Invasive Species Compendium

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Euphorbia lactea
(mottled spurge)

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Datasheet

Euphorbia lactea (mottled spurge)

Summary

  • Last modified
  • 22 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Euphorbia lactea
  • Preferred Common Name
  • mottled spurge
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • E. lactea has been widely commercialized as an ornamental plant and due to the presence of spines it is also used as a fence/hedge plant. Many cultivars have been developed for the horticultural trade (...

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Pictures

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PictureTitleCaptionCopyright
Euphorbia lactea (mottled spurge); habit. Honolua, Maui, Hawaii, USA. July 2009.
TitleHabit
CaptionEuphorbia lactea (mottled spurge); habit. Honolua, Maui, Hawaii, USA. July 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Euphorbia lactea (mottled spurge); habit. Honolua, Maui, Hawaii, USA. July 2009.
HabitEuphorbia lactea (mottled spurge); habit. Honolua, Maui, Hawaii, USA. July 2009.©Forest Starr & Kim Starr - CC BY 4.0
Euphorbia lactea (mottled spurge); habit, showing spines. Honolua, Maui, Hawaii, USA. July 2009.
TitleHabit
CaptionEuphorbia lactea (mottled spurge); habit, showing spines. Honolua, Maui, Hawaii, USA. July 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Euphorbia lactea (mottled spurge); habit, showing spines. Honolua, Maui, Hawaii, USA. July 2009.
HabitEuphorbia lactea (mottled spurge); habit, showing spines. Honolua, Maui, Hawaii, USA. July 2009.©Forest Starr & Kim Starr - CC BY 4.0
Euphorbia lactea (mottled spurge); habit. Old Kaumalapau Hwy, Lanai, Hawaii, usa. April 2007.
TitleHabit
CaptionEuphorbia lactea (mottled spurge); habit. Old Kaumalapau Hwy, Lanai, Hawaii, usa. April 2007.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Euphorbia lactea (mottled spurge); habit. Old Kaumalapau Hwy, Lanai, Hawaii, usa. April 2007.
HabitEuphorbia lactea (mottled spurge); habit. Old Kaumalapau Hwy, Lanai, Hawaii, usa. April 2007.©Forest Starr & Kim Starr - CC BY 4.0
Euphorbia lactea (mottled spurge); habit, as an ornamental. Kourou, French Guiana. October 2006.
TitleHabit
CaptionEuphorbia lactea (mottled spurge); habit, as an ornamental. Kourou, French Guiana. October 2006.
Copyright©Arria Belli/via wikipedia - CC BY-SA 3.0
Euphorbia lactea (mottled spurge); habit, as an ornamental. Kourou, French Guiana. October 2006.
HabitEuphorbia lactea (mottled spurge); habit, as an ornamental. Kourou, French Guiana. October 2006.©Arria Belli/via wikipedia - CC BY-SA 3.0
Euphorbia lactea (mottled spurge); habit, as an ornamental. Calle La Calera in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
TitleHabit
CaptionEuphorbia lactea (mottled spurge); habit, as an ornamental. Calle La Calera in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
Copyright©Frank Vincentz/via wikipedia - CC BY-SA 3.0
Euphorbia lactea (mottled spurge); habit, as an ornamental. Calle La Calera in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
HabitEuphorbia lactea (mottled spurge); habit, as an ornamental. Calle La Calera in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.©Frank Vincentz/via wikipedia - CC BY-SA 3.0
Euphorbia lactea (mottled spurge); habit, as an ornamental. Calle del Berrugo in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
TitleHabit
CaptionEuphorbia lactea (mottled spurge); habit, as an ornamental. Calle del Berrugo in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
Copyright©Frank Vincentz/via wikipedia - CC BY-SA 3.0
Euphorbia lactea (mottled spurge); habit, as an ornamental. Calle del Berrugo in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
HabitEuphorbia lactea (mottled spurge); habit, as an ornamental. Calle del Berrugo in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.©Frank Vincentz/via wikipedia - CC BY-SA 3.0
Euphorbia lactea (mottled spurge); habit, as an ornamental. Calle La Calera in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
TitleHabit
CaptionEuphorbia lactea (mottled spurge); habit, as an ornamental. Calle La Calera in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
Copyright©Frank Vincentz/via wikipedia - CC BY-SA 3.0
Euphorbia lactea (mottled spurge); habit, as an ornamental. Calle La Calera in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.
HabitEuphorbia lactea (mottled spurge); habit, as an ornamental. Calle La Calera in Playa Blanca, Yaiza, Lanzarote, Canary Islands. March 2011.©Frank Vincentz/via wikipedia - CC BY-SA 3.0

Identity

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Preferred Scientific Name

  • Euphorbia lactea Haw.

Preferred Common Name

  • mottled spurge

International Common Names

  • English: candelabra spurge; candelabra-cactus; dragon-bones; elkhorn; false cactus; frilled fan; hat-rack-cactus; mottled candlestick
  • Spanish: candelabro; candelero; cardon

Local Common Names

  • Colombia: lechero de lindero
  • Cuba: cardón; cruz de caravaca; escardon; tuna de cruz
  • Dominican Republic: cacto; candelabro; candelero; raqueta; reaqeta
  • Puerto Rico: escambrón; moteado
  • Sweden: Marmoreuforbia
  • United States Virgin Islands: euphorbia; Malayan spurge-tree; monkey- puzzle
  • USA: candelabra spurge; candelabra-cactus

Summary of Invasiveness

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E. lactea has been widely commercialized as an ornamental plant and due to the presence of spines it is also used as a fence/hedge plant. Many cultivars have been developed for the horticultural trade (USDA-ARS, 2016). It has escaped from cultivation and once naturalized, it often grows forming thickets mostly in disturbed sites, abandoned gardens, deciduous forests, coastal forests, and along roadsides (Little et al., 1974; PIER, 2016; PROTA, 2016). E. lactea spreads by seeds and vegetatively by cuttings and stem fragments (Little et al., 1974). Currently, this species is listed as invasive in Hawaii and Cuba (Oviedo Prieto et al., 2012; PIER, 2016). In Puerto Rico and the Virgin Islands, it is spreading and forming thickets in some places (Little et al., 1974).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Euphorbiales
  •                         Family: Euphorbiaceae
  •                             Genus: Euphorbia
  •                                 Species: Euphorbia lactea

Notes on Taxonomy and Nomenclature

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The family Euphorbiaceae includes around 218 genera and 6745 species with pantropical distribution (Stevens, 2012). Members of this family are mostly monoecious herbs, shrubs, and trees, sometimes succulent and cactus-like plants with xerophytic adaptations (Esser, 2009; Stevens, 2012). The Euphorbiaceae was considered as a morphologically well-defined family for a long time, based mostly on their unisexual flowers and the typical schizocarpous fruits. However, recent molecular studies have resulted in the recognition of several, mostly small families (i.e., Peraceae, Phyllanthaceae, Picrodendraceae, Putranjivaceae). In the most recent circumscription, Euphorbiaceae is defined as clearly monophyletic (Esser, 2009). The genus Euphorbia is among the largest genera of angiosperms, with about 2420 species that are renowned for their remarkably diverse growth forms (i.e., annual to perennial herbs, thorny/spiny stem or succulents to cactus-like and trees (Horn et al., 2012; Stevens, 2012).

Description

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A cactus-like, usually leafless but evergreen shrub or small tree up to 5 metres high and 15-20 cm in trunk diameter, with fleshy or succulent stems, much branched, hairless throughout, producing abundant milky sap when injured. Stems with whorls of branches nearly to base but on large plants shedding the spiny tissue and developing a rounded brown, fissured trunk. The 3-angled (sometimes 4-angled) branches are mostly joints 10-30 cm long and 2-5 cm across, slightly shiny dark green, with yellowish or whitish streak in the groove of the axis between the angles. The soft cut branches have a light green outer layer less than <0.5 cm thick, which yields latex, and within whitish watery tissue, slightly bitter. Raised leaf bases 0.5 cm high and about 1-2.5 cm apart along the edges of branches correspond to nodes and bear paired spreading gray spines (stipules). Leaves few, scattered, alternate, minute, stalkless, rounded, slightly shiny green, succulent, slightly thick and shedding early, or absent. Flowers small, yellowish green, borne intermittently in small clusters along the stem’s edge (Little et al., 1974).

Plant Type

Top of page Perennial
Seed propagated
Shrub
Succulent
Vegetatively propagated

Distribution

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E. lactea is native to Sri Lanka (Govaerts, 2016). It has been introduced in tropical and subtropical areas and now it can be found cultivated and naturalized in tropical Asia, tropical America, the West Indies, and many islands in the Pacific Ocean (see distribution table for details; Acevedo-Rodríguez and Strong, 2012; Govaerts, 2016; PIER, 2016; USDA-ARS, 2016).

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

Asia

BangladeshPresentIntroducedGovaerts (2016)
IndiaPresentIntroducedGovaerts (2016)
Indonesia
-Maluku IslandsPresentNative and IntroducedGovaerts (2016)Listed as both native and introduced
PakistanPresentIntroducedGovaerts (2016)
SingaporePresentIntroducedChong et al. (2009)Cultivated
Sri LankaPresentNativeGovaerts (2016)
ThailandPresentIntroducedGovaerts (2016)

North America

AnguillaPresentIntroducedInvasiveConnor (2008)
BahamasPresentIntroducedAcevedo-Rodríguez and Strong (2012)
British Virgin IslandsPresentIntroducedAcevedo-Rodríguez and Strong (2012)Guana and Tortola
CubaPresentIntroducedInvasiveOviedo Prieto et al. (2012)
CuraçaoPresentIntroducedKairo et al. (2003)aturalized
Dominican RepublicPresentIntroducedAcevedo-Rodríguez and Strong (2012)
Puerto RicoPresentIntroducedAcevedo-Rodríguez and Strong (2012)
U.S. Virgin IslandsPresentIntroducedAcevedo-Rodríguez and Strong (2012)St Thomas, St John
United States
-FloridaPresentIntroducedUSDA-NRCS (2016)
-HawaiiPresentIntroducedInvasiveStaples et al. (2000)

Oceania

Cook IslandsPresentIntroducedMcCormack (2007)Cultivated
Federated States of MicronesiaPresentIntroducedLorence and Flynn (2010); Herrera et al. (2010)Kosrae
French PolynesiaPresentIntroducedFlorence et al. (2013)
Marshall IslandsPresentIntroducedWhistler (1999)Cultivated
PalauPresentIntroducedPIER (2016)Cultivated

South America

ColombiaPresentIntroducedLittle et al. (1974)
EcuadorPresentIntroducedGovaerts (2016)Guayas, Los Rios, Pichincha
-Galapagos IslandsPresentIntroducedCharles Darwin Foundation (2008)
VenezuelaPresentIntroducedAcevedo-Rodríguez and Strong (2012)Margarita Island

History of Introduction and Spread

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E. lactea as well as many of its cultivars have been widely commercialized and introduced into new habitats to be used as an ornamental and fence/hedge plant (USDA-ARS, 2016). However, there is limited information available on the dates of introduction of this species. In the West Indies, original introductions probably occurred around the 1900s. By 1918 it was reported for Bermuda (Britton, 1918) and by 1920, it was listed as widely established in Bahamas and Cuba (Britton and Millspaugh, 1920).

Risk of Introduction

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The risk of introduction of E. lactea is moderate to high. A risk assessment performed for this species on Hawaii scored 8, resulting in a level of “high risk” (PIER, 2016). E. lactea is an attractive succulent cactus-like species widely commercialized as an ornamental and hedge plant. It has the potential to escape from cultivation and colonize new habitats forming dense thickets (Little et al., 1974). Therefore, the likelihood of this species colonizing new areas remains high.

Habitat

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E. lactea grows mostly in dry deciduous forests. It is a common element of dry disturbed sites, waste areas, roadsides, and dry and semiarid environments. It is also an abundant weed of coastal areas and sand dunes along ocean fronts (Little et al., 1974; Kairo et al., 2003; USDA-ARS, 2016).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedDisturbed areas Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Natural
Urban / peri-urban areas Present, no further details Productive/non-natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Principal habitat Natural
Natural grasslands Present, no further details Harmful (pest or invasive)
Natural grasslands Present, no further details Natural
Natural grasslands Present, no further details Productive/non-natural
Scrub / shrublands Present, no further details Harmful (pest or invasive)
Scrub / shrublands Present, no further details Natural
Scrub / shrublands Present, no further details Productive/non-natural
Arid regions Present, no further details Harmful (pest or invasive)
Arid regions Present, no further details Natural
Arid regions Present, no further details Productive/non-natural
Littoral
Coastal areas Present, no further details Harmful (pest or invasive)
Coastal areas Present, no further details Natural
Coastal areas Present, no further details Productive/non-natural

Biology and Ecology

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Genetics

The chromosome number reported for E. lactea is 2n=40. Euphorbia species are extremely variable from the standpoint of chromosome numbers. The chromosome numbers determined are inferred to belong to a primary system of n = 8 and to secondary systems of n= 6, 7, 9 and 10. In this vast genus both aneuploidy and polyploidy have been effective in speciation (Perry, 1943).

Reproductive Biology

The flowers of E. lactea are small and inconspicuous, borne intermittently (Little et al., 1974).

Associations

E. lactea often grows in dry and semiarid environments, associated with xerophytic vegetation such as real cactus species, other Euphorbiaceous species and thorny shrubs (Esser, 2009).

Environmental Requirements

E. lactea prefers to grow on partial to full sunlight exposures, on sandy and sandy-loam soils with pH ranging from 6.1 to 7.8. This species is well-adapted to dry regions and water-stressed habitats and has succulent stems and water storage tissues. Like cacti, Euphorbia species have developed similar compact forms with reduced surface and less water loss, also green stems replacing leaves (Little et al., 1974; Staples et al., 2000; PIER, 2016; USDA-ARS, 2016).

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])
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
BW - Desert climate Tolerated < 430mm annual precipitation

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 17 35

Rainfall

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ParameterLower limitUpper limitDescription
Dry season duration05number of consecutive months with <40 mm rainfall
Mean annual rainfall7502000mm; lower/upper limits

Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • neutral

Soil texture

  • light
  • medium

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Viscum minimum Parasite Whole plant not specific

Notes on Natural Enemies

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The fungus Coniothyrium euphorbiae [Phyllostictella euphorbiae] often causes stem and root rot, principally in plants growing in greenhouses and gardens (Pirone, 1978).

Means of Movement and Dispersal

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E. lactea spreads by seeds and vegetatively by cuttings and stem fragments (Little et al., 1974; PIER, 2016).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Botanical gardens and zoosOften planted to exemplify xerophytic adaptation of plants Yes Yes
DisturbanceNaturalized and spreading on disturbed and waste sties Yes Yes Little et al., 1974
Escape from confinement or garden escapeEscape from gardens and yards Yes Little et al., 1974
Hedges and windbreaksOccasionally planted as a hedge plant Yes Yes Little et al., 1974
HorticultureWidely commercialized as ornamental Yes Yes USDA-ARS, 2016
Ornamental purposesWidely commercialized as ornamental Yes Yes USDA-ARS, 2016

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Debris and waste associated with human activitiesStem fragments and seeds escaped from cultivation Yes Yes Little et al., 1974

Impact Summary

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

Environmental Impact

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E. lactea grows forming dense thickets that displace native vegetation mostly in dry and semiarid habitats. It competes with native plants for water and nutrients. This species also form spiny dense thickets that affect the movement of wildlife in highly invaded areas (Staples et al., 2000; Oviedo Prieto et al., 2012; USDA-ARS, 2016).

Social Impact

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The latex or milky juice of E. lactea is poisonous and caustic for humans, livestock, cats and dogs, as well as to wild animals. It should not come in contact with sores, eyes, mouths, or mucous membranes. Reportedly it injures the eyes and can cause blindness (Little et al., 1974; PROTA, 2016; USDA-ARS, 2016).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside 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
  • Benefits from human association (i.e. it is a human commensal)
  • Long lived
  • Gregarious
  • Reproduces asexually
Impact outcomes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts human health
  • Negatively impacts animal health
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Causes allergic responses
  • Competition - monopolizing resources
  • Competition - smothering
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately

Uses

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E. lactea is widely commercialized as an ornamental plant and due to the presence of spines it is also used as a fence/hedge plant. Many cultivars have been developed and are also commercialized in the horticultural trade (USDA-ARS, 2016). Euphorbia sap is used in India and in Africa to treat warts and in China to treat skin diseases. Latex composition in the genus is very diverse, including toxic compounds as well as other interesting and potentially bioactive molecules such as diterpenes and triterpenes (Ferreira et al. 2002). A study evaluating the anti-inflammatory of “tirucallo”, a tetracyclic triterpene isolated from E. lactea latex showed that the topical application of this compound effectively suppressed ear oedema in mice and exerts a topical anti-inflammatory effect in vivo (Fernandez-Arche et al., 2010).

Uses List

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Environmental

  • Amenity
  • Boundary, barrier or support

General

  • Botanical garden/zoo

Ornamental

  • Seed trade

Similarities to Other Species/Conditions

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Euphorbia species with succulent stems are commonly confused with cacti. However, the Euphorbia produce copious, milky latex, a trait that is absent in cacti. Additionally, flowers of Euphorbias are usually tiny and inconspicuous while cactus flowers are larger and showy.

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.

The herbicide Fluazifop-P-butyl, often used to control grass weeds, can also be used to control E. lactea invasions (EPA, 2003).

References

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Acevedo-Rodríguez P, Strong MT, 2012. Catalogue of the Seed Plants of the West Indies. Smithsonian Contributions to Botany, No. 98. Washington DC, USA: Smithsonian Institution, 1192 pp. http://botany.si.edu/Antilles/WestIndies/catalog.htm

Britton NL, 1918. Flora of Bermuda. New York, USA: Charles Scribner's Sons. 585 pp.

Britton NL, Millspaugh CF, 1920. The Bahama Flora. New York, USA: NL Britton & CF Millspaugh.

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

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. http://lkcnhm.nus.edu.sg/nus/pdf/PUBLICATION/LKCNH%20Museum%20Books/LKCNHM%20Books/flora_of_singapore_tc.pdf

Connor RA, 2008. Anguilla Invasive Species Strategy. Draft. http://www.gov.ai/documents/Anguilla%20Invasive%20Species%20Strategy%202008%20%282%29.pdf

EPA, 2003. U.S. Environmental Protection Agency. EH 951 Grass herbicide Report. http://www3.epa.gov/pesticides/chem_search/ppls/002217-00751-20030917.pdf

Esser HJ, 2009. Neotropical Euphorbiaceae. In: Milliken W, Klitgård B, Baracat A (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. www.kew.org/neotropikey

Fernandez-Arche A, Saenz MT, Arroyo M, Puerta R de la, Garcia MD, 2010. Topical anti-inflammatory effect of tirucallol, a triterpene isolated from Euphorbia lactea late, Phytomedicine, 17(2):146-148

Ferreira AMV, Carvalho LH, Carvalho MJM, Sequeira MM, Silva AM, 2002. Jatrophane and lathyrane diterpenoids from Euphorbia hyberna L., Phytochemistry, 61:373-377

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

Govaerts R, 2016. World Checklist of Euphorbiaceae. Richmond, UK: Royal Botanic Gardens, Kew. http://apps.kew.org/wcsp/

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

Horn JW, Ee BW van, Morawetz JJ, Riina R, Steinmann VW, Berry PE, Wurdack KJ, 2012. Phylogenetics and the evolution of major structural characters in the giant genus Euphorbia L.(Euphorbiaceae)., Molecular Phylogenetics and Evolution, 63:305-326

Kairo M, Ali B, Cheesman O, Haysom K, Murphy S, 2003. Invasive species threats in the Caribbean region. Report to the Nature Conservancy. Curepe, Trinidad and Tobago: CAB International, 132 pp. http://www.issg.org/database/species/reference_files/Kairo%20et%20al,%202003.pdf

Little EL Jr, Woodbury RO, Wadsworth FH, 1974. Trees of Puerto Rico and the Virgin Islands, Vol. 2. Agriculture Handbook No. 449. Washington, DC, USA: USDA, 1024 pp.

Lorence DH, Flynn T, 2010. Checklist of the plants of Kosrae. Unpublished checklist. Lawai, Hawaii: National Tropical Botanical Garden, 26 pp.

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

Oviedo Prieto R, Herrera Oliver P, Caluff MG, et al. , 2012. National list of invasive and potentially invasive plants in the Republic of Cuba - 2011. (Lista nacional de especies de plantas invasoras y potencialmente invasoras en la República de Cuba - 2011). Bissea: Boletín sobre Conservación de Plantas del Jardín Botánico Nacional de Cuba, 6(Special Issue 1):22-96.

Perry BA, 1943. Chromosome number and phylogenetic relationships in the Euphorbiaceae., American Journal of Botany, 30:527-543

PIER, 2016. Pacific Islands Ecosystems at Risk. HEAR, University of Hawaii, Honolulu, USA. http://www.hear.org/pier/index.html

Pirone PP, 1978. Diseases and pests of ornamental plants. : John Wiley & Sons.,: John Wiley & Sons.

PROTA, 2016. PROTA4U web database. Grubben GJH, Denton OA, eds. Wageningen, Netherlands: Plant Resources of Tropical Africa. http://www.prota4u.org/search.asp

Staples GW, Herbst DR, Imada CT, 2000. Survey of invasive or potentially invasive cultivated plants in Hawaii. Bishop Museum Occasional Papers, 65:35 pp. Honolulu, Hawaii, USA: Bishop Museum

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

USDA-ARS, 2016. Germplasm Resources Information Network (GRIN). Online Database. National Germplasm Resources Laboratory, Beltsville, USA. http://www.ars-grin.gov/cgi-bin/npgs/html/tax_search.pl

USDA-NRCS, 2016. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Whistler WA, 1999. Botanical survey of the United States of America Kwajalein Atoll (USAKA) Islands. Prepared for Oak Ridge Institute for Science and Education and the U. S. Army Environmental Center. 111 pp.

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

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

Chong K Y, Tan H T W, Corlett R T, 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. https://lkcnhm.nus.edu.sg/app/uploads/2017/04/flora_of_singapore_tc.pdf

Connor RA, 2008. Anguilla Invasive Species Strategy., http://www.gov.ai/documents/Anguilla%20Invasive%20Species%20Strategy%202008%20%282%29.pdf

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

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Links to Websites

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

Contributors

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19/05/16 Original text by:

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

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