, plantations and orchards Present, no further details Harmful (pest or invasive)
Terrestrial ManagedManaged forests, plantations and orchards Present, no further details Natural
Terrestrial ManagedManaged forests, plantations and orchards Present, no further details Productive/non-natural
Terrestrial ManagedManaged grasslands (grazing systems) Present, no further details Natural
Terrestrial ManagedManaged grasslands (grazing systems) Present, no further details Productive/non-natural
Terrestrial ManagedDisturbed areas Present, no further details Harmful (pest or invasive)
Terrestrial ManagedDisturbed areas Present, no further details Natural
Terrestrial ManagedDisturbed areas Present, no further details Productive/non-natural
Terrestrial ManagedRail / roadsides Present, no further details Harmful (pest or invasive)
Terrestrial ManagedRail / roadsides Present, no further details Natural
Terrestrial ManagedRail / roadsides Present, no further details Productive/non-natural
Terrestrial Natural / Semi-naturalNatural grasslands Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural grasslands Present, no further details Natural
Terrestrial Natural / Semi-naturalNatural grasslands Present, no further details Productive/non-natural
Terrestrial Natural / Semi-naturalRiverbanks Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalRiverbanks Present, no further details Natural
Terrestrial Natural / Semi-naturalRiverbanks Present, no further details Productive/non-natural
Terrestrial Natural / Semi-naturalWetlands Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalWetlands Present, no further details Natural
Terrestrial Natural / Semi-naturalWetlands Present, no further details Productive/non-natural
FreshwaterIrrigation channels Present, no further details Harmful (pest or invasive)
FreshwaterIrrigation channels Present, no further details Natural
FreshwaterIrrigation channels Present, no further details Productive/non-natural
FreshwaterLakes Present, no further details Harmful (pest or invasive)
FreshwaterLakes Present, no further details Natural
FreshwaterLakes Present, no further details Productive/non-natural
FreshwaterRivers / streams Present, no further details Harmful (pest or invasive)
FreshwaterRivers / streams Present, no further details Natural
FreshwaterRivers / streams Present, no further details Productive/non-natural
FreshwaterPonds Present, no further details Harmful (pest or invasive)
FreshwaterPonds Present, no further details Natural
FreshwaterPonds Present, no further details Productive/non-natural

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Elaeis guineensis (African oil palm)ArecaceaeOther
    Hevea brasiliensis (rubber)EuphorbiaceaeOther
      Oryza sativa (rice)PoaceaeOther

        Biology and Ecology

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        Genetics

        The chromosome number reported for U. mutica is 2n = 36 (Barkworth et al., 2003). 

        Reproductive Biology

        U. mutica has been reported as a short-day species that flowers most prolifically in humid environments at latitudes of 10–20º (Cook et al., 2005). Pollination is apparently wind-aided and little or no flowering is reported at subtropical latitudes. In Florida, it flowers from September through December (Langeland et al., 2008) and in northern Australia it flowers in late April/early May and seeds in May (Cook et al., 2005). 

        Longevity

        U. mutica is a long-lived perennial species (Barkworth et al., 2003). 

        Environmental Requirements

        U. mutica is native to floodplains in sub-Saharan tropical Africa; thus this species prefers to grow in flat, poorly drained, seasonal floodplains or high rainfall environments in tropical and subtropical regions of the world (mostly in areas with full sunlight; Cook et al., 2005; Queensland Department of Primary Industries and Fisheries, 2011). It is also adapted to grow in wetlands, ponds, and along rivers, creeks and lakes from sea level to about 1500 metres in elevation. U. mutica is well adapted to a wide range of soil types (from sandy to clay soils), and tolerates moderate salinity, low pH to 4.5 and the high levels of trace elements normally produced under water-logged conditions. It is also adapted to high temperatures (20-35°C) but growth is restricted by temperatures below 15ºC (Cook et al., 2005).

        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]))
        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])
        BS - Steppe climate Preferred > 430mm and < 860mm annual precipitation
        BW - Desert climate Tolerated < 430mm annual precipitation
        Cf - Warm temperate climate, wet all year Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
        Cs - Warm temperate climate with dry summer Tolerated 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) 18 27.5

        Rainfall

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        ParameterLower limitUpper limitDescription
        Mean annual rainfall8704100mm; lower/upper limits

        Soil Tolerances

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        Soil drainage

        • seasonally waterlogged

        Soil reaction

        • acid

        Soil texture

        • heavy
        • light
        • medium

        Special soil tolerances

        • shallow

        Natural enemies

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        Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
        Capnodium Mycoparasite Plants|Growing point not specific N
        Mocis latipes Herbivore All Stages not specific N
        Rhizoctonia Pathogen All Stages not specific N

        Notes on Natural Enemies

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        According to the Purdue University NewCROP web site (based on Duke, 1983), the following fungi have been reported on U. mutica:

        • Epicoccum andropogonis
        • Thanatephorus cucumeris
        • Gibberella pulicaris
        • Helminthosporium sp.
        • Marasmius sacchari
        • Mayriogenospora paspali
        • Myrothecium striatosporum
        • Khuskia oryzae
        • Nigrospora panici
        • Perisporium zeae
        • Pythium artorogus
        • Pythium arrhenomanes
        • Magnaporthe grisea
        • Uromyces setariae-italicae 

        This species is also attacked by the bacterium Pectobacterium carotovorum var. graminarum and the list of nematode species isolated from this grass includes:

        • Dolichodorus nigeriensis
        • Helicotylenchus pseudorobustus
        • Hemicriconemoides cocophilus
        • Scutellonema clathricaudatum
        • Tylenchorhynchus sp.
        • Xiphinema ifacolum

        Means of Movement and Dispersal

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        U. mutica spreads by seed and vegetatively by stolons. It can grow up to 5 metres in a year (Cook et al., 2005). U. mutica has water-borne seed and consequently seeds and stem fragments can be spread by floodwaters. Seeds and stem segments can also be dispersed by animals such as birds and by cattle. Long-distance dispersal occurs principally through its use as a pasture grass (Cook et al., 2005; Queensland Department of Primary Industries and Fisheries, 2011).

        Pathway Causes

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        CauseNotesLong DistanceLocalReferences
        Animal productionPasture grass Yes Yes Cook et al. (2005)
        Escape from confinement or garden escapeEscaped from cultivation Yes Yes Cook et al. (2005)
        ForagePasture grass Yes Yes Cook et al. (2005)

        Pathway Vectors

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        VectorNotesLong DistanceLocalReferences
        Debris and waste associated with human activitiesSeeds, stem fragments Yes Yes Cook et al. (2005)
        Host and vector organismsBirds spread seeds and stem fragments Yes Yes Smith (2002)
        Land vehiclesSeeds, stem fragments Yes Yes Cook et al. (2005)
        LivestockSeeds, runners and cuttings Yes Yes Cook et al. (2005)
        Machinery and equipmentSeeds, stem fragments Yes Yes Cook et al. (2005)
        MailSeeds Yes Yes Cook et al. (2005)
        Soil, sand and gravelSeeds, stem fragments Yes Yes Cook et al. (2005)
        WaterSeeds, stem fragments Yes Yes Smith (2002)

        Impact Summary

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

        Environmental Impact

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        U. mutica is a fast-growing species which grows forming very dense infestations that smother riverbanks, wetlands, and floodplain vegetation, and it also floats out over the water surface reducing areas available for waterfowl and water-birds. It is invasive in riparian habitats, wetlands, and swamps in Australia, the United States (i.e., Florida and Hawaii), Mexico, Central America, Cuba, Puerto Rico, and islands in the Pacific (Holm et al., 1977; Wagner et al., 1999; Villaseñor and Espinosa-Garcia, 2004; Langeland et al., 2005; Chacon and Saborio, 2012; Gonzalez et al., 2012; PIER, 2014; USDA-NRCS, 2014). U. mutica also invades areas of remnant vegetation away from water, especially in coastal areas and disturbed sites.

        In Australia, U. mutica is considered a serious environmental weed in wetlands in the Western Territory, Northern Territory, Queensland, and New South Wales, where it is destroying water-bird breeding habitats and replacing native vegetation along streams and in riparian zones. Here, this grass is destroying the breeding habitat of the magpie goose (Anseranas semipalmata) and reducing the ability of this bird to feed in open water. It is also one of the major environmental weeds infesting floodplains in the Northern Territory and contributing to the decline of the endangered yellow chat Epthianura crocea tunneyi (Queensland Department of Primary Industries and Fisheries, 2011; Hannan-Jones and Csurhes, 2012).

        In Florida, U. mutica invades disturbed low channels, lake shorelines, coastal berms, hardwood hammocks, mesic and wet flatwoods, bottomland forests, floodplain forests, streams, spring shores, marshes, swamps and ruderal communities where it is displacing native vegetation (Langeland et al., 2008).

        U. mutica can also change the fire regime in invaded habitats because during the dry season the aboveground portion of the grass dries out becoming a potential “fuel activator” for fires. It also has the potential to alter the water carrying capacity of streams and riparian areas invaded, causing increased flooding in infested water systems (Queensland Department of Primary Industries and Fisheries, 2011). In Brazil, U. mutica is one of several aquatic plant species that has caused significant damage to infrastructure associated with hydroelectric dams (Costa et al., 2006). 

        Threatened Species

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        Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
        Epthianura crocea tunneyi (Yellow Chat (tunneyi))National list(s)Australian Northern TerritoryCompetition - smothering; Competition - stranglingQueensland Department of Primary Industries and Fisheries (2011)
        Megalagrion nigrohamatum nigrolineatum (blackline Hawaiian damselfly)USA ESA listing as endangered speciesHawaiiEcosystem change / habitat alterationUS Fish and Wildlife Service (2014)
        Megalagrion oceanicum (oceanic Hawaiian damselfly)VU (IUCN red list: Vulnerable); USA ESA listing as endangered speciesHawaiiEcosystem change / habitat alterationUS Fish and Wildlife Service (2014)
        Megalagrion xanthomelas (orangeblack Hawaiian damselfly)VU (IUCN red list: Vulnerable); USA ESA listing as endangered speciesHawaiiEcosystem change / habitat alterationUS Fish and Wildlife Service (2014)

        Risk and Impact Factors

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        Invasiveness
        • Proved invasive outside its native range
        • Has a broad native range
        • Abundant in its native range
        • Highly adaptable to different environments
        • Is a habitat generalist
        • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
        • Pioneering in disturbed areas
        • Highly mobile locally
        • Benefits from human association (i.e. it is a human commensal)
        • Long lived
        • Fast growing
        • Has high reproductive potential
        • Reproduces asexually
        Impact outcomes
        • Altered trophic level
        • Damaged ecosystem services
        • Ecosystem change/ habitat alteration
        • Infrastructure damage
        • Modification of fire regime
        • Modification of hydrology
        • Modification of nutrient regime
        • Modification of successional patterns
        • Monoculture formation
        • Negatively impacts agriculture
        • Reduced native biodiversity
        • Soil accretion
        • Threat to/ loss of endangered species
        • Threat to/ loss of native species
        • Transportation disruption
        Impact mechanisms
        • Allelopathic
        • Competition - monopolizing resources
        • Competition - shading
        • Competition - smothering
        • Competition - strangling
        • Rapid growth
        • Rooting
        Likelihood of entry/control
        • Highly likely to be transported internationally accidentally
        • Difficult to identify/detect as a commodity contaminant

        Uses

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        U. mutica has been used as forage, fodder and pasture grass in waterlogged conditions and ponded pastures. It has also been used to control soil erosion on sloping fields and in seasonally waterlogged areas (Cook et al., 2005; PROTA, 2014).

        Uses List

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        Animal feed, fodder, forage

        • Fodder/animal feed
        • Forage

        Environmental

        • Erosion control or dune stabilization

        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.

        A combination of manual and chemical methods is recommended for the management of infestations of U. mutica. In the case of smaller infestations, plants can be cut out and all stolons must be removed. Larger infestations can be controlled by cutting the foliage and the aboveground segments of the grass. Long-term control of treated areas is recommended. The herbicide glyphosate can be applied to actively growing plants at the early head stage, but not to weeds growing over water (Queensland Department of Primary Industries and Fisheries, 2011).

        In Florida, a study evaluating the use of non-chemical control of U. mutica showed that burning, cutting, or roller-chopping should be applied in conjunction with flooding for effective management. This study shows that roller-chopping followed by flooding, and burning followed by flooding, can be options to control this grass species in areas where herbicides cannot be applied (Chaudhari et al., 2012).

        References

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        Acevedo-Rodríguez P, Strong MT, 2012. Catalogue of the Seed Plants of the West Indies. Smithsonian Contributions to Botany, 98:1192 pp. Washington DC, USA: Smithsonian Institution. http://botany.si.edu/Antilles/WestIndies/catalog.htm

        Barkworth ME, Capels KM, Long S, Piep MB, 2003. Urochloa. Flora of North America, volume 25. http://herbarium.usu.edu/webmanual/

        Chacón E, Saborío G, 2012. Red Interamericana de Información de Especies Invasoras, Costa Rica ([English title not available]). San José, Costa Rica: Asociación para la Conservación y el Estudio de la Biodiversidad. http://invasoras.acebio.org

        Chaudhari S, Sellers BA, Rockwood SV, Ferrell JA, MacDonald GE, Kenworthy KE, 2012. Nonchemical methods for paragrass (Urochloa mutica) control. Invasive Plant Science and Management, 5(1):20-26. http://wssajournals.org/loi/ipsm

        Chong KY, Tan HTW, Corlett, RT, 2009. A checklist of the total vascular plant flora of Singapore: native, naturalised and cultivated species. A checklist of the total vascular plant flora of Singapore. National University of Singapore, Singapore: Raffles Museum of Biodiversity Research, 273 pp

        Clayton WD, Govaerts R, Harman KT, Williamson H, Vorontsova M, 2014. World Checklist of Poaceae. Richmond, UK: Royal Botanic Gardens, Kew. http://apps.kew.org/wcsp/

        Cook B, Pengelly B, Brown S, Donnelly J, Eagle D, Franco A, Hanson J, Mullen B, Partridge I, Peters M, Schultze-Kraft R, 2005. Tropical Forages: an interactive selection tool. Brisbane, Australia: CSIRO, DPI&F (Qld), CIAT and ILRI. http://www.tropicalforages.info/

        Costa NV, Rodella RA, Martins D, 2006. Differentiation of aquatic weeds by multivariate analysis of foliar structural characters. (Diferenciação de espécies daninhas aquáticas pela análise multivariada de caracteres estruturais foliares.) Planta Daninha, 24(1):13-20. http://www.scielo.br/pdf/pd/v24n1/a02v24n1.pdf

        DAISIE, 2014. Delivering Alien Invasive Species Inventories for Europe. European Invasive Alien Species Gateway. www.europe-aliens.org/default.do

        Duke JA, 1983. Bracheri mutica (Forsk). Stapf. Handbook of Energy Crops. Unpublished. West Lafayette, Indiana, USA: Centre for New Crops and Plant Products, Purdue University. http://www.hort.purdue.edu/newcrop/duke_energy/Brachiaria_mutica.html#Yields%20and%20Economics

        Florence J, Chevillotte H, Ollier C, Meyer J-Y, 2013. Base de données botaniques Nadeaud de l'Herbier de la Polynésie Française (PAP) (Botanical database of the Nadeaud Herbarium of French Polynesia). http://www.herbier-tahiti.pf

        Florida Exotic Pest Plant Council, 2011. Florida EPPC's 2011 Invasive Plant Species List. http://www.fleppc.org/list/11list.html

        Funk V, Hollowell T, Berry P, Kelloff C, Alexander SN, 2007. Checklist of the plants of the Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana). Contributions from the United States National Herbarium, 584 pp

        González-Torres LR, Rankin R, Palmarola A (eds), 2012. Invasive plants in Cuba. (Plantas Invasoras en Cuba.) Bissea: Boletin sobre Conservacion de Plantad del Jardin Botanico Nacional, 6:1-140

        Graveson R, 2012. The Plants of Saint Lucia (in the Lesser Antilles of the Caribbean). The Plants of Saint Lucia (in the Lesser Antilles of the Caribbean). http://www.saintlucianplants.com

        Hannan-Jones M, Csurhes S, 2012. Para grass- Urichloa mutica. Invasive species risk assessment., Australia: Queensland Department of Agriculture, Fisheries and Forestry. http://www.daff.qld.gov.au/__data/assets/pdf_file/0004/65254/IPA-Para-Grass-Risk-Assessment.pdf

        Holm LG, Plucknett DL, Pancho JV, Herberger JP, 1977. The World's Worst Weeds. Distribution and Biology. Honolulu, Hawaii, USA: University Press of Hawaii

        I3N Brasil, 2014. National database of exotic invasive species (Base de dados nacional de espécies exóticas invasoras). Florianópolis - SC, Brazil: I3N Brasil, Instituto Hórus de Desenvolvimento e Conservação Ambiental. http://i3n.institutohorus.org.br

        Langeland KA, Cherry HM, McCormick CM, Craddock Burks KA, 2008. Identification and Biology of Non-native Plants in Florida's Natural Areas. Gainesville, Florida, USA: University of Florida IFAS Extension

        MacKee HS, 1994. Catalogue of introduced and cultivated plants in New Caledonia. (Catalogue des plantes introduites et cultivées en Nouvelle-Calédonie.) Paris, France: Muséum National d'Histoire Naturelle, unpaginated

        McCormack G, 2013. Cook Islands Biodiversity Database, Version 2007. Cook Islands Biodiversity Database. Rarotonga, Cook Islands: Cook Islands Natural Heritage Trust. http://cookislands.bishopmuseum.org/search.asp

        Morrone O, Zuloaga FO, 1992. A revision of the native and introduced South American species of Brachiaria (Trin.) Griseb. and Urochloa P. Beauv. (Poaceae: Panicoideae: Paniceae). (Revisión de las especies sudamericanas nátivas e introducidas de los géneros Brachiaria y Urochloa (Poaceae: Panicoideae: Paniceae).) Darwiniana, 31(1-4):43-109

        Morrone O, Zuloaga FO, 1993. Synopsis of the genus Urochloa (Poaceae: Panicoideae: Paniceae) from Mexico and Central America. (Sinopsis del género Urochloa (Poaceae: Panicoideae: Paniceae) para Mexico y América Central.) Darwiniana, 32:59-75

        PARSONS JJ, 1972. Spread of African pasture grasses to the American tropics. Journal of Range Management, 25(1):12-17

        PIER, 2014. Pacific Islands Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

        PROTA, 2014. PROTA4U web database. Grubben GJH, Denton OA, eds. Wageningen, Netherlands: Plant Resources of Tropical Africa. http://www.prota4u.org/search.asp

        Queensland Department of Primary Industries and Fisheries, 2011. Special edition of Environmental Weeds of Australia for Biosecurity Queensland., Australia: The University of Queensland and Department of Primary Industries and Fisheries. http://keyserver.lucidcentral.org/weeds/data/03030800-0b07-490a-8d04-0605030c0f01/media/Html/Index.htm

        Randall RP, 2012. A Global Compendium of Weeds. Perth, Australia: Department of Agriculture and Food Western Australia, 1124 pp. http://www.cabi.org/isc/FullTextPDF/2013/20133109119.pdf

        Sharp D, Simon BK, 2002. AusGrass1: Grasses of Australia. Canberra and Queensland, Australia: Australian Biological Resources Study and Environmental Protection Agency

        Shirasuna RT, 2014. Urochloa in Lista de Espécies da Flora do Brasil ([English title not available]). Rio de Janeiro, Brazil: Jardim Botânico do Rio de Janeiro. http://reflora.jbrj.gov.br/jabot/floradobrasil/FB24316

        Smith AC, 1979. Flora Vitiensis nova: A new flora of Fiji. Volume I. Lawai, Kauai, Hawaii, USA: National Tropical Botanical Garden, 494 pp

        Smith NM, 2002. Weeds of the wet/dry tropics of Australia - a field guide., Australia: Environment Centre NT, Inc, 112 pp

        Space JC, Flynn T, 2000. Observations on invasive plant species in American Samoa. Honolulu, USA: USDA Forest Service, 51 pp

        Space JC, Flynn T, 2001. Report to the Kingdom of Tonga on invasive plant species of environmental concern. Hawaii, USA: USDA Forest Service, Institute of Pacific Islands Forestry, 79 pp

        Space JC, Waterhouse B, Miles JE, Tiobech J, Rengulbai K, 2003. Report to the Republic of Palau on invasive plant species of environmental concern. Honolulu, Hawaii, USA: USDA Forest Service, 174 pp

        Space JC, Waterhouse BM, Newfield M, Bull C, 2004. Report to the Government of Niue and the United Nations Development Programme: Invasive plant species on Niue following Cyclone Heta. 80 pp. [UNDP NIU/98/G31 - Niue Enabling Activity.] http://www.hear.org/pier/reports/niue_report_2004.htm

        Stevens PF, 2012. Angiosperm Phylogeny Website. http://www.mobot.org/MOBOT/research/APweb/

        Stone BC, 1970. The flora of Guam. Micronesia, 6:1-659

        Sykes WR, 1977. Kermadec Islands flora: an annotated checklist. New Zealand Department of Scientific and Industrial Research Bulletin, 219:216 pp

        Torres González AM, Morton CM, 2005. Molecular and morphological phylogenetic analysis of Brachiaria and Urochloa (Poaceae). Molecular Phylogenetics and Evolution, 37(1):36-44. http://www.sciencedirect.com/science/journal/10557903

        US Fish and Wildlife Service, 2014. U.S. Fish and Wildlife Service species assessment and listing priority assignment form: Megalagrion xanthomelas. In: U.S. Fish and Wildlife Service species assessment and listing priority assignment form: Megalagrion xanthomelas : US Fish and Wildlife Service.14 pp. http://ecos.fws.gov/docs/candidate/assessments/2014/r1/I063_I01.pdf

        USDA-ARS, 2014. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

        USDA-NRCS, 2014. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

        Veldkamp JF, 2004. Miscellaneous notes on mainly Southeast Asian gramineae. Reinwardtia, 12:135-140

        Villaseñor JL, Espinosa-Garcia FJ, 2004. The alien flowering plants of Mexico. Diversity and Distributions, 10(2):113-123

        Wagner WI, Herbst DR, Sohmer SH, 1999. Manual of the Flowering Plants of Hawaii, revised edition. Honolulu, Hawaii, USA: University of Hawaii Press

        Waterhouse DF, 1993. The Major Arthropod Pests and Weeds of Agriculture in Southeast Asia. ACIAR Monograph No. 21. Canberra, Australia: Australian Centre for International Agricultural Research, 141 pp

        Webster RD, 1987. Australian Paniceae (Poaceae). Berlin, Germany: J. Cramer, 322pp

        Webster RD, 1988. Genera of the North American Paniceae (Poaceae: Panicoideae). Systematic Botany, 13(4):576-609

        Wu T-L, 2001. Check List of Hong Kong Plants. Hong Kong Herbarium and the South China Institute of Botany. Agriculture, Fisheries and Conservation Department Bulletin (revised), 1:384 pp

        Zuloaga FO, Morrone O, 2003. Brachiaria, Urochloa. Contributions of the US National Herbarium, 46:141-143, 629-634. [Catalogue of New World Grasses (Poaceae): III. Subfamilies Panicoideae, Aristidoideae, Arundionoideae, and Danthonioideae.]

        Distribution References

        Acevedo-Rodríguez P, Strong M T, 2012. Catalogue of the Seed Plants of the West Indies. Washington, DC, USA: Smithsonian Institution. 1192 pp. http://botany.si.edu/Antilles/WestIndies/catalog.htm

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

        Chacón E, Saborío G, 2012. [English title not available]. (Red Interamericana de Información de Especies Invasoras, Costa Rica)., San José, Costa Rica: Asociación para la Conservación y el Estudio de la Biodiversidad. http://invasoras.acebio.org

        Chaudhari S, Sellers B A, Rockwood S V, Ferrell J A, MacDonald G E, Kenworthy K E, 2012. Nonchemical methods for paragrass (Urochloa mutica) control. Invasive Plant Science and Management. 5 (1), 20-26. http://wssajournals.org/loi/ipsm DOI:10.1614/IPSM-D-11-00026.1

        Chaudhari S, Sellers B A, Rockwood S V, Ferrell J A, MacDonald G E, Kenworthy K E, 2012a. Integrating chemical and cultural practices to control para grass (Urochloa mutica). Journal of Aquatic Plant Management. 39-45. http://www.apms.org/wp/wp-content/uploads/2012/01/2-17661-p039-45-APMdj.pdf

        Chong KY, Tan HTW, Corlett RT, 2009. A checklist of the total vascular plant flora of Singapore: native, naturalised and cultivated species. In: A checklist of the total vascular plant flora of Singapore, Singapore: National University of Singapore, Raffles Museum of Biodiversity Research. 273 pp.

        Clayton WD, Govaerts R, Harman KT, Williamson H, Vorontsova M, 2014. World Checklist of Poaceae., Richmond, UK: Royal Botanic Gardens, Kew. http://apps.kew.org/wcsp/

        DAISIE, 2014. Delivering Alien Invasive Species Inventories for Europe. http://www.europe-aliens.org/

        Douglas M M, O'Connor R A, 2003. Effects of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain. Freshwater Biology. 48 (6), 962-971. DOI:10.1046/j.1365-2427.2003.01072.x

        Florence J, Chevillotte H, Ollier C, Meyer J-Y, 2013. Botanical database of the Nadeaud Herbarium of French Polynesia. (Base de données botaniques Nadeaud de l'Herbier de la Polynésie Française (PAP))., http://www.herbier-tahiti.pf

        Florida Exotic Pest Plant Council, 2011. Florida EPPC's 2011 Invasive Plant Species List., http://www.fleppc.org/list/11list.html

        Funk V, Hollowell T, Berry P, Kelloff C, Alexander S N, 2007. Contributions from the United States National Herbarium, Washington, USA: Department of Systematic Biology - Botany, National Museum of Natural History, Smithsonian Institution. 55, 584 pp.

        González-Torres LR, Rankin R, Palmarola A, 2012. Invasive plants in Cuba. (Plantas Invasoras en Cuba). In: Bissea: Boletin sobre Conservacion de Plantad del Jardin Botanico Nacional, 6 [ed. by González-Torres LR, Rankin R, Palmarola A]. 1-140.

        Graveson R, 2012. The Plants of Saint Lucia (in the Lesser Antilles of the Caribbean). In: The Plants of Saint Lucia (in the Lesser Antilles of the Caribbean), http://www.saintlucianplants.com

        Holm L G, Plucknett D L, Pancho J V, Herberger J P, 1977. The world's worst weeds. Distribution and biology. Honolulu, Hawaii, USA: University Press of Hawaii. 610 pp.

        I3N Brasil, 2014. National database of exotic invasive species. (Base de dados nacional de espécies exóticas invasoras)., Florianópolis - SC, Brazil: I3N Brasil, Instituto Hórus de Desenvolvimento e Conservação Ambiental. http://i3n.institutohorus.org.br

        MacKee H S, 1994. Catalogue des plantes introduites et cultivées en Nouvelle-Calédonie. Paris, France: Muséum National d'Histoire Naturelle. unpaginated.

        McCormack G, 2007. Cook Islands Biodiversity Database, Version 2007.2. In: Cook Islands Biodiversity Database, Version 2007.2, Rarotonga: Cook Islands Natural Heritage Trust. http://cookislands.bishopmuseum.org

        PIER, 2014. Pacific Islands Ecosystems at Risk., Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

        Shirasuna RT, 2014. [English title not available]. (Urochloa in Lista de Espécies da Flora do Brasil)., Rio de Janeiro, Brazil: Jardim Botânico do Rio de Janeiro. http://reflora.jbrj.gov.br/jabot/floradobrasil/FB24316

        Smith AC, 1979. Flora Vitiensis nova: A new flora of Fiji., I Lawai, Kauai, Hawaii, USA: National Tropical Botanical Garden. 494 pp.

        Smith NM, 2002. Weeds of the wet/dry tropics of Australia - a field guide., Australia: Environment Centre NT, Inc. 112 pp.

        Space J C, Waterhouse B M, Newfield M, Bull C, 2004. Report to the Government of Niue and the United Nations Development Programme: Invasive plant species on Niue following Cyclone Heta. [UNDP NIU/98/G31 - Niue Enabling Activity. In: Report to the Government of Niue and the United Nations Development Programme: Invasive plant species on Niue following Cyclone Heta. [UNDP NIU/98/G31 - Niue Enabling Activity.]. 76 pp. http://www.hear.org/pier/reports/niue_report_2004.htm

        Space JC, Flynn T, 2000. Observations on invasive plant species in American Samoa., Honolulu, USA: USDA Forest Service. 51 pp.

        Space JC, Flynn T, 2001. Report to the Kingdom of Tonga on invasive plant species of environmental concern., Hawaii, USA: USDA Forest Service, Institute of Pacific Islands Forestry. 79 pp.

        Space JC, Waterhouse B, Miles JE, Tiobech J, Rengulbai K, 2003. Report to the Republic of Palau on invasive plant species of environmental concern., Honolulu, Hawaii, USA: USDA Forest Service. 174 pp.

        Stone B C, 1970. The flora of Guam. A manual for the identification of the vascular plants of the island. Guam: University of Guam. vi + 659 pp.

        Sykes WR, 1977. Kermadec Islands flora: an annotated checklist. In: New Zealand Department of Scientific and Industrial Research Bulletin, 219 216 pp.

        USDA-ARS, 2014. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

        USDA-NRCS, 2014. The PLANTS Database. Greensboro, North Carolina, USA: National Plant Data Team. https://plants.sc.egov.usda.gov

        Villaseñor J L, Espinosa-Garcia F J, 2004. The alien flowering plants of Mexico. Diversity and Distributions. 10 (2), 113-123. DOI:10.1111/j.1366-9516.2004.00059.x

        Wagner WI, Herbst DR, Sohmer SH, 1999. Manual of the Flowering Plants of Hawaii, revised edition., Honolulu, Hawaii, USA: University of Hawaii Press.

        Waterhouse D F, 1993. The major arthropod pests and weeds of agriculture in Southeast Asia. Canberra, Australia: ACIAR. v + 141 pp.

        Wu TL, 2001. Check List of Hong Kong Plants. Hong Kong Herbarium and the South China Institute of Botany. In: Agriculture, Fisheries and Conservation Department Bulletin (revised), 1 384 pp.

        Links to Websites

        Top of page
        WebsiteURLComment
        Catalogue of Seed Plants of the West Indieshttp://botany.si.edu/antilles/WestIndies/catalog.htm
        Grasses in North Americahttp://herbarium.usu.edu/webmanual/
        Pacific Island Ecosystems at Risk (PIER): Plant threats to Pacific ecosystemshttp://www.hear.org/pier/index.html
        Plant Resources of Tropical Africahttp://www.prota.org
        Tropical Forageshttp://www.tropicalforages.info/index.htm

        Contributors

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        12/02/14 Original text by:

        Julissa Rojas-Sandoval, Department of Botany-Smithsonian NMNH, Washington DC, USA

        Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

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