Brachypodium distachyon (purple false brome)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Air Temperature
- Soil Tolerances
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Environmental Impact
- Threatened Species
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Brachypodium distachyon (L.) P. Beauv.
Preferred Common Name
- purple false brome
Other Scientific Names
- Bromus distachyos L.
- Trachynia distachya (L.) Link
International Common Names
- English: annual false brome; false brome; purple false brome; stiff-brome
Local Common Names
- China: er sui duan bing cao
- Finland: aroluste
- France: brachypode à deux épis; brachypode de Pau; brachypode de Phénicie; brachypode des bois; cutandie maritime; vulpie genouillée
- Germany: istrianer zwenke; zweiährige zwenke; zweigrannige zwenke
- Sweden: grusskafting
Summary of InvasivenessTop of page
B. distachyon, commonly known as purple false brome, is a grass species that is related to the major cereal grain species. It is native to southern Europe, northern Africa and southwestern Asia east to India but is widely introduced and naturalized elsewhere. It occurs broadly as an alien species throughout North America and has been reported as invasive in China, Chile, Australia and California. Brachypodium species are invasive weeds that dominate areas where they are planted or have become established and this species can equally form dense stands, reducing diversity and preventing the establishment of native species. The California Invasive Plant Council (Cal-IPC) classifies its potential impact on native ecosystems as ‘Moderate’.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Monocotyledonae
- Order: Cyperales
- Family: Poaceae
- Genus: Brachypodium
- Species: Brachypodium distachyon
Notes on Taxonomy and NomenclatureTop of page
Brachypodium is a genus of about 18 species. About 15 species, among them Brachypodium distachyon, occur naturally in Eurasia, centered in the Mediterranean. Three species occur in the Western Hemisphere, centered in Mexico (Flora of North America Editorial Committee, 2016).
B. distachyon was first described by A.M.F.J. Palisot de Beauvois in 1812. The phylogenetic status of B. distachyon has been full of controversy. B. distachyon is sometimes treated as the only member of Trachynia Link. However, recently scientists have reclassified it into its own tribe Brachypodieae based upon cytological, anatomical and physiological studies (Hasterok et al., 2004; Flora of North America Editorial Committee, 2016; Gramene, 2016).
The Plant List (2016) includes at least 40 synonyms showing its previous inclusion in the genera Agropyron, Brachypodium, Bromus, Festuca, Secale, Trachynia, Triticum anmd Zerna.
Brachypodium is related to the major cereal grain species. But it is understood to be more closely related to the Triticeae (wheat, barley) than to the other cereals (Gramene, 2016).
Brachy is Greek for "short" and podion is Greek for "little foot". "Short footed" refers to the small pedicels of the spikelets (Watson and Dallwitz, 2016).
DescriptionTop of page
Bright green or greenish-blue leaves; leaf-sheaths with slightly stiffened distinct hairs ascending from the surface. Ligule an eciliate membrane. Leaf-blades 1–12 cm long; 1–6 mm wide; glaucous. Leaf-blade surface with slightly stiffened distinct hairs ascending from the surface; sparsely hairy.
Inflorescence composed of racemes. Racemes 1; single; bilateral; 2–8(–10) cm long; bearing 1–7 fertile spikelets on each. Spikelet packing broadside to rhachis. Spikelets solitary. Fertile spikelets pedicelled. Pedicels oblong; 0.5–1 mm long.
Spikelets comprising 5–16 fertile florets; with diminished florets at the apex. Spikelets oblong; laterally compressed; 10–30 mm long; breaking up at maturity; disarticulating below each fertile floret.
Glumes persistent; dissimilar; shorter than spikelet. Lower glume lanceolate; 5–6 mm long; 0.75 length of upper glume; coriaceous; without keels; 5–7 -veined. Lower glume lateral veins ribbed. Lower glume apex acute. Upper glume oblong; 7–8 mm long; 0.75 length of adjacent fertile lemma; coriaceous; without keels; 7 -veined. Upper glume lateral veins ribbed. Upper glume apex acute.
Fertile lemma oblong; laterally compressed; 8–10 mm long; coriaceous; glaucous; without keel; 7 -veined. Lemma surface glabrous, or pilose. Lemma margins ciliolate; hairy above. Lemma apex acute; awned; 1 -awned. Principal lemma awn 8–12 mm long overall. Apical sterile florets resembling fertile though underdeveloped.
Lodicules 2; ciliate. Anthers 3; 0.5–1 mm long. Ovary pubescent on apex.
Plant TypeTop of page Annual
Grass / sedge
DistributionTop of page
B. distachyon is native to the Madeira Islands, Canary Islands and northern Africa: Ethiopia; Algeria; Egypt; Libya; Morocco and Tunisia; southwestern Asia east to Pakistan: Azerbaijan; Russian Federation (Dagestan); Turkmenistan; Uzbekistan; Afghanistan; Cyprus; Egypt (Sinai); Iran; Iraq; Israel; Jordan; Lebanon; Syria; Turkey and Pakistan; and east and southern Europe: Ukraine; Albania; Bulgaria; Croatia; Greece; Italy; Slovenia; France; Portugal and Spain. It is uncertain whether it’s native to the Azores, Portugal (DAISIE, 2016; GBIF, 2016; USDA-ARS, 2016).
This species is widely introduced and naturalized elsewhere (USDA-ARS, 2016). It occurs broadly as an alien species throughout North America and has specifically been reported from Texas, Colorado, California, Oregon, Hawai’i and New Jersey (USDA-NRCS, 2016). It has been reported as invasive in China, Chile (offshore islands), Australia (New South Wales) (PIER, 2016) and in California, USA (Calflora, 2016).
Distribution TableTop of page
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|
|Egypt||Present||Native||USDA-ARS (2016); Encyclopedia of Life (2016)||Sinaï|
|Belgium||Absent, Eradicated||DAISIE (2016)|
|-Azores||Absent, Unconfirmed presence record(s)||USDA-ARS (2016); DAISIE (2016)|
|Russia||Present||CABI (Undated)||Present based on regional distribution.|
|-Southern Russia||Present||Native||USDA-ARS (2016)||Dagestan|
|-Canary Islands||Present||Native||USDA-ARS (2016)|
|United Kingdom||Present||Introduced||DAISIE (2016)|
|United States||Present||Introduced||PIER (2016)|
|-California||Present||Introduced||Invasive||Calflora (2016); GBIF (2016)|
|-Hawaii||Present||Introduced||ITIS (2016); USDA-NRCS (2016)|
|Australia||Present||CABI (Undated)||Present based on regional distribution.|
|-New South Wales||Present||Introduced||Invasive||PIER (2016)||Uncommon, in disturbed areas|
|Chile||Present||Introduced||Invasive||PIER (2016)||Isla Más Afuera (Alejandro Selkirk Island)|
Risk of IntroductionTop of page
The California Invasive Plant Council (Cal-IPC) classifies its potential impact on native ecosystems as ‘Moderate’, meaning that “the species has substantial and apparent, but generally not severe, ecological impacts on physical processes, plant and animal communities, and vegetation structure. Their reproductive biology and other attributes are conducive to moderate to high rates of dispersal, though establishment is generally dependent upon ecological disturbance. Ecological amplitude and distribution may range from limited to widespread”. Based on impact, invasiveness and distribution, Cal-IPC (2016) has given B. distachyon the alert status: ‘No Alert’. There is significant risk of introduction of B. distachyon as a contaminant of cereal or grass seeds and carried in wool. Deliberate introduction can also occur when it is used as a research tool.
HabitatTop of page
This species is native to dry, open habitats where it grows on sandy or rocky soils. Where it is introduced and naturalized, it is often found in disturbed areas (Flora of North America Editorial Committee, 2016). The Encyclopedia of Life (2016) also mentions forest edges, abandoned plantations, beaches, coastal dunes, orchards and waste places as habitats where B. distachyon can be found.
In California it is reported to invade undisturbed/disturbed grasslands, dry slopes and fields, roadsides, margins of shrub thickets and oak woodlands. It tolerates partial shade in oak woodlands (Cal-IPC, 2016).
In Australia it grows in dry, disturbed areas on sandy or rocky soils (Flora of North America Editorial Committee, 2016).
Habitat ListTop of page
|Terrestrial – Managed||Managed forests, plantations and orchards||Present, no further details|
|Disturbed areas||Present, no further details|
|Rail / roadsides||Present, no further details|
|Terrestrial ‑ Natural / Semi-natural||Natural forests||Present, no further details|
|Natural grasslands||Present, no further details|
|Arid regions||Present, no further details|
|Coastal dunes||Present, no further details|
Biology and EcologyTop of page
Chromosome number: 5; 2n = 2x = 10, 4x = 20, 6x = 30 polyploidy (Gramene, 2016).
B. distachyon is a winter annual that reproduces only by seed. It has a rapid life cycle, it reaches reproductive maturity within eight to twelve weeks (Li et al., 2012). Bloom period is from April to July. This species is self compatible (Cal-IPC, 2016). Seed longevity in the soil is expected to last two years (DiTomaso et al., 2013). Germination is variable within the species. There may be varying duration of after-ripening requirement and response to light; in general, light tended to reduce germination rather than enhance it (Barrero et al., 2011)..
Physiology and Phenology
B. disatachyon has C3 physiology. Aronson et al. (1992) have compared the dynamics of vegetative and reproductive growth in matched pairs of Mediteranean and desert populations of three unrelated annual species, among them B. distachyon, under high and low levels of water availability in a common-environment experiment. B. distachyon showed earlier switches to reproductive development and to subsequent phenophases, and the transition to flowering occurred at smaller plant sizes. Water stress slightly accelerated the transition to flowering in B. distachyon by 4–6 days. The duration of life cycle was greatly shortened by water stress.
The experiment also showed that the Mediterranean population of B. distachyon was much more plastic in its response to water stress than the desert population in its transition to plant ageing. It appeared that plants from the desert populations were adapted to shorter, more compact growth cycles, culminating in earlier dates of seed maturation and plant ageing. They also showed larger phenotypic plasticity in the transition to plant ageing, which trait was enhanced or magnified by sustained or repeated lack of water. By contrast, plants from Mediterranean populations delayed switchover from one phenophase to the next, seeming to “bet” on more water in the near future (Aronson et al., 1992).
B. distachyon is a winter annual but seeds may persist in the soil for at least 2 years (DiTomaso et al., 2013).
This species tolerates arid environments with annual rainfalls between 279 mm and 1702 mm (Calflora, 2016). According to Cal-IPC (2016), it can be found at elevations up to 600 m. Calflora (2016) reports on altitudes from 4 m up to 950 m.
B. distachyon grows on non-saline soils with a pH of 3.4 to 7.6. It tolerates a very low concentrate of CaCO3 (3%). The grass requires a minimum depth of 20 cm and grows on fine, medium and coarse soil textures (Calflora, 2016).
ClimateTop of page
|BS - Steppe climate||Preferred||> 430mm and < 860mm annual precipitation|
|BW - Desert climate||Preferred||< 430mm annual precipitation|
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Preferred||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
Air TemperatureTop of page
|Parameter||Lower limit||Upper limit|
|Mean maximum temperature of hottest month (ºC)||37|
|Mean minimum temperature of coldest month (ºC)||1|
RainfallTop of page
|Parameter||Lower limit||Upper limit||Description|
|Mean annual rainfall||279||1702||mm; lower/upper limits|
Soil TolerancesTop of page
- very acid
Means of Movement and DispersalTop of page
This species is spread by water (Cal-IPC, 2016).
Vector Transmission (Biotic)
Florets fall near the plant and can be dispersed by animals (Cal-IPC, 2016).
The florets of B. distachyon can also be dispersed by vehicle tyres and human activities, especially in contaminated hay (Cal-IPC, 2016).
Researchers use this species as a model for the grasses used to produce biomass, food, feed and forage (USDA-JGI, 2016).
Pathway CausesTop of page
Pathway VectorsTop of page
|Germplasm||Used as a model for the grasses used to produce biomass, food, feed and forage||Yes||USDA-JGI, 2016|
|Land vehicles||Florets can be dispersed by vehicle tyres||Yes||Cal-IPC, California Invasive Plant Council|
|Water||B. distachyon is spread by waterways||Yes||Cal-IPC, California Invasive Plant Council|
Impact SummaryTop of page
Environmental ImpactTop of page
Brachypodium genera are invasive weeds that dominate areas where they grow (USDA Forest Service, 2003). In the USA, B. distachyon is a weed of urban bushland and of the western edge of the wheatbelt. It is specifically reported from California, Oregon, Texas, Colorado, Hawaii and New Jersey. Generally, it is under-reported, probably because of superficial similarity to Bromus species (Hussey et al, 1997; Gramene, 2016). B. distachyon is locally abundant in California. It seems to still be spreading in California, but probably at a relatively slow rate statewide. Regionally it can spread rapidly (Cal-IPC, 2016).
Impact on Habitats
B. distachyon can form dense stands that can change fire regimes and build up thick litter layers (Cal-IPC, 2016). An increase in woody vegetation and nonnative plant species in some habitats has resulted in less available prairie habitat overall.
Impact on Biodiversity
Cal-IPC (2016) evaluated the impact of B. distachyon on plant community composition, structure, and interactions as ‘Moderate’. Their reproductive biology and other attributes are conducive to moderate to high rates of dispersal, though establishment is generally dependent upon ecological disturbance. Ecological amplitude and distribution may range from limited to widespread. It can form dense stands in some locations, particularly in oak woodlands. These near monotypic stands can reduce diversity and prevent native species from establishing. The impact on higher trophic levels is evaluated as ‘Moderate’. The fibrous stems make it a poor forage for animals. The long awns can also injure animals (Cal-IPC, 2016).
It is recorded, among others, as being one of the most prominent nonnative species threatening the endangered Acanthomintha ilicifolia, competing for nutrients, light, water and space (US Fish and Wildlife Service, 2009). It is also noted as an invasive of concern to the protection of the endangered species Sibara filifolia (US Fish and Wildlife Service, 2006).
B. distachyon can contribute to the loss of prairie habitat, and subsequently lead to habitat that is avoided by some butterflies and birds, such as the Taylor’s checkerspot butterflies (Euphydryas editha taylori) and streaked horned larks (Eremophila alpestris strigata) (US Fish and Wildlife Service, 2013).
Based on impact, invasiveness and distribution, Cal-IPC (2016) has given B. distachyon the alert status: ‘No Alert’.
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Acanthomintha ilicifolia||NatureServe; USA ESA listing as threatened species||California||Competition - monopolizing resources; Competition - smothering||US Fish and Wildlife Service, 2009|
|Eremophila alpestris strigata (streaked horned lark)||USA ESA listing as threatened species||California||US Fish and Wildlife Service, 2013||Ecosystem change/ habitat alteration|
|Euphydryas editha taylori||National list(s)||California||US Fish and Wildlife Service, 2013||Ecosystem change/ habitat alteration|
|Sibara filifolia (Santra Cruz Island Rockcress)||USA ESA listing as endangered species||California||Competition - monopolizing resources; Competition - smothering||US Fish and Wildlife Service, 2006|
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Has a broad native range
- Pioneering in disturbed areas
- Tolerant of shade
- Fast growing
- Ecosystem change/ habitat alteration
- Modification of fire regime
- Modification of successional patterns
- Monoculture formation
- Reduced native biodiversity
- Threat to/ loss of native species
- Competition - monopolizing resources
- Competition - smothering
- Highly likely to be transported internationally deliberately
- Difficult to identify/detect as a commodity contaminant
UsesTop of page
B. distachyon is used by researchers as a model for the grasses used to produce biomass, food, feed and forage. The species has many qualities that make it an excellent model organism for functional genomics research in temperate grasses, cereals, and dedicated biofuel crops. These attributes include the relatively small size of its genome which makes it useful for genetic mapping and sequencing. It has diploid accessions and a series of polyploid accessions. It is small sized, has a short lifecycle and it is self compatible. As a weed it has simple growth requirements (USDA-JGI, 2016).
Studies in B. distachyon have provided new insight into the structure and physiology of plant cell walls, the development and chemical composition of endosperm, and the genetic basis for cold tolerance. Recent work on auxin transport has also uncovered mechanisms that apply to all angiosperms other than Arabidopsis (Kellogg, 2015).
The species is a poor forage grass for livestock (DiTomaso et al., 2013).
Uses ListTop of page
- Research model
Similarities to Other Species/ConditionsTop of page
B. distachyon can be distinguished from other species of Brachypodium by having shorter pedicels and anthers, laterally compressed spikelets, and fewer spikelets per raceme. This species is the only annual member of the Brachypodieae and its flowers are self-pollinating and set fertile seed without the flower opening (Flora of North America Editorial Committee, 2016; Gramene, 2016). In the UK it is distinguished from native Brachypodium species by its annual habit (Stace, 1991).
It can be confused with Bromus hordeaceus (soft brome), an annual grass which is hairy all over, and has rounder, hairier, softer seedheads (Cal-IPC, 2016).
Prevention and ControlTop of page
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.
Cultural Control and Sanitary Measures
Prescribed burning in early summer has successfully reduced the size of populations. It is a poor forage grass and so grazing is unlikely to be a useful control measure (DiTomaso et al., 2013).
Hand pulling, mowing or tillage may be suitable for small infestations. Mowing should be undertaken before viable seed production but after most of the soil moisture has been depleted to prevent regrowth (DiTomaso et al., 2013).
In California B. distachyon has reportedly been controlled by the grass-specific herbicide fluazifop-p-butyl and by the non-selective glyphosate (Conservation Biology Institute, 2016). No other specific chemicals are reported, although it is likely that similar herbicides used to control other grasses may be relevant for this species (DiTomaso et al., 2013). In Israel it is reported to be resistant to triazine herbicides (Regev and Kleifeld, 1982).
ReferencesTop of page
Barrero JM, Jacobsen JV, Talbot MJ, White RG, Swain SM, Garvin DF, Gubler F, 2011. Grain dormancy and light quality effects on germination in the model grass Brachypodium distachyon. New Phytologist, 193(2):376-386.
Calflora, 2016. Information on California plants for education, research, and conservation. Berkeley, California, USA: Calflora Database. http://www.calflora.org
Cal-IPC (California Invasive Plant Council), 2016. California Invasive Plants Council. Berkeley, California, USA: California Invasive Plant Council. http://www.cal-ipc.org/
Clayton WD, Vorontsova MS, Harman KT, Williamson H, 2016. GrassBase - The Online World Grass Flora. http://www.kew.org/data/grasses-db.html
Conservation Biology Institute, 2016. Brachypodium Control: Experimental Treatments to Control Brachypodium - An Adaptive Approach for Conserving Endemic Species, San Diego County, California. California, USA: Conservation Biology Institute. http://consbio.org/products/reports/brachypodium-control
DAISIE, 2016. Delivering Alien Invasive Species Inventories for Europe. European Invasive Alien Species Gateway. www.europe-aliens.org/default.do
DiTomaso JM, Kyser GB, Oneto SR, Wilson RG, Orloff SB, Anderson LW, Wright SD, Roncoroni JA, Miller TL, Prather TS, Ransom C, Beck KG, Duncan C, Wilson KA, Mann JJ, 2013. Weed Control in Natural Areas in the Western United States. Davis, California, USA: Weed Research and Information Center, University of California, 544 pp.
Encyclopedia of Life, 2016. Encyclopedia of Life. http://www.eol.org
Flora of North America Editorial Committee, 2016. Flora of North America North of Mexico. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=1
GBIF, 2016. Global Biodiversity Information Facility. http://www.gbif.org/species
Gramene, 2016. The Gramene database. Release #39. http://www.gramene.org/
ITIS, 2016. Integrated Taxonomic Information System online database. http://www.itis.gov
Kaye T, 2003. Invasive Plant Alert False-brome (Brachypodium sylvaticum). Oregon, USA: Institute for Applied Ecology, 2 pp. http://appliedeco.org/wp-content/uploads/brsybrochure.pdf
Li ChuAn, Rudi H, Stockinger EJ, Cheng HongMei, Cao Moju, Fox SE, Mockler TC, Westereng B, Fjellheim S, Rognli OA, Sandve SR, 2012. Comparative analyses reveal potential uses of Brachypodium distachyon as a model for cold stress responses in temperate grasses. BMC Plant Biology, 12(65):(8 May 2012). http://www.biomedcentral.com/content/pdf/1471-2229-12-65.pdf
PIER, 2016. Pacific Island Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html
The Plant List, 2013. The Plant List: a working list of all plant species. Version 1.1. London, UK: Royal Botanic Gardens, Kew. http://www.theplantlist.org
US Fish and Wildlife Service, 2006. 5-Year Review: Summary and Evaluation: Species Reviewed: Sibara filifolia. California, USA: US Fish and Wildlife Service.
US Fish and Wildlife Service, 2009. 5-Year Review: Summary and Evaluation: Species Reviewed: Acanthomintha ilicifolia. California, USA: US Fish and Wildlife Service.
US Fish and Wildlife Service, 2009. In: Acanthomintha ilicifolia (San Diego thornmint). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 39 pp.. http://ecos.fws.gov/docs/five_year_review/doc2571.pdf
US Fish and Wildlife Service, 2013. Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for the Taylor's Checkerspot Butterfly and Threatened Status for the Streaked Horned Lark. Federal Register, 78(192). Washington DC, USA: Department of the Interior, 53.
USDA-ARS, 2016. Germplasm Resources Information Network (GRIN). National Plant Germplasm System. Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx
USDA-JGI, 2016. Brachypodium genome resources project. Joint Genome Institute. Berkeley, USA: Joint Genome Institute, Lawrence Berkeley National Laboratory.
USDA-NRCS, 2016. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/
Watson L, Dallwitz MJ, 2016. The grass genera of the world: descriptions, illustrations, identification, and information retrieval; including synonyms, morphology, anatomy, physiology, phytochemistry, cytology, classification, pathogens, world and local distribution, and references. Institute of Botany, Chinese Academy of Sciences. http://delta-intkey.com
CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
Calflora, 2016. Information on California plants for education, research, and conservation., Berkeley, California, USA: Calflora Database. http://www.calflora.org
DAISIE, 2016. Delivering Alien Invasive Species Inventories for Europe. http://www.europe-aliens.org/
Encyclopedia of Life, 2016. Encyclopedia of Life., http://www.eol.org
GBIF, 2016. Global Biodiversity Information Facility. http://www.gbif.org/species
ITIS, 2016. Integrated Taxonomic Information System online database., http://www.itis.gov
PIER, 2016. Pacific Island Ecosystems at Risk., Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html
USDA-ARS, 2016. 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, 2016. The PLANTS Database. Greensboro, North Carolina, USA: National Plant Data Team. https://plants.sc.egov.usda.gov
ContributorsTop of page
02/03/16 Original text by:
Diana Quiroz, Naturalis Biodiversity Center, Netherlands
Distribution MapsTop of page
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