Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Cyanthillium cinereum
(little ironweed)

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Datasheet

Cyanthillium cinereum (little ironweed)

Summary

  • Last modified
  • 10 December 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Host Plant
  • Preferred Scientific Name
  • Cyanthillium cinereum
  • Preferred Common Name
  • little ironweed
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • C. cinereum is a cosmopolitan weed common in disturbed areas in tropical and subtropical regions of the world (Randall, 2012). It is a fast-gr...

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Pictures

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PictureTitleCaptionCopyright
Cyanthillium cinereum (little ironweed); Flowers. Hana LZ Hana Ranch, Maui, Hawaii. February 2009.
TitleFlowers
CaptionCyanthillium cinereum (little ironweed); Flowers. Hana LZ Hana Ranch, Maui, Hawaii. February 2009.
Copyright©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowers. Hana LZ Hana Ranch, Maui, Hawaii. February 2009.
FlowersCyanthillium cinereum (little ironweed); Flowers. Hana LZ Hana Ranch, Maui, Hawaii. February 2009.©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowers. Kipahulu LZ Haleakala National Park, Maui, Hawaii. February 2009.
TitleFlowers
CaptionCyanthillium cinereum (little ironweed); Flowers. Kipahulu LZ Haleakala National Park, Maui, Hawaii. February 2009.
Copyright©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowers. Kipahulu LZ Haleakala National Park, Maui, Hawaii. February 2009.
FlowersCyanthillium cinereum (little ironweed); Flowers. Kipahulu LZ Haleakala National Park, Maui, Hawaii. February 2009.©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowers. Wailua, Maui, Hawaii. September 2014.
TitleFlowers
CaptionCyanthillium cinereum (little ironweed); Flowers. Wailua, Maui, Hawaii. September 2014.
Copyright©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowers. Wailua, Maui, Hawaii. September 2014.
FlowersCyanthillium cinereum (little ironweed); Flowers. Wailua, Maui, Hawaii. September 2014.©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Seedhead. Kerala, India. August 2015
TitleSeedhead
CaptionCyanthillium cinereum (little ironweed); Seedhead. Kerala, India. August 2015
Copyright©Vengolis/via Wikimedia Commons - CC BY-SA 4.0
Cyanthillium cinereum (little ironweed); Seedhead. Kerala, India. August 2015
SeedheadCyanthillium cinereum (little ironweed); Seedhead. Kerala, India. August 2015©Vengolis/via Wikimedia Commons - CC BY-SA 4.0
Cyanthillium cinereum (little ironweed); Seedhead. Arakoon, New South Wales, Australia. May 2009.
TitleSeedhead
CaptionCyanthillium cinereum (little ironweed); Seedhead. Arakoon, New South Wales, Australia. May 2009.
Copyright©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Seedhead. Arakoon, New South Wales, Australia. May 2009.
SeedheadCyanthillium cinereum (little ironweed); Seedhead. Arakoon, New South Wales, Australia. May 2009.©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowers. Hakioawa, Kahoolawe, Hawaii. December 2015.
TitleFlowers
CaptionCyanthillium cinereum (little ironweed); Flowers. Hakioawa, Kahoolawe, Hawaii. December 2015.
Copyright©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowers. Hakioawa, Kahoolawe, Hawaii. December 2015.
FlowersCyanthillium cinereum (little ironweed); Flowers. Hakioawa, Kahoolawe, Hawaii. December 2015.©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowerheads. Dungog, New South Wales, Australia. January 2015.
TitleFlowerhead
CaptionCyanthillium cinereum (little ironweed); Flowerheads. Dungog, New South Wales, Australia. January 2015.
Copyright©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Flowerheads. Dungog, New South Wales, Australia. January 2015.
FlowerheadCyanthillium cinereum (little ironweed); Flowerheads. Dungog, New South Wales, Australia. January 2015.©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Cauline leaf.  Dungog, New South Wales, Australia. January 2015.
TitleCauline leaf
CaptionCyanthillium cinereum (little ironweed); Cauline leaf. Dungog, New South Wales, Australia. January 2015.
Copyright©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Cauline leaf.  Dungog, New South Wales, Australia. January 2015.
Cauline leafCyanthillium cinereum (little ironweed); Cauline leaf. Dungog, New South Wales, Australia. January 2015.©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Leaves. Dungog, New South Wales, Australia. January 2015.
TitleLeaves
CaptionCyanthillium cinereum (little ironweed); Leaves. Dungog, New South Wales, Australia. January 2015.
Copyright©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Leaves. Dungog, New South Wales, Australia. January 2015.
LeavesCyanthillium cinereum (little ironweed); Leaves. Dungog, New South Wales, Australia. January 2015.©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Basal rosette of leaves. Dungog, New South Wales, Australia. January 2015.
TitleBasal leaves
CaptionCyanthillium cinereum (little ironweed); Basal rosette of leaves. Dungog, New South Wales, Australia. January 2015.
Copyright©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Basal rosette of leaves. Dungog, New South Wales, Australia. January 2015.
Basal leavesCyanthillium cinereum (little ironweed); Basal rosette of leaves. Dungog, New South Wales, Australia. January 2015.©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Plant. Dungog, New South Wales, Australia. January 2015.
TitlePlant
CaptionCyanthillium cinereum (little ironweed); Plant. Dungog, New South Wales, Australia. January 2015.
Copyright©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Plant. Dungog, New South Wales, Australia. January 2015.
PlantCyanthillium cinereum (little ironweed); Plant. Dungog, New South Wales, Australia. January 2015.©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Habit. Wailua, Maui, Hawaii. September 2014.
TitleHabit
CaptionCyanthillium cinereum (little ironweed); Habit. Wailua, Maui, Hawaii. September 2014.
Copyright©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Habit. Wailua, Maui, Hawaii. September 2014.
HabitCyanthillium cinereum (little ironweed); Habit. Wailua, Maui, Hawaii. September 2014.©Forest and Kim Starr/via Starr Environmental - CC BY 2.0
Cyanthillium cinereum (little ironweed); Seedling. Arakoon, New South Wales, Australia. May 2009.
TitleSeedling
CaptionCyanthillium cinereum (little ironweed); Seedling. Arakoon, New South Wales, Australia. May 2009.
Copyright©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Seedling. Arakoon, New South Wales, Australia. May 2009.
SeedlingCyanthillium cinereum (little ironweed); Seedling. Arakoon, New South Wales, Australia. May 2009.©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Leaves, ventral (left) and dorsal (right). Dungog, New South Wales, Australia. July 2015.
TitleLeaves
CaptionCyanthillium cinereum (little ironweed); Leaves, ventral (left) and dorsal (right). Dungog, New South Wales, Australia. July 2015.
Copyright©Harry Rose/via Flickr - CC BY 2.0
Cyanthillium cinereum (little ironweed); Leaves, ventral (left) and dorsal (right). Dungog, New South Wales, Australia. July 2015.
LeavesCyanthillium cinereum (little ironweed); Leaves, ventral (left) and dorsal (right). Dungog, New South Wales, Australia. July 2015.©Harry Rose/via Flickr - CC BY 2.0

Identity

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Preferred Scientific Name

  • Cyanthillium cinereum (L.) H. Rob

Preferred Common Name

  • little ironweed

Other Scientific Names

  • Blumea esquirolii H.Lev. & Vaniot
  • Cacalia arguta Kuntze
  • Cacalia cinerea (L.) Kuntze
  • Cacalia erigerodes Kuntze
  • Cacalia exilis Kuntze
  • Cacalia kroneana Kuntze
  • Cacalia linfolia DC.
  • Cacalia rotundifolia Willd.
  • Cacalia vialis Kuntze
  • Calea cordata Lour.
  • Cineraria glaberrima Spreng. ex DC.
  • Conyza chinensis L.
  • Conyza cinerea L.
  • Conzya heterophylla Lam.
  • Conzya incana DC.
  • Conzya prolifera Lam.
  • Crassocephalum flatmense Hochst. & Steud. ex DC.
  • Cyanopis decurrens Zoll. & Mor.
  • Eupatorium arboreum Reinw. ex de Vriese
  • Eupatorium myosotifolium Jacq.
  • Pteronia tomentosa Lour.
  • Seneciodes cinereum (L.) Kuntze
  • Serratula cinerea (L.) Roxb.
  • Vernonia abbreviata DC.
  • Vernonia arguta Baker
  • Vernonia betonicaefolia Baker
  • Vernonia cinerea (L.) Less.
  • Vernonia exilis Miq.
  • Vernonia fasciculata Blume
  • Vernonia kroneana Miq.
  • Vernonia vialis DC.

International Common Names

  • English: ironweed; small ironweed; vernonia
  • Spanish: machadita; rabo de buey; yerba morada
  • French: ayapana sauvage
  • Chinese: ye xiang niu

Local Common Names

  • Cuba: machadita
  • Fiji: kaukamea; tho vuka; vutikaumondro
  • India: ankari; ankta; pokasubgo; sahaderi; sandri
  • Indonesia: maryuna
  • Japan: mura-saki-mukashi-yomogi; reinan-nogiku; yambaru-higotai
  • Lesser Antilles: inflammation bush; iron bush; measle bush
  • Malaysia: rumput taki babi; tambak-tambak
  • Mauritius: ayapa sauvage
  • Nigeria: bojure
  • Philippines: agas-moro; bulak-manok; kolong-kugon
  • Puerto Rico: rabo de buey; yerba socialista
  • Sri Lanka: alavangu pillu; monara kudumbiya
  • Thailand: kaan thuup; yaa dok khaao; yaa-saam-wan; ya-la-ong

Summary of Invasiveness

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C. cinereum is a cosmopolitan weed common in disturbed areas in tropical and subtropical regions of the world (Randall, 2012). It is a fast-growing, annual herb with the capacity to form dense patches in gardens, roadsides, waste grounds and pasture (Holm et al., 1997). Currently, this species is considered invasive in many islands in the Pacific Ocean (e.g. Hawaii, Fiji, French Polynesia and Micronesia), New Zealand, Singapore, Costa Rica, Guatemala, Nicaragua, Panama, Galápagos Islands, Cuba, Puerto Rico and the Virgin Islands (see distribution table for details; Chong et al., 2009; Chacon and Saborio, 2012; González-Torres et al., 2012; PIER, 2013).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Asterales
  •                         Family: Asteraceae
  •                             Genus: Cyanthillium
  •                                 Species: Cyanthillium cinereum

Notes on Taxonomy and Nomenclature

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The family Asteraceae is one of the most diverse groups among flowering plants including 1620 genera and about 23,600 species (Stevens, 2012). Species in the Asteraceae are very variable vegetatively, but may be recognized by their “capitulate” and involucrate inflorescences in which numerous small flowers open first on the outside and are infrequently subtended by bracts. The anthers in this family are usually fused and form a tube through which the style extends before the two stigmatic lobes separate and become recurved. The rather small, single-seeded fruits usually have a plumose “pappus” and are frequently dispersed by wind (Stevens, 2012).

The genus Cyanthillium has been used as a synonym of Vernonia, while other botanists have treated both genera as distinct. Currently, both genera are treated separately and Cyanthillium is distinguished from Vernonia by its smooth to finely 5-striate (vs. 5-10-ribbed) achenes, and by its echinolophate (vs. tricolporate) pollen, and by base chromosome number usually reported as 2n = 18 or 20 (vs. 2n = 34). Currently five species are recognized within Cyanthillium (Funk and Pruski, 1996).

Description

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Annual herb, to 80 (-150) cm tall. Stems unbranched or more commonly few-branched. Leaves petiolate or distal ones sessile; petioles to 1.5 cm, narrowly winged; blade 1.5-5 × 1-1.5(4) cm, obovate or less often elliptic or ovate, base cuneate, apex acute to obtuse, lower surface pilose, usually glandular. Inflorescence of 5 to numerous heads, 13-20-flowered; involucre 2.5-3 mm long; outer phyllaries much reduced, inner phyllaries subequal. Corolla 3-4 mm, exserted 1.5-2 mm from involucre, the tube long and narrow, 2-3 mm, the limbs short, pilose. Achenes 1.2–2 mm long, subfusiform, terete not ribbed, inner pappus white, exserted from involucre and nearly as long as the corollas (Funk and Pruski, 1996).

Plant Type

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Annual
Herbaceous
Seed propagated

Distribution

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C. cinereum is native to Africa, tropical and temperate Asia, and Australia (USDA-ARS, 2013). It is widely naturalized in tropical regions in America and islands in the Pacific (PIER, 2013; USDA-ARS, 2013).   

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: 10 Jan 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

BeninPresentNative
BurundiPresentNative
CameroonPresentNative
Central African RepublicPresentNative
Congo, Democratic Republic of thePresentNative
Côte d'IvoirePresentNative
EritreaPresentNative
EswatiniPresentNative
EthiopiaPresentNative
GabonPresentNative
GhanaPresentNative
GuineaPresentNative
Guinea-BissauPresentNative
KenyaPresentNative
LiberiaPresentNative
MadagascarPresentNative
MaliPresentNative
MayottePresentIntroducedInvasive
NigerPresentNative
RéunionPresentNative
RwandaPresentNative
SenegalPresentNative
SeychellesPresentIntroducedInvasive
Sierra LeonePresentNative
SomaliaPresentNative
South AfricaPresentNative
TanzaniaPresentNative
TogoPresentNative
UgandaPresentNative

Asia

BangladeshPresentWeed
BhutanPresentNative
British Indian Ocean Territory
-Chagos ArchipelagoPresentIntroducedInvasive
ChinaPresentPresent based on regional distribution.
-FujianPresentNativeConsidered a weed
-GuangdongPresentNativeConsidered a weed
-GuangxiPresentNativeConsidered a weed
-HubeiPresentNativeConsidered a weed
-HunanPresentNativeConsidered a weed
-JiangxiPresentNativeConsidered a weed
-SichuanPresentNativeConsidered a weed
-YunnanPresentNativeConsidered a weed
-ZhejiangPresentNativeConsidered a weed
IndiaPresentNative
IndonesiaPresentNative
JapanPresentNative
-KyushuPresentNative
-Ryukyu IslandsPresentNative
MalaysiaPresentNative
MaldivesPresent
MyanmarPresentNative
OmanPresentNative
PhilippinesPresentNative
SingaporePresentInvasive
Sri LankaPresentNative
TaiwanPresentNativeConsidered a weed
ThailandPresentNative
VietnamPresentNative
YemenPresentNative

North America

AnguillaPresentIntroduced
Antigua and BarbudaPresentIntroduced
BahamasPresentIntroduced
BarbadosPresentIntroduced
BelizePresentIntroduced
British Virgin IslandsPresentIntroducedGuana, Tortola, Virgin Gorda
Costa RicaPresentIntroducedInvasive
CubaPresentIntroducedInvasiveHighly invasive
DominicaPresentIntroduced
Dominican RepublicPresentIntroduced
GrenadaPresentIntroduced
GuatemalaPresentIntroducedInvasive
HaitiPresentIntroduced
HondurasPresentIntroducedInvasive
JamaicaPresentIntroduced
MartiniquePresentIntroduced
MexicoPresentIntroducedWeed
MontserratPresentIntroduced
Netherlands AntillesPresentIntroducedSaba, St. Marteen
NicaraguaPresentIntroducedInvasive
PanamaPresentIntroducedInvasive
Puerto RicoPresentIntroducedInvasive
Saint Kitts and NevisPresentIntroduced
Saint LuciaPresentIntroduced
Saint Vincent and the GrenadinesPresentIntroduced
Trinidad and TobagoPresentIntroduced
U.S. Virgin IslandsPresentIntroducedInvasive
United StatesPresentPresent based on regional distribution.
-FloridaPresentIntroduced
-HawaiiPresentIntroducedInvasive

Oceania

American SamoaPresentIntroducedInvasive
AustraliaPresentPresent based on regional distribution.
-New South WalesPresentNative
-Northern TerritoryPresentNative
-QueenslandPresentNative
-Western AustraliaPresentNative
Christmas IslandPresentIntroducedInvasive
Cook IslandsPresentIntroduced
Federated States of MicronesiaPresentInvasiveThis species has been listed as native for this archipelago
FijiPresentIntroducedInvasive
French PolynesiaPresentIntroducedInvasive
GuamPresentIntroducedInvasive
KiribatiPresentIntroducedInvasiveFormerly listed as native
Marshall IslandsPresentInvasiveListed as native for some islands and naturalised and invasive for other islands
NauruPresentIntroducedInvasiveFormerly listed as native
New CaledoniaPresentIntroducedInvasive
NiuePresentIntroducedInvasive
Northern Mariana IslandsPresentInvasiveListed as native for some islands and naturalised and invasive for other islands
PalauPresentNative
Papua New GuineaPresentNative
PitcairnPresentIntroducedInvasive
SamoaPresent
Solomon IslandsPresent
TongaPresentIntroducedInvasive
TuvaluPresent
U.S. Minor Outlying IslandsPresentIntroducedInvasive
VanuatuPresent
Wallis and FutunaPresentIntroducedInvasive

South America

BrazilPresentPresent based on regional distribution.
-AmazonasPresentIntroduced
-BahiaPresentIntroduced
-Minas GeraisPresentIntroduced
-ParaPresentIntroduced
-PernambucoPresentIntroduced
-Rio de JaneiroPresentIntroduced
-Sao PauloPresentIntroduced
EcuadorPresentPresent based on regional distribution.
-Galapagos IslandsPresentIntroducedInvasive
French GuianaPresentIntroducedNaturalizedNaturalised
GuyanaPresentIntroducedNaturalizedNaturalised
SurinamePresentIntroducedNaturalizedNaturalised
VenezuelaPresentIntroducedNaturalizedNaturalised

History of Introduction and Spread

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The history of introduction of C. cinereum is uncertain. It is highly probable that the species has been introduced accidentally as a contaminant or as a weed in nursery materials (Holm et al., 1997). For the West Indies, herbarium collections show that it was first collected in Jamaica in 1882 (US Herbarium Collection). In Central America the date of introduction is unclear but since the 1950s the species appears listed as a weed (Cardenas and Coulston, 1967).

Risk of Introduction

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The risk of introduction of C. cinereum is high. This species is easily dispersed by wind and has the potential to grow as a weed in ruderal areas, and agricultural and pasture lands. C. cinereum has been reported as a weed in 27 crops in 47 countries in Asia, East and West Africa and the Caribbean (Holm et al., 1997). Consequently, this species has the potential to spread much further than it has to date.

Habitat

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C. cinereum is a common weed in coastal areas, beaches, cultivated areas, disturbed areas, pastures, roadsides, rocky areas, savannas, secondary vegetation, and gardens. In Africa (within its native distribution range), this species grows at lower altitudes (0-800 m) in woodlands and grasslands (PROTA4U, 2013).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial ManagedCultivated / agricultural land Present, no further details Harmful (pest or invasive)
Terrestrial ManagedCultivated / agricultural land Present, no further details 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 grasslands (grazing systems) Present, no further details Harmful (pest or invasive)
Terrestrial ManagedManaged grasslands (grazing systems) Present, no further details Natural
Terrestrial ManagedDisturbed areas Present, no further details Harmful (pest or invasive)
Terrestrial ManagedDisturbed areas Present, no further details Natural
Terrestrial ManagedRail / roadsides Present, no further details Harmful (pest or invasive)
Terrestrial ManagedRail / roadsides Present, no further details Natural
Terrestrial ManagedUrban / peri-urban areas Present, no further details Harmful (pest or invasive)
Terrestrial ManagedUrban / peri-urban areas Present, no further details Natural
Terrestrial Natural / Semi-naturalNatural forests Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural forests Present, no further details Natural
Terrestrial Natural / Semi-naturalNatural grasslands Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural grasslands Present, no further details Natural
LittoralCoastal areas Present, no further details Harmful (pest or invasive)
LittoralCoastal areas Present, no further details Natural

Hosts/Species Affected

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C. cinereum is reported as a weed in 27 crops in 47 countries (Holm et al., 1997).

Serious weed in:

  • sugarcane, cotton, groundnuts and wheat in India
  • pastures in Australia, India, Nigeria and Thailand
  • rice in Philippines;
  • taro in Samoa. 

Common weed in:

  • banana in Surinam and Tonga
  • cassava in Surinam
  • cocoa in Indonesia
  • citrus in Surinam
  • cotton in the Philippines
  • maize in India
  • oil palm in Surinam
  • pastures in Australia, Dominican Republic, Puerto Rico and Jamaica
  • pineapple in Hawaii
  • rice in Surinam, India, Indonesia and Sri Lanka
  • rubber in Indonesia and Thailand
  • sugarcane in Bangladesh, Hawaii, and the Philippines
  • taro in Tonga
  • tea in India and Indonesia
  • vegetables in Surinam and Thailand 

Unranked weed in:

  • abaca (Musa textilis) in the Philippines
  • cocoa in Dominican Republic
  • cassava in India, Indonesia, and Nigeria
  • coconut in Sri Lanka and Surinam
  • coffee in Dominican Republic
  • cotton in Mozambique and Tanzania
  • legumes and tomatoes in the Philippines
  • macadamia nut in Hawaii
  • maize in Cambodia, Gambia, Indonesia, Nigeria, the Philippines and Zambia
  • pastures in the Philippines
  • groundnuts in Indonesia and Nigeria
  • rice in Laos, Thailand and Vietnam
  • rubber and tea in Sri Lanka
  • sugarcane in British Guiana, Dominican Republic, Laos and Vietnam
  • tobacco in the Philippines

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Ananas comosus (pineapple)BromeliaceaeMain
    Arachis hypogaea (groundnut)FabaceaeMain
      CitrusRutaceaeMain
        Cocos nucifera (coconut)ArecaceaeMain
          Coffea arabica (arabica coffee)RubiaceaeMain
            Colocasia esculenta (taro)AraceaeMain
              Elaeis guineensis (African oil palm)ArecaceaeMain
                Gossypium (cotton)MalvaceaeMain
                  Hevea brasiliensis (rubber)EuphorbiaceaeMain
                    Macadamia integrifolia (macadamia nut)ProteaceaeMain
                      Manihot esculenta (cassava)EuphorbiaceaeMain
                        Musa (banana)MusaceaeMain
                          NicotianaSolanaceaeMain
                            Oryza sativa (rice)PoaceaeMain
                              SaccharumPoaceaeMain
                                Solanum lycopersicum (tomato)SolanaceaeMain
                                  Theobroma cacao (cocoa)MalvaceaeMain
                                    Triticum (wheat)PoaceaeMain
                                      Zea mays (maize)PoaceaeMain

                                        Growth Stages

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                                        Flowering stage, Vegetative growing stage

                                        Biology and Ecology

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                                        Genetics

                                        C. cinereum plants are diploid with a chromosome number of 2n = 18 (Holm et al., 1997; Pruski 2013). 

                                        Reproductive Biology

                                        Flowers in C. cinereum are pollinated by wind. Probably the most common reproduction system in this genus is allogamy with a sporophytic self-incompatibility (Holm et al., 1997; Flora of China Editorial Committee, 2012).

                                        Physiology and Phenology

                                        C. cinereum is an annual herb and under favourable environmental conditions it produces flowers and seeds for many months (Holm et al., 1997; Flora of China Editorial Committee, 2012).   

                                        Environmental Requirements

                                        C. cinereum usually grows as a weed, thus it needs full sunlight and moderate water availability to grow. It prefers sandy-loam soils but can be found growing on a range of soils with pH ranging from 4 to 6. It is able to tolerate semiarid conditions as well as partial salinity conditions (PROTA4U, 2013).

                                        Climate

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                                        ClimateStatusDescriptionRemark
                                        Af - Tropical rainforest climate Preferred > 60mm precipitation per month
                                        Am - Tropical monsoon climate Preferred Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
                                        As - Tropical savanna climate with dry summer Preferred < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
                                        Aw - Tropical wet and dry savanna climate Preferred < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
                                        BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
                                        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)

                                        Rainfall Regime

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                                        Bimodal

                                        Soil Tolerances

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                                        Soil drainage

                                        • free

                                        Soil reaction

                                        • acid
                                        • neutral

                                        Soil texture

                                        • heavy
                                        • light
                                        • medium

                                        Special soil tolerances

                                        • saline

                                        Means of Movement and Dispersal

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                                        C. cinereum spreads by seeds which are adapted to wind dispersal. In addition, seeds may be secondarily dispersed as a contaminant in crop seeds, pasture seeds and in agricultural machinery (Holm et al., 1997; Pruski 2013).

                                        Pathway Causes

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                                        CauseNotesLong DistanceLocalReferences
                                        Crop productionProbably dispersed as a seed and crop contaminant Yes Yes Holm et al., 1997
                                        DisturbanceCommon weed Yes Yes Holm et al., 1997
                                        Escape from confinement or garden escapeCommon weed Yes Yes Holm et al., 1997
                                        ForageCommon weed in pasturelands Yes Yes Holm et al., 1997
                                        Garden waste disposalCommon weed in gardens Yes Yes Holm et al., 1997
                                        HorticultureCommon weed Yes Yes Holm et al., 1997
                                        Medicinal useUsed in African traditional herb medicine Yes Yes Prota4U, 2013

                                        Pathway Vectors

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                                        VectorNotesLong DistanceLocalReferences
                                        Machinery and equipmentSeeds Yes Yes Holm et al., 1997
                                        Soil, sand and gravelSeeds Yes Yes Holm et al., 1997
                                        WaterSeeds Yes Yes Holm et al., 1997

                                        Impact Summary

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

                                        Economic Impact

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                                        C. cinereum is a weed with important economic impacts mainly in agriculture, pasture and garden activities. It is listed as a weed in about 27 different crops in 47 countries in Asia, Australia, Africa and America. In addition, C. cinereum has the potential to negatively impact natural and seminatural habitats in coastal areas, forest edges, secondary forests, upland forests, and grasslands (Pruski, 2013).

                                        C. cinereum can host the tobacco leaf curl virus and the root-knot nematode (Holm et al., 1997).

                                        Threatened Species

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                                        Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
                                        Schiedea spergulina var. leiopodaNational list(s); USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resourcesUS Fish and Wildlife Service, 2010

                                        Risk and Impact Factors

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                                        Invasiveness
                                        • Invasive in its native range
                                        • Proved invasive outside its native range
                                        • Has a broad native range
                                        • Abundant in its native range
                                        • Highly adaptable to different environments
                                        • Is a habitat generalist
                                        • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
                                        • Pioneering in disturbed areas
                                        • Highly mobile locally
                                        • Benefits from human association (i.e. it is a human commensal)
                                        • Fast growing
                                        Impact outcomes
                                        • Damaged ecosystem services
                                        • Ecosystem change/ habitat alteration
                                        • Loss of medicinal resources
                                        • Modification of successional patterns
                                        • Monoculture formation
                                        • Negatively impacts agriculture
                                        • Negatively impacts cultural/traditional practices
                                        • Negatively impacts forestry
                                        Impact mechanisms
                                        • Competition - monopolizing resources
                                        • Pest and disease transmission
                                        • Rapid growth
                                        Likelihood of entry/control
                                        • Highly likely to be transported internationally accidentally
                                        • Difficult to identify/detect as a commodity contaminant
                                        • Difficult/costly to control

                                        Uses

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                                        Leaves of C. cinereum have some antibiotic activity and consequently they are frequently used in traditional medicine in India to treat conjunctivitis, dropsy and urinary disorders. In the Philippines, this species is used as an infusion for cough and skin diseases. A poultice from leaves reduces headaches while a root decoction relieves stomach aches and diarrhea. In Thailand, the leaves are used in the treatment of asthma and bronchitis (Holm et al., 1997; PROTA4U, 2013). The young shoots are eaten as a cooked vegetable in Java (PROTA4U, 2013).

                                        Uses List

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                                        Medicinal, pharmaceutical

                                        • Traditional/folklore

                                        Similarities to Other Species/Conditions

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                                        Species of Cyanthillium can be confused with species in the genus Vernonia. These two genera can be distinguished by the following characters:

                                        1. In Cyanthillium the cypselae is smooth to finely 5-striate while in Vernonia cypselae is 5-10-ribbed.
                                        2. Pollen in Cyanthillium is echinolophate while in Vernonia pollen is tricolporate
                                        3. The chromosome number usually reported for Cyanthillium species is 2n = 18 or 20 and for Vernonia species is 2n = 34.

                                        The species C. cinereum is recognized by the slender, ribbed stem, shallowly toothed leaves and the branched corymbs with bright purple to pinkish violet flower heads (Holm et al., 1997).

                                        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

                                        Balick MJ, Nee M, Atha DE, 2000. Checklist of the vascular plants of Belize. Memoirs of the New York Botanical Garden, 85:1-246.

                                        Cardenas J, Coulston L, 1967. Weeds: A List of Common and Scientific Names for Brazil, Colombia, Puerto Rico, and Central America.

                                        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

                                        Charles Darwin Foundation, 2008. Database inventory of introduced plant species in the rural and urban zones of Galapagos. Galapagos, Ecuador: Charles Darwin Foundation.

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

                                        Flora of China Editorial Committee, 2012. Flora of China Web. Cambridge, USA: Harvard University Herbaria. http://flora.huh.harvard.edu/china/

                                        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

                                        Forzza RC, Leitman PM, Costa AF, Carvalho Jr AA, et al. , 2012. List of species of the Flora of Brazil (Lista de espécies Flora do Brasil). Rio de Janeiro, Brazil: Rio de Janeiro Botanic Garden. http://floradobrasil.jbrj.gov.br/2012/

                                        Fosberg FR, Stoddart DR, 1994. Flora of the Phoenix Islands, central Pacific. Atoll Research Bulletin, 393:60 pp.

                                        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.

                                        Funk VA, Pruski J, 1996. Asteraceae. Memoirs of the New York Botanical Garden, 78:85-122. [Flora of St. John, US Virgin Islands.]

                                        Gargominy O, Bouchet P, Pascal M, Jaffre T, Tourneu JC, 1996. [English title not available]. (Consequences des introductions d'especes animals et vegetales sur la biodiversite en Nouvelle-Caledonie.) Rev. Ecol. (Terre Vie), 51:375-401.

                                        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. Plants of Saint Lucia. http://www.saintlucianplants.com

                                        Herrera K, Lorence D, Flynn T, Balick MJ, 2010. Checklist of the vascular plants of Pohnpei with local names and uses. Lawai, Hawaii, USA: National Tropical Botanical Garden, 146 pp.

                                        Hokche O, Berry PE, Huber O, 2008. Nuevo Catálogo de la Flora Vascular de Venezuela (New catalogue of the vascular flora of Venezuela). Caracas, Venezuela: Fundación Instituto Botánico de Venezuela, 860 pp.

                                        Holm L, Doll J, Holm E, Pancho J, Herberger J, 1997. World Weeds. Natural Histories and Distribution. New York, USA: John Wiley and Sons, Inc.

                                        Madagascar Catalogue, 2012. Catalogue of the vascular plants of Madagascar. Missouri Botanical Garden. St Louis, USA: Missouri Botanical Garden and Madagascar Research and Conservation Program, Antananarivo. http://www.efloras.org/madagascar

                                        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

                                        Meyer JY, 2007. Rapport de mission sur l'Ile d'Uvea (Wallis & Futuna) du 6 au 17 Novembre 2007: Inventaire preliminaire de la flore vasculaire secondaire ([English title not available]). Papeete, Tahiti: Ministère de l'Education, l'Enseignement Supérieur et la Recherche, 39 pp. http://www.li-an.fr/jyves/Meyer_2007_Rapport_Plantes_Introduites_Wallis.pdf

                                        Orchard AE, 1993. Flora of Australia. Vol. 50, Oceanic islands 2. Canberra: Australian Government Publishing Service.

                                        PIER, 2013. Pacific Islands Ecosystems at Risk. Honolulu, Hawaii, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

                                        Prota4U, 2013. PROTA4U web database. Grubben GJH, Denton OA, eds. Wageningen, Netherlands: Plant Resources of Tropical Africa. http://www.prota4u.org/search.asp

                                        Pruski JF, 2013. Asteraceae. In: Flora Mesoamericana, Volume 5 (2) [ed. by Davidse, G. \Sousa Sánchez, M. \Knapp, S. \Chiang Cabrera, F.]., Mexico: Universidad Nacional Autónoma de México. http://www.tropicos.org/docs/meso/asteraceae.pdf

                                        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

                                        Smith AC, 1991. Flora Vitiensis nova: A new flora of Fiji. Lawai, Kauai, Hawai`i. National Tropical Botanical Garden, Volume 5, 626 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. 76 pp. http://www.hear.org/pier/pdf/niue_report_20041217.pdf

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

                                        Stone BC, 1970. The Flora of Guam. Micronesica, 6:1-659.

                                        Thaman RR, Fosberg FR, Manner HI, Hassall DC, 1994. The flora of Nauru. Atoll Research Bulletin, 392:1-223.

                                        US Fish and Wildlife Service, 2010. In: Schiedea spergulina var. leiopoda (no common name). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 11 pp.

                                        USDA-ARS, 2013. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

                                        Vibrans H, 2009. Malezas de México- Pennisetum purpureum ([English title not available]). http://www.conabio.gob.mx/malezasdemexico/poaceae/pennisetum-purpureum/fichas/ficha.htm

                                        Wagner WL, Herbst DR, Sohmer SH, 1999. Manual of the flowering plants of Hawaii. Revised edition. Honolulu, Hawaii, USA: University of Hawaii Press/Bishop Museum Press. [Bernice P. Bishop Museum special publication.]

                                        Wagner WL, Herbst DR, Weitzman A, Lorence DH, 2013. Flora of Micronesia. Flora of Micronesia (online). National Tropical Botanical Garden and the Smithsonian Institution. http://botany.si.edu/pacificislandbiodiversity/micronesia/index.htm

                                        Whistler WA, 1983. The flora and vegetation of Swains Island. Atoll Research Bulletin, 262:25 pp.

                                        Whistler WA, 1988. Checklist of the weed flora of Western Polynesia. An annotated list of the weed species of Samoa, Tonga, Niue, and Wallis and Futuna, along with the earliest dates of collection and the local names. Technical Paper, South Pacific Commission, No. 194:69 pp.

                                        Whistler WA, 1996. Botanical survey of Diego Garcia, Chagos Archipelago, British Indian Ocean Territory. Isle Botanica (online), 49 pp. http://www.zianet.com/tedmorris/dg/2005NRMP-Appendixe-botanicalsurvey.pdf

                                        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

                                        Anon, 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.

                                        Balick MJ, Nee M, Atha DE, 2000. Checklist of the vascular plants of Belize. In: Memoirs of the New York Botanical Garden, 85 1-246.

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

                                        CABI, Undated a. CABI Compendium: Status as determined by CABI editor. 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

                                        Charles Darwin Foundation, 2008. Database inventory of introduced plant species in the rural and urban zones of Galapagos., Galapagos, Ecuador: Charles Darwin Foundation.

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

                                        Flora of China Editorial Committee, 2012. Flora of China Web., Cambridge, USA: Harvard University Herbaria. http://flora.huh.harvard.edu/china/

                                        Florence J, Chevillotte H, Ollier C, Meyer JY, 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

                                        Forzza RC, Leitman PM, Costa AF, Carvalho Jr AA et al, 2012. List of species of the Flora of Brazil. (Lista de espécies Flora do Brasil)., Rio de Janeiro, Brazil: Rio de Janeiro Botanic Garden. http://floradobrasil.jbrj.gov.br/2012/

                                        Fosberg FR, Stoddart DR, 1994. Flora of the Phoenix Islands, central Pacific. In: Atoll Research Bulletin, 393 60 pp.

                                        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.

                                        Gargominy O, Bouchet P, Pascal M, Jaffre T, Tourneu J C, 1996. The consequences of introducing animal and plant species on the biodiversity of New Caledonia. (Consequences des introductions d'especes animals et vegetales sur la biodiversite en Nouvelle-Caledonie.). Revue d'Ecologie (La Terre et la Vie). 375-401.

                                        Graveson R, 2012. Plants of Saint Lucia., http://www.saintlucianplants.com

                                        Herrera K, Lorence D, Flynn T, Balick MJ, 2010. Checklist of the vascular plants of Pohnpei with local names and uses., Lawai, Hawaii, USA: National Tropical Botanical Garden. 146 pp.

                                        Hokche O, Berry PE, Huber O, 2008. New catalogue of the vascular flora of Venezuela. (Nuevo Catálogo de la Flora Vascular de Venezuela)., Caracas, Venezuela, Fundación Instituto Botánico de Venezuela. 860 pp.

                                        Holm L, Doll J, Holm E, Pancho J, Herberger J, 1997. World weeds: natural histories and distribution. New York, USA: John Wiley and Sons. xv + 1129 pp.

                                        Madagascar Catalogue, 2012. Catalogue of the vascular plants of Madagascar. In: Missouri Botanical Garden, St Louis, USA: Missouri Botanical Garden and Madagascar Research and Conservation Program, Antananarivo. http://www.efloras.org/madagascar

                                        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

                                        Meyer JY, 2007. [English title not available]. (Rapport de mission sur l'Ile d'Uvea (Wallis & Futuna) du 6 au 17 Novembre 2007: Inventaire preliminaire de la flore vasculaire secondaire)., Papeete, Tahiti, Ministère de l'Education, l'Enseignement Supérieur et la Recherche. 39 pp. http://www.li-an.fr/jyves/Meyer_2007_Rapport_Plantes_Introduites_Wallis.pdf

                                        Orchard AE, 1993. Flora of Australia. In: Oceanic islands 2, 50 Canberra, Australian Government Publishing Service.

                                        PIER, 2013. Pacific Islands Ecosystems at Risk., Honolulu, Hawaii, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

                                        Pruski JF, 2013. Asteraceae. In: Flora Mesoamericana, 5 (2) [ed. by Davidse G, Sousa Sánchez M, Knapp S, Chiang Cabrera F]. Mexico: Universidad Nacional Autónoma de México. http://www.tropicos.org/docs/meso/asteraceae.pdf

                                        Smith AC, 1991. Flora Vitiensis nova: A new flora of Fiji., 5 Lawai, Kauai, Hawai`i, National Tropical Botanical Garden. 626 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., 76 pp. http://www.hear.org/pier/pdf/niue_report_20041217.pdf

                                        Stone BC, 1970. The Flora of Guam. In: Micronesica, 6 1-659.

                                        Thaman RR, Fosberg FR, Manner HI, Hassall DC, 1994. The flora of Nauru. In: Atoll Research Bulletin, 392 1-223.

                                        USDA-ARS, 2013. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

                                        Vibrans H, 2009. [English title not available]. (Malezas de México- Pennisetum purpureum)., http://www.conabio.gob.mx/malezasdemexico/poaceae/pennisetum-purpureum/fichas/ficha.htm

                                        Wagner WL, Herbst DR, Sohmer SH, 1999. Manual of the flowering plants of Hawaii. Revised edition., Honolulu, Hawaii, USA: University of Hawaii Press/Bishop Museum Press. [Bernice P. Bishop Museum special publication].

                                        Wagner WL, Herbst DR, Weitzman A, Lorence DH, 2013. Flora of Micronesia. Flora of Micronesia (online)., National Tropical Botanical Garden and the Smithsonian Institution. http://botany.si.edu/pacificislandbiodiversity/micronesia/index.htm

                                        Whistler W A, 1988. Checklist of the weed flora of Western Polynesia. An annotated list of the weed species of Samoa, Tonga, Niue, and Wallis and Futuna, along with the earliest dates of collection and the local names. In: Technical Paper, South Pacific Commission, 69 pp.

                                        Whistler WA, 1983. The flora and vegetation of Swains Island. In: Atoll Research Bulletin, 262 25 pp.

                                        Whistler WA, 1996. Botanical survey of Diego Garcia, Chagos Archipelago, British Indian Ocean Territory., Isle Botanica. 49 pp. http://www.zianet.com/tedmorris/dg/2005NRMP-Appendixe-botanicalsurvey.pdf

                                        Contributors

                                        Top of page

                                        10/01/14 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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