Lawsonia inermis
(Egyptian privet)

Pictures

Picture	Title	Caption	Copyright
Title Inflorescence Caption Copyright ©A.R. Pittaway	Inflorescence		©A.R. Pittaway
Title Line artwork Caption 1. lower (to left) and upper (to right) part of flowering branch 2. flower 3. fruit Copyright PROSEA Foundation	Line artwork	1. lower (to left) and upper (to right) part of flowering branch 2. flower 3. fruit	PROSEA Foundation

Identity

Preferred Scientific Name

• Lawsonia inermis L.

Preferred Common Name

• Egyptian privet

Other Scientific Names

• Alcanna spinosa (L.) Gaertn.
• Casearia multiflora Spreng.
• Lawsonia alba Lam.
• Lawsonia purpurea Lam.
• Lawsonia speciosa L.
• Lawsonia spinosa L.
• Rotantha combretoides Baker

International Common Names

• English: camphire; henna; henna tree; mignonette tree
• Spanish: alcana; alhena; jenna; reseda
• French: henne; henné
• Arabic: hinná; yoranna
• Portuguese: alfeneiro; hena; hésia

Local Common Names

• Cambodia: krâpéén
• Dominican Republic: resedón
• Germany: Hennastrauch
• India: hena; marithoni; maruthani; mehedi; mendhi; mendi
• Indonesia: inai
• Indonesia/Java: pacar kuku
• Italy: alcanna vera; arbusto della henna
• Laos: kaaw
• Lesser Antilles: copiae
• Malaysia: hinna; inai; pacar kuku
• Myanmar: dan
• Pakistan: hena; mendhi
• Philippines: cinamomo
• Vietnam: lá mòn; nhuôm móng

EPPO code

• LAWIN (Lawsonia inermis)

Trade name

• henna
• mendhi

Summary of Invasiveness

L. inermis is a shrub or small tree widely cultivated as an ornamental and hedge plant and for the commercial production of henna, a dye extracted from its leaves (Oyen, 1991; Orwa et al., 2009; PROTA, 2017; USDA-ARS, 2017). This species has often become naturalized, particularly on alluvial soils along rivers. In some African regions (e.g., Madagascar) this species has become so common that there is no need for cultivation (PROTA, 2017). It is listed as invasive in Cuba (Oviedo Prieto et al., 2012) and Singapore (Chong et al., 2009).

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Myrtales
•                         Family: Lythraceae
•                             Genus: Lawsonia - (Withdrawn)
•                                 Species: Lawsonia inermis

Notes on Taxonomy and Nomenclature

The family Lythraceae comprises 31 genera and 620 species of herbs, shrubs and trees that are recognizable by their flaky bark; opposite, entire leaves that lack gland dots; ovary is more or less superior. The fruit is usually a capsule; the testa of about half the genera has distinctive, initially inverted mucilaginous hairs (Stevens, 2012). Lawsonia is a small genus of just two species: Lawsonia inermis and Lawsonia odorata (The Plant List, 2013). The specific epithet inermis means unarmed or without spines (Orwa et al., 2009).

Description

L. inermis is a much-branched, glabrous shrub or small tree, 2-6 m tall, with greyish-brown bark, unarmed when young, older plants with spine-tipped branchlets. Young branches quadrangular. Leaves opposite, entire and sub-sessile, elliptic to broadly lanceolate, 1.5-5 cm x 0.5-2 cm. Flowers numerous in large, pyramidal, terminal cymes, fragrant, 1 cm across and 4-merous, calyx with 2 mm long tube, and 3 mm long spreading lobes; petals orbicular or obovate, white or red; stamens 8, inserted in pairs on the rim of the calyx tube; ovary 4-celled, style up to 5 mm long, erect. Fruit a globose capsule, 4-8 mm in diameter, many-seeded, opening irregularly. Seeds 3 mm long, angular, with thick coat (Oyen, 1991). 

Plant Type

Distribution

The native distribution range of L. inermis is uncertain, but this species probably originates in Asia (USDA-ARS, 2017). Linguistic and biogeographic evidence supports an origin in the area of Baluchistan (Iran/Pakistan) to western India, where it can still be found growing in the wild (PROTA, 2017). From there it would have spread to the rest of India, Indonesia, the Sahel, Central Africa, and the Middle East. L. inermis is now distributed throughout tropical and subtropical regions (Oyen, 1991; Orwa et al., 2009; PROTA, 2017). Both USDA-ARS (2017) and PROTA (2017) suggest that the origin is likely to be South Asia, but also include much of Africa and parts of Asia and the Middle East in the listed native distribution.

Distribution Table

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.

History of Introduction and Spread

L. inermis occurs wild from Iran and Pakistan to western India. From there it would have spread eastward to the rest of India and Indonesia, and westward to the Middle East where it became one of the important plants of Islam. Aronson et al. (2017) suggest that it was introduced to the Middle East 1500-2500 years ago. Later, Islamic armies and traders from Arabia introduced this species in areas as far as Spain, northern Africa, Madagascar, the Moluccas, Indo-China and Japan (PROTA, 2017). Currently, this species can be found throughout the tropics and subtropics. It is often grown in home gardens and commercial production is limited to a few places in India, Pakistan, Iran, Egypt, Libya, Niger and Sudan (Oyen, 1991; PROTA, 2017).

The earliest known use of the dye (henna) prepared from L. inermis dated to the Ancient Egypt. Egyptians used this plant to stain mummies and mummy’s wrappings. Later, between 1400-1500 BC, Greeks probably moved this species from the Middle East and North Africa to the Mediterranean islands and Europe. In classical Greece, henna was well known as a medicinal and cosmetic plant (Zavada, 1993).

Risk of Introduction

The likelihood of continued introduction of L. inermis is moderate to high. This species has been widely introduced across tropical and subtropical regions to be used as an ornamental and for henna dye production (USDA-ARS, 2017). The international demand for henna dye is still growing, and consequently new introductions are highly likely to occur (Oyen, 1991).

Habitat

L. inermis is well adapted to a wide range of environmental conditions. It grows in dry and semi-arid habitats, in coastal secondary scrubs, along watercourses and in temporarily flooded riverbeds and riverine thickets (Orwa et al., 2009; PROTA, 2017). In Africa it has often become naturalized, particularly on alluvial soils along rivers and in areas near villages (PROTA, 2017). In Central America and Mexico, it can also be naturalized in secondary vegetation (Davidse et al., 2009).

Biology and Ecology

Genetics

The chromosome number reported for L. inermis is 2n = 30 (Oyen, 1991; Hanson et al., 2001).

Reproductive Biology

L. inermis is listed as a “self-pollinated” plant (Phirke and Saha, 2013).

Physiology and Phenology

L. inermis produces flowers all year round (PROTA, 2017).

Longevity

L. inermis is a perennial shrub or small tree. In cultivation, while highest yields occur at 4-8 years after planting, plants are often left in the field for 12-25 (max. 40) years (Orwa et al., 2009).

Activity Patterns

L. inermis requires high temperatures (~25°C) for germination, growth and establishment (PROTA, 2017).

Environmental Requirements

L. inermis grows best in areas where mean annual temperatures are within the range 19-27°C (but can tolerate 13-33°C) and mean annual rainfall ranging from 500 mm to 1500 mm, but can tolerate ranges varies form 200 mm to 4200 mm. It can grow on poor, stony, and sandy soils, but it is also adapted to heavy, fertile clay soils with pH ranging from 4.3 to 8. This species requires high temperatures (optimum daily average about 25°C) for germination, growth and development. L. inermis tolerates drought and semiarid conditions. It often grows in temporarily flooded, riverbeds and riverine thickets at elevations up to 2000 m (Oyen, 1991; Orwa et al., 2009; FAO, 2017; PROTA, 2017).

Latitude/Altitude Ranges

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free
• impeded

Soil reaction

• acid
• alkaline
• neutral

Soil texture

• heavy
• light
• medium

Special soil tolerances

• infertile
• saline
• shallow

Natural enemies

Notes on Natural Enemies

Pests and diseases have been reported attacking L. inermis including: Alternaria alternata, Alternaria tenuissima, the beetle Pachnoda interrupta, Phenacoccus solenopsis (cotton mealybug), the aphid Sarucallis kahawaluokalani and Corticium koleroga. In India, a black root rot caused by Corticium koleroga and a bacterial leaf spot caused by Xanthomonas lawsoniae have been reported on this species (Orwa et al., 2009).

Means of Movement and Dispersal

Birds feed on the fruits of L. inermis and probably disperse the seeds (Orwa et al., 2009).

Intentional Introduction

L. inermis has been widely introduced to be used as ornamental and hedge plants, and for the commercial production of henna, a dye extracted from its leaves (Oyen, 1991; Orwa et al., 2009; PROTA, 2017; USDA-ARS, 2017).

Pathway Causes

Pathway Vectors

Environmental Impact

Impact on habitats

L. inermis removes a large quantity of nutrients from the soil. For example, a yield of 1000 kg dry leaves removes about 180-190 kg of nitrogen, 100-150 kg of potassium and 10-30 kg of phosphate (Orwa et al., 2009).

Impact on biodiversity

L. inermis can grow forming dense thickets which are able to cause environmental degradation by out-competing and replacing native vegetation, mostly native vegetation on alluvial soils and in riparian zones (PROTA, 2017).

Social Impact

The henna dye is known to be dangerous to people with glucose-6-phosphate dehydrogenase deficiency and may cause allergic reaction and injuries to the skin (FDA, 2017).

Risk and Impact Factors

• Proved invasive outside its 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
• Benefits from human association (i.e. it is a human commensal)
• Long lived
• Fast growing
• Gregarious

• Altered trophic level
• Ecosystem change/ habitat alteration
• Monoculture formation
• Negatively impacts human health
• Reduced native biodiversity
• Threat to/ loss of native species

• Causes allergic responses
• Competition - monopolizing resources
• Competition - smothering
• Herbivory/grazing/browsing
• Rapid growth
• Rooting

• Highly likely to be transported internationally deliberately

Uses

Economic Value

L. inermis is widely cultivated in tropical and subtropical regions of the world. It is often planted as ornamental for its fragrant flowers. It is also cultivated for the production of henna: an orange-red dye obtained from macerated, triturated or powdered leaves. Henna dye is used for dyeing cloth and hair, for staining nails, palms and soles, and in traditional medicine. Leaves are browsed by livestock and the wood is used to make tent pegs and tool handles (Oyen, 1991; Orwa et al., 2009; PROTA, 2017; USDA-ARS, 2017).

Environmental Services

L. inermis has been successfully planted in soil conservation projects (Orwa et al., 2009). It can also be grown as a live fence.

Uses List

Animal feed, fodder, forage

• Fodder/animal feed

Environmental

• Amenity
• Boundary, barrier or support
• Erosion control or dune stabilization
• Revegetation
• Shade and shelter

Fuels

• Fuelwood

Human food and beverage

• Food additive

Materials

• Baskets
• Carved material
• Dye/tanning
• Dyestuffs
• Essential oils
• Fibre
• Pesticide
• Wood/timber

Medicinal, pharmaceutical

• Source of medicine/pharmaceutical
• Traditional/folklore

Ornamental

• Christmas tree
• Cut flower
• garden plant
• Potted plant
• Propagation material
• Seed trade

Wood Products

Woodware

• Industrial and domestic woodware
• Tool handles

References

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

Ahmad MB, Rauf A, Osman SM, 1989. Physio-chemical analysis of seven seed oils. Journal of the Oil Technologists' Association of India, 21(3):46-47; 5 ref.

Ali BH, Bashir AK, Tanira MOM, 1995. Anti-inflammatory, antipyretic, and analgesic effects of Lawsonia inermis L. (henna) in rats. Pharmacology, 51(6):356-363; 11 ref.

Anand N, 1983. The market for annatto and other natural colouring materials, with special reference to the United Kingdom. Report, Tropical Development and Research Institute, UK, No. G174, 32 pp.; 15 ref.

Anttila LS, Alakoski-Johansson GM, Johansson SG, 1993. Browse preference of Orma livestock and chemical composition of Prosopis juliflora and nine indigenous woody species in Bura, Eastern Kenya. East African Agricultural and Forestry Journal, 58(Special Issue):83-90; 28 ref.

Aronson I, Aronson TB, Patzelt A, Knees SG, Lewis GP, Lupton D, Taifour H, Gardner MF, Thompson H, Al-Hatmi S, Al-Khulaidi AW, 2017. Paleorelicts or archaeophytes: enigmatic trees in the Middle East. Journal of Arid Environments, 137: 69-82

Bajrang Singh, Khanduja SD, Srivastava GS, 1984. Qualitative analysis of some firewood shrubs. Biomass, 5(4):317-320; 3 ref.

Broome R, Sabir K, Carrington S, 2007. Plants of the Eastern Caribbean. Online database. Barbados: University of the West Indies. http://ecflora.cavehill.uwi.edu/index.html

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

Datta BK, Amhed M, Banoo R, Talukdar SA, 1989. Antifungal and antibacterial activity of the seed oil of Lawsonia alba Lam. studied in vitro. Bangladesh Journal of Microbiology, 6(2):49-52; 5 ref.

Davidse G, Sousa Sánchez M, Knapp S, Chiang Cabrera F, 2009. Flora Mesoamericana. St. Louis, USA: Missouri Botanical Garden

FAO, 2017. Ecocrop Website. http://ecocrop.fao.org/ecocrop/srv/en/home

FDA, 2007. Temporary Tattoos, Henna/Mehndi, and "Black Henna": Fact Sheet. http://www.fda.gov/MedicalDevices/ProductsandMedicalProcedures/HomeHealthandConsumer/ConsumerProducts/ContactLenses/ucm108569.htm

Fernandes A, 1980. Flora of Mozambique. 75. Sonneratiaceae. [Flora de Mocambique. 75. Sonneratiaceae.] 1980, i + 5 pp.

Flora of China Editorial Committee, 2013. Flora of China. Beijing & St. Louis: Missouri Botanical Garden Press

Funk VA, Berry PE, Alexander S, Hollowell TH, Kelloff CL, 2007. Checklist of the plants of the Guiana Shield. Contributions from the United States National Herbarium, 55: 1–584

Hanson L, McMahon KA, Johnson MAT, Bennett MD, 2001. First nuclear DNA C-values for 25 angiosperm families. Annals of Botany, 87: 251–258

Hokche O, Berry PE, Huber O, 2008. Nuevo Catálogo de la Flora Vascular de Venezuela. 1–859. Caracas, Venezuela: Fundación Instituto Botánico de Venezuela

Jørgensen PM, León-Yánez S, 1999. Catalogue of the vascular plants of Ecuador. Monographs in Systematic Botany from the Missouri Botanical Garden. 75:1-1182

Jørgensen PM, Nee MH, Beck SG, 2014. Catalago de las plantas vasculares de Bolivia. Monographs in Systematic Botany from the Missouri Botanical Garden, 127: 1-1744

Kolarkar AS, Singh N, Shankarnarayan Ka, 1981. Note on Mehendi (Lawsonia inermis L.) cultivation in normal and degraded lands of Western Rajastan. Indian Journal of Soil Conservation, 9:71-74.

Koshcheev AV, Bukin VP, 1988. The effect of soil salinity on the productivity of henna. Agrokhimicheskoe i Pochvennoe Issledovanie v Botanicheskikh Sadakh, 68-72; 3 ref.

Lemmens RHMJ, Wulijarni-Soetjipto N (Editors), 1991. Plant resources of South-East Asia. No. 3. Dye and tannin-producing plants. pp.195.

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

Mohammed-Ali, 1996. Chemical and medical evaluation of Lawsonia inermia (henna). Hamard Medicus, 39(4):43-48; 100 ref.

Orwa C, Mutua A, Kindt R, Jamnadass R, Anthony S, 2009. Agroforestree Database:a tree reference and selection guide version 4.0 (http://www.worldagroforestry.org/sites/treedbs/treedatabases.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

Oyen LPA, 1991. Lawsonia inermis L. Record from Proseabase. In: Lemmens RHMJ, Wulijarni-Soetjipto N, Eds. Bogor, Indonesia: PROSEA (Plant Resources of South-East Asia) Foundation. http://www.proseanet.org

Parker RN, 1956. A forest flora for the Punjab with Hazara and Delhi. Lahore, Pakistan: Government Printing Press.

Patel MK, Bhatt VV, Kulkarni Ys, 1951. Three new bacterial diseases of plants from Bombay. Curr. Sci. 20 (12), (326-7).

Phirke SS, Saha M, 2013. Lawsonia inermis: A rainfed ratoon crop. National Conference on Biodiversity: Status and Challenges in Conservation–FAVEO, pp. 189-193

PIER, 2017. Pacific Island Ecosystems at Risk. http://www.hear.org/pier/index.html

Prendergast HDV, 1994. Tree seeds from the Sultanate of Oman in the Kew Seed Bank. Plant Genetic Resources Newsletter, No. 97, 37-42; 24 ref.

PROTA, 2017. Plant Resources of Tropical Africa. http://www.prota4u.org/search.asp

Quiniones SS, Dayan MP, 1981. Notes on the diseases of forest species in the Philippines. Sylvatrop, 6(2):61-67; 10 ref.

Russel S, Ifiorah CC, 1990. Occurrence of dinitrogen-fixing bacteria and acetylene reduction activity in [the] rhizosphere of five selected plants from the tropics. Annals of Warsaw Agricultural University SGGW AR, Agriculture, No.22, 63-66; 13 ref.

Salgare SA, Mamta Rawal, Rawal M, 1990. Effect of auto-exhaust pollution at Andheri (West), Bombay on the anatomy of some cultivated plants (monsoon season). Journal of Ecobiology, 2(4):273-280; 9 ref.

Sharma ND, Singh SR, 1992. Some records of fungi from central India. JNKVV Research Journal, 26(2):47-48; 10 ref.

Smith AC, 1985. Flora Vitiensis nova: a new flora of Fiji. National Tropical Botanical Garden, Lawai, Kauai, Hawaii. Volume 3. 758 pp.

Stephan ZA, 1988. Newly reported hosts of root-knot nematodes in Iraq. International Nematology Network Newsletter, 5(3):36-43

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

Swailem SM, Awadallah KT, Shaheen AA, 1976. Abundance of Lindingaspis rossi Mask. on ornamental host plants in Giza and Zagazig regions, Egypt (Hemiptera-Homoptera: Diaspididae). Bulletin of the Entomological Society of Egypt, No. 60:257-263

Takeda Y, Fatope MO, 1988. New phenolic glucosides from Lawsonia inermis. Journal of Natural Products, 51(4):725-729; 11 ref.

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

USDA-ARS, 2017. 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

Wong KC, Teng TE, 1995. Volatile components of Lawsonia inermis L. flowers. Journal of Essential Oil Research, 7(4):425-428; 21 ref.

Zavada MS, 1993. The historical use of Henna (Lawsonia inermis L.) in the Balkans. Thaiszia Journal of Botany, 3: 97-100

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

Broome R, Sabir K, Carrington S, 2007. Plants of the Eastern Caribbean. Online database. In: Plants of the Eastern Caribbean. Online database. Barbados: University of the West Indies. http://ecflora.cavehill.uwi.edu/index.html

CABI, Undated. Compendium record. Wallingford, UK: CABI

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

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

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

Davidse G, Sousa Sánchez M, Knapp S, Chiang Cabrera F, 2009. (Flora Mesoamericana). In: Flora Mesoamericana, St. Louis, USA: Missouri Botanical Garden.

Flora of China Editorial Committee, 2013. Flora of China., Beijing & St. Louis, Missouri Botanical Garden Press.

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.

Hokche O, Berry P E, Huber O, 2008. Nuevo Catálogo de la Flora Vascular de Venezuela. Caracas, Venezuela: Fundación Instituto Botánico de Venezuela. 860 pp.

Jørgensen PM, León-Yánez S, 1999. Catalogue of the vascular plants of Ecuador. In: Monographs in Systematic Botany from the Missouri Botanical Garden, 75 1-1181.

Jørgensen PM, Nee MH, Beck SG, 2014. (Catalago de las plantas vasculares de Bolivia). In: Monographs in Systematic Botany from the Missouri Botanical Garden, 127 1-1744.

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

Orwa C, Mutua A, Kindt R, Jamnadass R, Simons A, 2009. Agroforestree Database: a tree reference and selection guide. Version 4. In: Agroforestree Database: a tree reference and selection guide. Version 4. Nairobi, Kenya: World Agroforestry Centre. http://www.worldagroforestry.org/sites/treedbs/treedatabases.asp

Oviedo Prieto R, Herrera Oliver P, Caluff M G, 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 No. 1), 22-96.

PIER, 2017. Pacific Islands Ecosystems at Risk. In: Pacific Islands Ecosystems at Risk. Honolulu, Hawaii, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

PROTA, 2017. Plant Resources of Tropical Africa., http://www.prota4u.org/search.asp

Smith A C, 1985. Flora Vitiensis nova: a new flora of Fiji (Spermatophytes only). Volume 3: Angiospermae: Dicotyledones, families 117-163. Lawai, Hawaii, USA: Pacific Tropical Botanical Garden. vi + 758 pp.

USDA-ARS, 2017. Germplasm Resources Information Network (GRIN). Online Database. In: Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

Contributors

11/03/17 Original text by:

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

Distribution Maps