Thunbergia fragrans
(whitelady)

Identity

Preferred Scientific Name

• Thunbergia fragrans Roxb.

Preferred Common Name

• whitelady

Other Scientific Names

• Flemingia grandiflora Rottler
• Roxburghia rostrata Russell ex Nees
• Thunbergia bodinieri H. Lév.
• Thunbergia fragrans subsp. hainanensis (C.Y. Wu & H.S. Lo) H.P. Cui
• Thunbergia hainanensis C.Y. Wu & H.S. Lo
• Thunbergia volubilis Pers.

International Common Names

• English: angel wings; angelwings; fragrant thunbergia; sweet clock vine; sweet clockvine; sweet clock-vine; thunbergia; white clockvine; white lady; white thunbergia
• Spanish: flor de nieve; velo de novia
• French: bec martin; liane toupie
• Chinese: wan hua cao

Local Common Names

• Cuba: flor de nieve; jazmín del Vedado
• Dominican Republic: jazmín de maya
• Jamaica: white nightshade
• Puerto Rico: susana blanca
• United States Virgin Islands: white Susan vine

Summary of Invasiveness

T. fragrans is an herbaceous fast-growing vine widely cultivated as an ornamental in tropical and subtropical regions of the world, but it is also a common weed in moist disturbed areas, in particular along roadsides (Starr et al., 2003; Randall, 2012). In most cases, this species has been intentionally introduced as an ornamental and it has escaped from cultivation and naturalized in both relatively unaltered and disturbed forests, riversides, roadsides and urban bushland (Starr et al., 2003; Meyer and Lavergne, 2004; Queensland Department of Primary Industries and Fisheries, 2011). T. fragrans is included in the Global Compendium of Weeds where is listed as an “environmental weed,” and it is also listed as invasive in Australia, Japan, Singapore, Cuba, Puerto Rico, Hawaii and French Polynesia among others (Meyer and Lavergne, 2004; Mito and Uesugi, 2004; Chong et al., 2009; Queensland Department of Primary Industries and Fisheries, 2011; Oviedo-Prieto et al., 2012; PIER, 2014).

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Scrophulariales
•                         Family: Acanthaceae
•                             Genus: Thunbergia
•                                 Species: Thunbergia fragrans

Notes on Taxonomy and Nomenclature

The family Acanthaceae includes about 221 genera and 4000 species widespread in both New and Old World Tropics (Scotland and Vollesen, 2000; Stevens, 2012). Species within this family are herbs or woody shrubs, lianas and trees (Stevens, 2012). The subfamily Thunbergioideae comprise five genera, the largest of which, Thunbergia, contains about 100 species restricted to tropical and subtropical regions of Africa, Madagascar, Asia, and Australia (McDade et al., 2000; Borg et al., 2008). The subfamily Thunbergioideae is characterized by a predominantly twining habit, enlarged bracteoles, and a reduced calyx. Furthermore, Thunbergioideae lack the retinaculate fruits found in most Acanthaceae species, instead possessing either dry and/or dehiscent capsules without retinacula (Borg et al., 2008). The genus Thunbergia is named after the Swedish botanist and explorer, Carl Peter Thunberg (1743-1822).

Description

T. fragrans is an herbaceous vine, twining, 2-3 m in length. Stems cylindrical, striate, slender, puberulous. Leaves opposite; blades 6.5-11 × 1.8-6 cm, ovate to lanceolate, chartaceous, the apex acute, the base truncate or subcordiform; margins undulate and ciliate; upper surface dark green, glabrous or somewhat scabrous; lower surface pale green, dull, puberulous, with prominent venation; petiole 2-3.5 cm long, slender, pubescent, sulcate, with the base somewhat dilated. Flowers axillary, solitary or in pairs; pedicels pubescent, 5-7 cm long, striate; bracts green, membranaceous, ovate, pubescent, 1.6-2 cm long, covering the calyx. Calyx green, of 15-20 sepals, lanceolate, 3-5 mm long; corolla white, infundibuliform, with 5 lobes, the tube 2.5-4 cm long, narrow at the base, yellow inside, the limb 4-5 cm in diameter. Capsules 1-2.5 cm long, depressed-globose at the base, the upper half in the form of a beak, dehiscent in two halves; seeds 4, globose, approximately 5 mm in diameter, pubescent, with a depression at the base (Acevedo-Rodríguez, 2005). Variation in the shape, size, pubescence, and margin form of the leaves is extensive in T. fragrans, and taxa have been recognized based on these characters (Flora of China Editorial Committee, 2014).

Plant Type

Distribution

T. fragrans is native to India, Southeastern Asia and Malesia (Daniel, 2010; USDA-ARS, 2014). It has been widely cultivated as an ornamental and now it is cultivated and naturalized in North, Central and South America, the Caribbean and on many islands in the Indian and Pacific Ocean (see distribution table for details; Daniel, 1995, 2001, 2005; Meyer and Lavergne, 2004; Queensland Department of Primary Industries and Fisheries, 2011; PIER, 2014; USDA-NRCS, 2014).

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

T. fragrans was probably introduced as an ornamental in the Caribbean during the nineteenth century. It is recorded in the US National Herbarium from collections made in 1845 on St Thomas Island (US Virgin Islands), in 1874 in Trinidad, and in 1885 in Puerto Rico. By 1900, I. Urban listed this species as “spontaneous” for the islands of Cuba, Jamaica, St. Thomas, St. Croix, Puerto Rico, Guadeloupe, Martinique, St. Vincent, Barbados and Trinidad in his Symbolae Antillanae. In Hawaii, T. fragrans was first collected on Kaua'i in 1916 (Wagner et al. 1999). In Australia, T. fragrans is considered a species of “recent introduction” and by the year 2011 this species had been recorded at several locations in the coastal districts of eastern Queensland (Queensland Department of Primary Industries and Fisheries, 2011).

Risk of Introduction

The risk of introduction of T. fragrans is very high. It is a fast-growing vine which has been intentionally introduced as an ornamental in many tropical, subtropical and warm temperate regions where it has escaped and naturalized becoming a serious threat for native plant communities (Starr et al., 2003; Meyer and Lavergne, 2004). Considering that T. fragrans spreads sexually by seeds and vegetatively by cuttings, stem fragments, and roots, the likelihood of invading and colonizing new habitats remains high. In addition, seeds are available from nurseries on the internet.

Habitat

Within its native distribution range (i.e., India and China), T. fragrans grows in thickets, forest borders, roadsides and scrub jungles from 400-2300 m (Flora of China Editorial Committee, 2014; India Biodiversity, 2014). In Australia, T. fragrans grows as a weed in riverbanks (i.e., riparian vegetation), closed forests, forest margins, plantation crops, roadsides, urban bushland, disturbed sites and waste areas in tropical and subtropical habitats (Queensland Department of Primary Industries and Fisheries, 2011). In Hawaii, T. fragrans is a common weed in moist disturbed lowland areas (Wagner et al., 1999). In Fiji, T. fragrans grows in thickets, along roadsides, and in coconut plantations near sea level (Smith, 1981). In the West Indies, this species grows as a weed in disturbed areas and along forest edges (Adams, 1972; Broome et al., 2007; Oviedo Prieto et al., 2012).

Habitat List

Biology and Ecology

Reproductive Biology

Species in the genus Thunbergia have bisexual, zygomorphic, axillary, and usually solitary flowers (Acevedo-Rodríguez, 2005). In the case of T. fragrans, pollinators are unknown, but based on floral traits the species could be considered entomophilous (Starr et al., 2003). 

Physiology and Phenology

In China, T. fragrans has been recorded flowering from August to January and fruiting from November to March (Flora of China Editorial Committee, 2014). In Australia, flowering occurs throughout the year, but is most abundant during summer and autumn (Queensland Department of Primary Industries and Fisheries, 2011). In Jamaica and Puerto Rico, this species has been recorded flowering and fruiting throughout the year (Adams, 1972; Acevedo-Rodríguez, 2005). 

Environmental Requirements

T. fragrans grows best on fertile soils with good drainage and pH ranging from 6.1 to 7.8 (PROTA, 2014). It is able to grow beneath closed forests (i.e., shaded areas) as well as in areas with full-sunlight exposure (Queensland Department of Primary Industries and Fisheries, 2011; PROTA, 2014).

Climate

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil reaction

• neutral

Soil texture

• light
• medium

Notes on Natural Enemies

According to Brickell and Zuk (1997), Thunbergia species are susceptible to spider mites, whiteflies, and scale insects.

Means of Movement and Dispersal

T. fragrans is easily propagated by seeds, which it produces abundantly in its native range and in the tropical countries of introduction (Meyer and Lavergne, 2004). T. fragrans also spreads vegetatively by cuttings, stem fragments, and roots (Starr et al., 2003; PIER, 2014). Seeds and plant fragments can be spread in dumped garden waste, and by water, soil movement, garden tools, and vehicles (Starr et al., 2003; Queensland Department of Primary Industries and Fisheries, 2011).

Pathway Causes

Pathway Vectors

Impact Summary

Environmental Impact

T. fragrans grows climbing on other vegetation, and forming a dense blanket. It is most commonly found on roadsides, but can spread into forests and scrub. It is regarded as a potentially significant environmental weed in Queensland (Queensland Department of Primary Industries and Fisheries, 2011).

Risk and Impact Factors

• 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
• Tolerant of shade
• Highly mobile locally
• Benefits from human association (i.e. it is a human commensal)
• Long lived
• Fast growing
• Reproduces asexually

• Conflict
• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Host damage
• Modification of successional patterns
• Monoculture formation
• Reduced native biodiversity
• Threat to/ loss of native species

• Competition - monopolizing resources
• Competition - shading
• Competition - smothering
• Competition - strangling
• Rapid growth
• Rooting

• Highly likely to be transported internationally deliberately

Uses

T. fragrans has been widely used as an ornamental for its attractive flowers (Starr et al., 2003; Meyer and Lavergne, 2004). It is most often grown on trellises and fences, hanging baskets, and as a ground cover in gardens (Wagner et al., 1999; Queensland Department of Primary Industries and Fisheries, 2011; PIER, 2014; USDA-ARS, 2014).

Uses List

Environmental

• Amenity

General

• Souvenirs

Ornamental

• Potted plant
• Seed trade

Similarities to Other Species/Conditions

T. fragrans is relatively similar to Thunbergia grandiflora, Thunbergia laurifolia and Thunbergia alata. These three species can be distinguished by the following vegetative and reproductive characters (Queensland Department of Primary Industries and Fisheries, 2011):

• T. fragrans is an herbaceous creeper or climber bearing moderately large flowers (about 50 mm across). Corollas are entirely white and have a very narrow throat.
• T. grandiflora and T. laurifolia are relatively woody climbers bearing very large flowers (60-80 mm across). Corollas are usually blue or purplish with broad throats that are pale yellowish in colour.
• T. alata is a climber bearing relatively small flowers (25-40 mm across). Corollas are usually orange or yellow with conspicuous blackish-coloured throats that are quite narrow; petioles are winged.

Prevention and Control

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.

Small infestations should be removed manually. All plants segments and roots have to be removed in order to avoid re-sprouts. The use of this species as an ornamental should be discouraged (Starr et al., 2003; Queensland Department of Agriculture and Fisheries, 2011). 

In Australia, the only herbicide active registered for the control of Thunbergia species is imazapyr. This herbicide should be applied in a ratio of 7.5 ml/L water. For effective control, apply the herbicide when the plant is actively growing (Queensland Department of Agriculture and Fisheries, 2011).

References

Acevedo-Rodríguez P, 2005. Vines and climbing plants of Puerto Rico and the Virgin Islands. Contributions from the United States National Herbarium, 51:483 pp.

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

Adams CD, 1972. Flowering plants of Jamaica. Mona, Jamaica: University of the West Indies.

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

Borg AJ; McDade LA; Schönenberger J, 2008. Molecular phylogenetics and morphological evolution of Thunbergioideae (Acanthaceae). Taxon, 57(3):811-822.

Brickell C; Zuk JD, 1997. The American Horticultural Society A-Z Encyclopedia of Garden Plants. New York, USA: Dorling Kindersley Publishing, 1104 pp.

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

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.

Correa A; Galdames MDC; Stapf MNS, 2004. Catalogue of vascular plants of Panama (Catalogo de Plantas Vasculares de Panama.), Panama: Smithsonian Tropical Research Institute, 599 pp.

Daniel TF, 1995. Acanthaceae. 4. Flora Chiapas [ed. by Breedlove, D. E.]. San Francisco, USA: California Academy of Sciences, 1-158.

Daniel TF, 2001. Catalog of Acanthaceae in El Salvador. Contributions from the University of Michigan Herbarium, 23:115-137.

Daniel TF, 2005. Catalog of Honduran Acanthaceae with taxonomic and phytogeographic notes. Contributions from the University of Michigan Herbarium, 24:51-108.

Daniel TF, 2010. Catalog of Guatemalan Acanthaceae: taxonomy, ecology, and conservation. Proceedings of the California Academy of Science, 61:289-377.

Flora of China Editorial Committee, 2014. Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

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

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.

Hancock IR; Henderson CP, 1988. Flora of the Solomon Islands. Research Bulletin No. 7. Honiara, Solomon Islands: Dodo Creek Research Station.

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.

Idárraga-Piedrahita A; Ortiz RDC; Callejas Posada R; Merello M, 2011. Flora of Antioquia. (Flora de Antioquia.) Catálogo de las Plantas Vasculares, vol. 2. Listado de las Plantas Vasculares del Departamento de Antioquia:939 pp.

India Biodiversity, 2014. Online Portal of India Biodiversity. http://indiabiodiversity.org/species/list

Josekutty PC; Wakuk EE; Joseph MJ, 2002. Invasive weedy angiosperms in Kosrae, Federated States of Micronesia. Micronesica Supplement, 6:61-65. [Invasive species and their management.]

McDade LA; Masta SE; Moody ML; Waters E, 2000. Phylogenetic relationships among Acanthaceae: evidence from two genomes. Systematic Botany, 25:106-121.

Meyer JY; Lavergne C, 2004. Beautés fatales: Acanthaceae species as invasive alien plants on tropical Indo-Pacific islands. Diversity and Distributions, 10(5/6):333-347.

Mito T; Uesugi T, 2004. Invasive alien species in Japan: the status quo and the new regulation for prevention of their adverse effects. Global Environmental Research, 8(2):171-191.

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.

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

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

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

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

Scotland RW; Vollesen K, 2000. Classification of Acanthaceae. Kew Bulletin, 55:513-589.

Smith AC, 1981. Flora Vitiensis Nova: A new flora of Fiji. Lawai, Kauai, Hawaii. National Tropical Botanical Garden, 2:290.

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

Space JC; Flynn T, 2002. Report to the Government of Samoa on invasive plant species of environmental concern. Hawaii, USA: USDA Forest Service, Institute of Pacific Islands Forestry, 80 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

Starr F; Starr K; Loope LL, 2003. Thunbergia fragrans. Plants of Hawaii. Hawaii, USA: US Geological Survey, Biological Resources Division, Haleakala Field Station. http://www.hear.org/pier/pdf/pohreports/thunbergia_fragrans.pdf

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

Swarbrick JT, 1997. Weeds of the Pacific Islands. Technical paper No. 209. Noumea, New Caledonia: South Pacific Commission.

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

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

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

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

Whistler WA, 2000. Tropical ornamentals. Portland, Oregon, USA: Timber Press.

Wu TL, 2001. Check List of Hong Kong Plants. Agriculture, Fisheries and Conservation Department Bulletin 1 (revised):384 pp.

Zuloaga FO; Morrone O; Belgrano MJ; Marticorena C; Marchesi E, 2008. [English title not available]. (Catálogo de las Plantas Vasculares del Cono Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay).) Monographs in Systematic Botany from the Missouri Botanical Garden, 107:1-3348.

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

Adams C D, 1972. Flowering plants of Jamaica. Mona, Jamaica: University of the West Indies. 848 pp.

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.

Broome R, Sabir K, Carrington S, 2007. Plants of the Eastern Caribbean., 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 inferred from regional distribution. 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 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.

Correa A, Galdames MDC, Stapf MNS, 2004. Catalogue of vascular plants of Panama. (Catalogo de Plantas Vasculares de Panama)., Panama: Smithsonian Tropical Research Institute. 599 pp.

Daniel T F, 2001. Catalog of Acanthaceae in El Salvador. Contributions from the University of Michigan Herbarium. 115-137.

Daniel T F, 2005. Catalog of Honduran Acanthaceae with taxonomic and phytogeographic notes. Contributions from the University of Michigan Herbarium. 51-108.

Daniel TF, 1995. Acanthaceae. In: 4. Flora Chiapas, [ed. by Breedlove DE]. San Francisco, USA: California Academy of Sciences. 1-158.

Daniel TF, 2010. Catalog of Guatemalan Acanthaceae: taxonomy, ecology, and conservation. [Proceedings of the California Academy of Science], 61 289-377.

Flora of China Editorial Committee, 2014. Flora of China., St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

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

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.

Hancock I R, Henderson C P, 1988. Flora of the Solomon Islands. In: Research Bulletin - Dodo Creek Research Station, Honiara, Solomon Islands: ii + 203 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.

India Biodiversity, 2014. Online Portal of India Biodiversity., http://indiabiodiversity.org/species/list

Josekutty PC, Wakuk EE, Joseph MJ, 2002. Invasive weedy angiosperms in Kosrae, Federated States of Micronesia. In: Micronesica Supplement, 6 61-65.

Meyer J Y, Lavergne C, 2004. Beautés fatales: Acanthaceae species as invasive alien plants on tropical Indo-Pacific islands. Diversity and Distributions. 10 (5/6), 333-347. DOI:10.1111/j.1366-9516.2004.00094.x

Mito T, Uesugi T, 2004. Invasive alien species in Japan: the status quo and new regulations for prevention of their adverse effects. In: Global Environmental Research, 8 (2) 171-191.

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

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

Smith A C, 1981. Flora Vitiensis nova: a new flora of Fiji (spermatophytes only). Volume 2. In: Flora Vitiensis nova: a new flora of Fiji (spermatophytes only). Volume 2. Kauai, Hawaii, USA: Pacific Tropical Botanical Garden. 818 pp.

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

Space JC, Flynn T, 2002. Report to the Government of Samoa on invasive plant species of environmental concern., Hawaii, USA: USDA Forest Service, Institute of Pacific Islands Forestry. 80 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

Swarbrick JT, 1997. Weeds of the Pacific Islands. In: Technical paper No. 209, Noumea, New Caledonia, South Pacific Commission.

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

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

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

Wu TL, 2001. Check List of Hong Kong Plants. In: Agriculture, Fisheries and Conservation Department Bulletin, 1 384.

Zuloaga FO, Morrone O, Belgrano MJ, Marticorena C, Marchesi E, 2008. [English title not available]. (Catálogo de las Plantas Vasculares del Cono Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay)). In: Monographs in Systematic Botany from the Missouri Botanical Garden, 107 1-3348.

Links to Websites

Contributors

24/03/14 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

Distribution Maps