Invasive Species Compendium

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Datasheet

Barleria cristata
(Philippine violet)

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Datasheet

Barleria cristata (Philippine violet)

Summary

  • Last modified
  • 13 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Barleria cristata
  • Preferred Common Name
  • Philippine violet
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • B. cristata is a fast growing perennial plant often cultivated in gardens for its showy flowers. This species has repeatedly escaped from cultivation and naturalized in disturbed sites, abandoned gardens, river...

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Pictures

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PictureTitleCaptionCopyright
Barleria cristata (Philippine violet); flowers. Ananthagiri Hills, Rangareddy, Andhra Pradesh, India. October 2009.
TitleFlowers
CaptionBarleria cristata (Philippine violet); flowers. Ananthagiri Hills, Rangareddy, Andhra Pradesh, India. October 2009.
Copyright©J.M. Garg-2009/via wikipedia - CC BY 3.0
Barleria cristata (Philippine violet); flowers. Ananthagiri Hills, Rangareddy, Andhra Pradesh, India. October 2009.
FlowersBarleria cristata (Philippine violet); flowers. Ananthagiri Hills, Rangareddy, Andhra Pradesh, India. October 2009.©J.M. Garg-2009/via wikipedia - CC BY 3.0
Barleria cristata (Philippine violet); flowers and leaves. Kaluaaha, Molokai, Hawaii, USA. May 2005.
TitleFlowers and leaves
CaptionBarleria cristata (Philippine violet); flowers and leaves. Kaluaaha, Molokai, Hawaii, USA. May 2005.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Barleria cristata (Philippine violet); flowers and leaves. Kaluaaha, Molokai, Hawaii, USA. May 2005.
Flowers and leavesBarleria cristata (Philippine violet); flowers and leaves. Kaluaaha, Molokai, Hawaii, USA. May 2005.©Forest Starr & Kim Starr - CC BY 4.0
Barleria cristata (Philippine violet); habit. Kaluaaha, Molokai, Hawaii, USA. May 2005.
TitleHabit
CaptionBarleria cristata (Philippine violet); habit. Kaluaaha, Molokai, Hawaii, USA. May 2005.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Barleria cristata (Philippine violet); habit. Kaluaaha, Molokai, Hawaii, USA. May 2005.
HabitBarleria cristata (Philippine violet); habit. Kaluaaha, Molokai, Hawaii, USA. May 2005.©Forest Starr & Kim Starr - CC BY 4.0
Barleria cristata (Philippine violet); flowering habit. Narsapur, Andhra Pradesh, India. October 2008.
TitleFlowering habit
CaptionBarleria cristata (Philippine violet); flowering habit. Narsapur, Andhra Pradesh, India. October 2008.
Copyright©J.M. Garg-2008/via wikipedia - CC BY 3.0
Barleria cristata (Philippine violet); flowering habit. Narsapur, Andhra Pradesh, India. October 2008.
Flowering habitBarleria cristata (Philippine violet); flowering habit. Narsapur, Andhra Pradesh, India. October 2008.©J.M. Garg-2008/via wikipedia - CC BY 3.0

Identity

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Preferred Scientific Name

  • Barleria cristata L.

Preferred Common Name

  • Philippine violet

Other Scientific Names

  • Barleria alba Lodd.
  • Barleria ciliata Roxb.
  • Barleria cristata var. albida Haines
  • Barleria cristata var. dichotoma (Roxb.) Prain
  • Barleria cristata var. mairei H. Lév.
  • Barleria dichotoma Roxb.
  • Barleria indica Roxb. L. ex T. Anderson
  • Barleria laciniata Wall.
  • Barleria lactea Desf. ex Steud.
  • Barleria napalensis Nees
  • Barleria napalensis var. microphylla Nees
  • Barleria nuda Nees
  • Barleria prinoides Nees
  • Barleria venulosa Nees

International Common Names

  • English: barleria; bluebell barleria; crested Philippine violet
  • Chinese: jia du juan

Local Common Names

  • Australia: Philippine violet
  • Fiji: tombithi
  • Haiti: harré-volé; jolie-jolie
  • Puerto Rico: enana
  • Thailand: crested Philippine violet; kaan chang; luem thao yai; thong ra-aa
  • USA: crested Philippine violet; Philippine violet

EPPO code

  • BAECR (Barleria cristata)

Summary of Invasiveness

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B. cristata is a fast growing perennial plant often cultivated in gardens for its showy flowers. This species has repeatedly escaped from cultivation and naturalized in disturbed sites, abandoned gardens, riverbanks, and along roadsides (Daniel, 2001; Flora of China Editorial Committee, 2015). B. cristata has the potential to tolerate a remarkable range of habitats including human-disturbed sites, wastelands, and xeric vegetation (Meyer and Lavergne, 2004). Currently, it is listed as an invasive plant species in Australia, Fiji, Hawaii, French Polynesia, Mauritius, Réunion, and Puerto Rico (Smith, 1991; Meyer and Lavergne, 2004; PIER, 2015; Rojas-Sandoval and Acevedo-Rodriquez, 2015). In Australia, it is listed as an environmental weed (Weeds of Australia, 2015). 

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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Acanthaceae is a family of flowering plants comprising about 220 genera and about 4000 species widespread in both New and Old World Tropics (Scotland and Vollesen, 2000; Stevens, 2012). Species within the Acanthaceae includes herbs or woody shrubs, lianas and trees. Barleria is a large, polymorphic, widespread genus of herbs, shrubs and rarely climbers comprising approximately 300 species distributed worldwide (Balkwill and Balkwill, 1998). The greatest representation of this genus occurs in Africa and Asia, with its greatest centre of diversity in tropical East Africa (Balkwill and Balkwill, 1997). The genus Barleria is readily distinguished from other genera in the Acanthaceae by a combination of three features:

(1)  a four-merous calyx with two large outer sepals and two smaller inner ones.

(2)  globose, honey-combed pollen.

(3)  predominance of double cystoliths (calcium oxalate crystals) in the epidermal cells (Balkwill and Balkwill, 1998).

Although each of these features occurs in other Acanthaceae species, their regular co-occurrence is restricted to Barleria (Balkwill and Balkwill, 1997).

The genus name Barleria is in honour of Jacques Barrelier (1606-1673), a French Dominican monk who was a physician, botanist, plant collector and author; cristata is from the Latin cristatus, crested.

Description

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The following description is taken from Flora of China Editorial Committee (2015):

B. cristata is a perennial subshrub to 2 m tall. Stems terete, branched, covered with soft trichomes. Petiole (0-)0.3-1 cm; leaf blade elliptic to oblong to ovate, 2-10 × 1-4 cm, both surfaces villous especially along veins, secondary veins 4-7 on each side of midvein, base cuneate and decurrent onto petiole, apex acute to sometimes acuminate. Inflorescences axillary short and dense cymes, shortly pedunculate; bracts absent; bracteoles variable, linear to linear-lanceolate, 2.4-6.5 × 0.5-1.5 cm, 3-7-veined, base cuneate, margin usually spiny but sometimes bristly pilose and becoming spinescent with age, apex acuminate. Outer calyx lobes ovate to narrowly elliptic to lanceolate, 1.2-2.5 × 0.5-1.3 cm, pilose, reticulately veined, margin spiny, apex mucronate; inner calyx lobes linear to lanceolate, 6-12 mm, 1-veined, margin scarious. Corolla purplish blue, 4.5-6.5 cm, outside pilose; tube basally narrowly cylindric then gradually widened; lobes oblong-elliptic, approximately 1.5 cm, equal. Stamens 4, didynamous; filaments pilose especially toward base; staminode 1, filament sparsely pilose. Ovary oblong-ellipsoid, glabrous; style linear, ca. 2.5 cm, glabrous; stigma slightly inflated. Capsule 1.2-1.8 cm, glabrous, 4-seeded. Seeds subglobose to ovoid, 4-5 × 4 mm.

While flowers are generally purple-blue, a white-flowered cultivar ‘Alba’ is present among both cultivated and naturalized populations (Weeds of Australia, 2015). 

Distribution

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B. cristata is native to Asia including China, India, Laos, Myanmar, Nepal, Pakistan, Sri Lanka, Thailand, and Vietnam. Currently, it has a pantropical distribution and can be found naturalized in Africa, North America, Central America, the West Indies, and on islands in the Pacific Ocean (see distribution table for details; Flora of China Editorial Committee, 2015; PIER, 2015; USDA-ARS, 2015). 

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasivePlantedReferenceNotes

Asia

BangladeshPresentNativeUSDA-ARS, 2015
BhutanPresentNative
CambodiaPresentNativeFlora of China Editorial Committee, 2015
ChinaPresentPresent based on regional distribution.
-FujianPresentNativeFlora of China Editorial Committee, 2015
-GuangdongPresentNativeFlora of China Editorial Committee, 2015
-GuangxiPresentNativeFlora of China Editorial Committee, 2015
-GuizhouPresentNativeFlora of China Editorial Committee, 2015
-HainanPresentNativeFlora of China Editorial Committee, 2015
-Hong KongPresentIntroducedWu, 2001Cultivated
-SichuanPresentNativeFlora of China Editorial Committee, 2015
-TibetPresentNativeFlora of China Editorial Committee, 2015
-YunnanPresentNativeFlora of China Editorial Committee, 2015
Christmas Island (Indian Ocean)PresentIntroduced Invasive Swarbrick, 1997
IndiaPresent Natural USDA-ARS, 2015
-KarnatakaPresentNativeIndia Biodiversity Portal, 2016
-KeralaPresentNativeIndia Biodiversity Portal, 2016
-MaharashtraPresentNativeIndia Biodiversity Portal, 2016
-Tamil NaduPresentNativeIndia Biodiversity Portal, 2016
IndonesiaPresentIntroducedUSDA-ARS, 2015Cultivated and naturalized
LaosPresentNativeUSDA-ARS, 2015
MyanmarPresent Natural USDA-ARS, 2015
NepalPresentNativeUSDA-ARS, 2015
PakistanPresentNativeUSDA-ARS, 2015
PhilippinesPresentFlora of China Editorial Committee, 2015Listed as both native and introduced
SingaporePresentIntroducedChong et al., 2009Cultivated
Sri LankaPresentNativeFlora of China Editorial Committee, 2015
TaiwanPresentNativeFlora of China Editorial Committee, 2015
ThailandPresentNativeUSDA-ARS, 2015
VietnamPresentNativeUSDA-ARS, 2015

Africa

MauritiusPresentIntroduced Invasive Meyer and Lavergne, 2004
RéunionPresentIntroduced Invasive Meyer and Lavergne, 2004

North America

USAPresentPresent based on regional distribution.
-FloridaPresentIntroducedUSDA-NRCS, 2015
-HawaiiPresentIntroduced Invasive Wagner et al., 1999

Central America and Caribbean

BahamasPresentIntroducedAcevedo-Rodriguez and Strong, 2012
BarbadosPresentIntroducedAcevedo-Rodriguez and Strong, 2012
Dominican RepublicPresentIntroducedAcevedo-Rodriguez and Strong, 2012
El SalvadorPresentIntroducedDaniel, 2001
HaitiPresentIntroducedAcevedo-Rodriguez and Strong, 2012
HondurasPresentIntroducedDaniel, 2005
JamaicaPresentIntroducedAcevedo-Rodriguez and Strong, 2012
MartiniquePresentIntroducedUS National Herbarium
MontserratPresentIntroducedAcevedo-Rodriguez and Strong, 2012
PanamaPresentIntroducedCorrea et al., 2004
Puerto RicoPresentIntroduced Invasive Rojas-Sandoval and Acevedo-Rodríguez, 2015
Saint Vincent and the GrenadinesPresentIntroducedAcevedo-Rodriguez and Strong, 2012
United States Virgin IslandsPresentIntroduced Invasive Rojas-Sandoval and Acevedo-Rodríguez, 2015St Thomas

South America

VenezuelaPresentIntroducedHokche et al., 2008Bolivar, Delta Amacuro, Distrito Federal, Falcon, Merida, Nueva Esparta, Sucre

Oceania

AustraliaPresentPresent based on regional distribution.
-QueenslandPresentIntroducedWeeds of Australia, 2015Naturalized
FijiPresentIntroduced Invasive Smith, 1991
French PolynesiaPresentIntroduced Invasive Florence et al., 2013
GuamPresentIntroducedPIER, 2015Cultivated
Marshall IslandsPresentIntroducedWagner et al., 2013
Micronesia, Federated states ofPresentIntroducedLorence and Flynn, 2010
NauruPresentIntroducedThaman et al., 1994Cultivated
New CaledoniaPresentIntroduced Invasive MacKee, 1994
NiuePresentIntroducedPIER, 2015Cultivated
Northern Mariana IslandsPresentIntroducedPIER, 2015
PalauPresentIntroducedSpace et al., 2009Cultivated
Solomon IslandsPresentIntroducedHancock et al., 1988
TongaPresentIntroducedPIER, 2015

History of Introduction and Spread

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B. cristata has been intentionally introduced as an ornamental into tropical and subtropical regions of the world since the 1800s. In the West Indies, this species appears in a herbarium collection made in 1884 in Martinique. In Puerto Rico it was introduced before the 1900s (Rojas-Sandoval and Acevedo-Rodriquez, 2015). In Australia, B. cristata was also introduced as an ornamental and it has recently become naturalized in the sub-tropical and tropical regions of eastern Australia (Weeds of Australia, 2015). 

Risk of Introduction

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The risk of introduction of B. cristata is high, mainly because this species is widely cultivated as an ornamental plant. The species spreads by seeds and rhizomes and thus it has the capability to escape and colonize new habitats. Stem segments and stolons can also be dispersed as a contaminant of soil and as a garden-waste.

Habitat

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Within its native distribution range, B. cristata can be found growing along roadsides, slopes, streams, and xeric vegetation at elevations below 100 m up to 2600 m (Flora of China Editorial Committee, 2015). In Hawaii, B. cristata can be found naturalized in dry habitats (Wagner et al., 1999). In Fiji, B. cristata is often cultivated in villages and gardens and can be found naturalized along roadsides from near sea level to about 100 m (Smith, 1991). In the West Indies, B. cristata is a common ornamental plant in gardens and can be found naturalized in ruderal sites and semi-natural habitats in dry and wet regions (Rojas-Sandoval and Acevedo-Rodriguez, 2015). 

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 Natural
Rail / roadsides Present, no further details Harmful (pest or invasive)
Rail / roadsides Present, no further details Natural
Urban / peri-urban areas Present, no further details Harmful (pest or invasive)
Urban / peri-urban areas Present, no further details Natural
Urban / peri-urban areas Present, no further details Productive/non-natural
Terrestrial ‑ Natural / Semi-naturalNatural grasslands Present, no further details Harmful (pest or invasive)
Natural grasslands Present, no further details Natural
Riverbanks Present, no further details Harmful (pest or invasive)
Riverbanks Present, no further details Natural

Biology and Ecology

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Genetics

The chromosome number reported for B. cristata is 2n = 40 (Flora of China Editorial Committee, 2015).

Reproductive Biology

B. cristata has bisexual, zygomorphic flowers. In China, it has been recorded flowering in May and August-December (Flora of China Editorial Committee, 2015). In India it blooms in December (India Biodiversity Portal, 2016).

Environmental Requirements

B. cristata prefers to grow in warm and humid areas from sea level up to 2600 m on sandy and loam sandy soils with pH ranging from 5.6 to 7.5 (Flora of China Editorial Committee, 2015). It has the potential to establish in areas with full sunny to partial shaded conditions.

Climate

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ClimateStatusDescriptionRemark
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)

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 10 26

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall8002500mm; lower/upper limits

Means of Movement and Dispersal

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B. cristata spreads mostly by stolons (Swarbrick, 1997). This species is dispersed long distances mostly by people who use the plant as an ornamental (Flora of China Editorial Committee, 2015). Stem fragments and stolons can be spread in dumped garden waste, and by water, soil movements, and vehicles. 

Impact Summary

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CategoryImpact
Economic/livelihood Positive and negative
Environment (generally) Positive and negative

Environmental Impact

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B. cristata is sometimes weedy and has the potential to outcompete native plants for common resources such as water, space, and light (Aguilar, 2001). In Australia, B. cristata is seen as a threat to riparian areas and is also regarded as an environmental weed (Weeds of Australia, 2015). 

Risk and Impact Factors

Top of page Invasiveness
  • 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
  • Benefits from human association (i.e. it is a human commensal)
  • Long lived
  • Fast growing
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Loss of medicinal resources
  • Modification of successional patterns
  • Monoculture formation
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - smothering
  • Hybridization
  • Rapid growth
  • Rooting
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately

Uses

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B. cristata is often cultivated as a medicinal and ornamental plant. It is also cultivated as a hedge plant. In Asia, it is used in traditional medicine. The bitter juice of the leaves or roots is used as a diaphoretic and expectorant for serious catarrhal infections. An infusion of the roots and leaves is applied to boils and sores to reduce swellings (Aguilar, 2001). 

References

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Acevedo-Rodríguez P; Strong MT, 2012. Catalogue of the Seed Plants of the West Indies. Smithsonian Contributions to Botany, 98:1192 pp. Washington DC, USA: Smithsonian Institution. http://botany.si.edu/Antilles/WestIndies/catalog.htm

Aguilar NO, 2001. Barleria cristata L. Record from Proseabase. PROSEA (Plant Resources of South-East Asia) [ed. by Valkenburg, J. L. C. H. van \Bunyapraphatsara, N.]. Bogor, Indonesia: PROSEA (Plant Resources of South-East Asia) Foundation. http://www.proseanet.org

Balkwill MJ; Balkwill K, 1997. Delimitation and infra-generic classification of Barleria (Acanthaceae). Kew Bulletin, 1:535-573.

Balkwill MJ; Balkwill K, 1998. A preliminary analysis of distribution patterns in a large, pantropical genus, Barleria L. (Acanthaceae). Journal of Biogeography, 25:95-110.

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

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, 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.

Flora of China Editorial Committee, 2015. 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

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.

India Biodiversity Portal, 2016. Online Portal of India Biodiversity. http://indiabiodiversity.org/species/list

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

MacKee HS, 1994. Catalogue of introduced and cultivated plants in New Caledonia. (Catalogue des plantes introduites et cultivées en Nouvelle-Calédonie.) Paris, France: Muséum National d'Histoire Naturelle, unpaginated.

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.

PIER, 2015. Pacific Islands Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Rojas-Sandoval J; Acevedo-Rodríguez P, 2015. Naturalization and invasion of alien plants in Puerto Rico and the Virgin Islands. Biological Invasions, 17(1):149-163. http://rd.springer.com/article/10.1007/s10530-014-0712-3/fulltext.html

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

Smith AC, 1991. Flora Vitiensis nova: A new flora of Fiji. Lawai, Kauai, Hawai`i. National Tropical Botanical Garden, Volume 5, 626 pp.

Space JC; Lorence DH; LaRosa AM, 2009. Report to the Republic of Palau: 2008 update on Invasive Plant Species. Hilo, Hawaii, USA: USDA Forest Service, 227. http://www.sprep.org/att/irc/ecopies/countries/palau/48.pdf

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

Swarbrick JT, 1997. Environmental weeds and exotic plants on Christmas Island, Indian Ocean. Report to Parks Australia. J.T. Swarbrick, Weed Science Consultancy, 131 pp.

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

USDA-ARS, 2015. 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, 2015. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Wagner WL; Herbst DR; Sohmer SH, 1999. Manual of the Flowering Plants of Hawaii, Revised ed. Honolulu, USA: University of Hawaii Press.

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

Weeds of Australia, 2015. Weeds of Australia, Biosecurity Queensland Edition. http://keyserver.lucidcentral.org/weeds/data/03030800-0b07-490a-8d04-0605030c0f01/media/Html/search.html?zoom_query=

Wu TL, 2001. Check List of Hong Kong Plants. Hong Kong Herbarium and the South China Institute of Botany. Agriculture, Fisheries and Conservation Department Bulletin 1 (revised):384 pp. http://www.hkflora.com/v2/flora/plant_check_list.php

Contributors

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30/04/15 Original text by:

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

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