Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

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Triphasia trifolia
(limeberry)

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Datasheet

Triphasia trifolia (limeberry)

Summary

  • Last modified
  • 16 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Host Plant
  • Preferred Scientific Name
  • Triphasia trifolia
  • Preferred Common Name
  • limeberry
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • T. trifolia is an aggressive invasive plant with the capacity to grow in sunny open areas as well as in shaded areas beneath the canopy forest (

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Pictures

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PictureTitleCaptionCopyright
Trifoliate leaves and flowers of Triphasia trifolia
TitleTrifoliate leaves and flower
CaptionTrifoliate leaves and flowers of Triphasia trifolia
Copyright©Smithsonian Institution
Trifoliate leaves and flowers of Triphasia trifolia
Trifoliate leaves and flowerTrifoliate leaves and flowers of Triphasia trifolia©Smithsonian Institution
Flower of Triphasia trifolia
TitleFlower
CaptionFlower of Triphasia trifolia
Copyright©Smithsonian Institution
Flower of Triphasia trifolia
FlowerFlower of Triphasia trifolia©Smithsonian Institution
Ripe fruits of Triphasia trifolia
TitleRipe fruits
CaptionRipe fruits of Triphasia trifolia
Copyright©Smithsonian Institution
Ripe fruits of Triphasia trifolia
Ripe fruitsRipe fruits of Triphasia trifolia©Smithsonian Institution

Identity

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Preferred Scientific Name

  • Triphasia trifolia (Burm. f.) P. Wilson

Preferred Common Name

  • limeberry

Other Scientific Names

  • Limonia trifolia Burm. f.
  • Limonia trifoliata L. (nom. illeg.)
  • Triphasia trifoliata DC. (nom. illeg.)

International Common Names

  • English: orange berry; trifoliate limeberry; triphasia
  • Spanish: limón de Jerusalén; limoncito; mirto
  • French: orangine; petite citronelle

Local Common Names

  • Bahamas: chinese lemon; lime berry
  • Dominican Republic: naranjita de pegar
  • Guam: lemonchina; lemondichina
  • Lesser Antilles: chinese lemon; citronella; mutton lemon; myrtle lemon; myrtle lime; sweet lemon; sweet lime
  • Puerto Rico: china debakon; chinita
  • Saint Lucia: sitonnèl; sweet lime
  • Samoa: moli vai atigi lima; vali atigi lima
  • United States Virgin Islands: lime-berry

Summary of Invasiveness

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T. trifolia is an aggressive invasive plant with the capacity to grow in sunny open areas as well as in shaded areas beneath the canopy forest (Daley et al., 2012). It is a woody shrub classified as a “weed” and included in the Global Compendium of Weeds (Randall, 2012). T. trifolia spreads by seeds that can be easily dispersed by animals, mainly by birds, facilitating the invasion of this species into native forests. Seeds have high germination rates under natural conditions and grow forming dense-spiny thickets in the understory of the forest (Clark, 2003; Daley et al., 2012). For example, vegetation surveys performed in the understory of secondary forests on the islands of St. Croix (US Virgin Islands) have showed stem counts for this species exceeding 38,000 stems per hectare (Clark, 2003).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Rutales
  •                         Family: Rutaceae
  •                             Genus: Triphasia
  •                                 Species: Triphasia trifolia

Notes on Taxonomy and Nomenclature

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Rutaceae is a large plant family including about 154 genera and 2100 species distributed primarily in tropical and subtropical regions (Kubitzki et al., 2011). The genus Triphasia is located within the subfamily Aurantioideae along with nineteen genera including the genus Citrus. Triphasia includes three species native to South-East Asia (Indo-Malaysia), the Philippines and New Guinea (Kubitzki et al., 2011).

Description

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A glabrous shrub or small tree with paired spines in the axils of the leaves; leaves trifoliate; the terminal leaflet largest (2-4 cm long); lateral leaflets much smaller than the terminal one (1-2-2 x 0.8-1.2 cm), broadly rounded at the tip, cuneate at the base; petiolules very short (1.5-2 cm); petioles short (3-5 mm) and wingless. Flowers fragrant, calyx 1.5-2 mm long, 3-lobed, green, persistent; petals 3, white 8-9 mm long; stamens 6, filaments slender, glabrous, 5-7 mm long, anthers oblong, 2 x 1 mm; nectary disk annular or short-cylindric; ovary 3-locular. Fruits ellipsoid to globose, 1-1.5 cm long, dull reddish-orange or crimson when fleshy ripe; peel lemon-scented with many small oil glands; 1-3 seeds per fruit, immersed in mucilaginous pulpy flesh (Stone, 1970; Liogier, 1988; Acevedo-Rodríguez, 1996).

Plant Type

Top of page Perennial
Seed propagated
Shrub
Woody

Distribution

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T. trifolia is native to South-East Asia, Malaysia, and Christmas Islands. It was introduced as an ornamental and as a “hedge or boundary plant” in many tropical and subtropical countries and currently it can be found in Africa, the USA (Florida, Hawaii, and Texas), Mexico, Central America, the West Indies and the Pacific islands (PIER, 2012; USDA-ARS, 2012).

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 ReportedInvasiveReferenceNotes

Asia

CambodiaPresentNativePIER, 2012
Christmas Island (Indian Ocean)PresentNativePIER, 2012
IndonesiaPresentNativePIER, 2012
-JavaPresentNativePIER, 2012
LaosPresentNativePIER, 2012
MalaysiaPresentPresent based on regional distribution.
-Peninsular MalaysiaPresentNativePIER, 2012
PhilippinesPresentIntroduced Invasive PIER, 2012
SingaporePresentIntroducedPIER, 2012
ThailandPresentNativePIER, 2012
VietnamPresentNativePIER, 2012

Africa

RéunionPresentIntroduced Invasive PIER, 2012
Sierra LeonePresentIntroducedBurkill, 1994

North America

MexicoPresentIntroducedBreedlove, 1986
USAPresentPresent based on regional distribution.
-FloridaPresentIntroducedWunderlin and Hansen, 2008; USDA-ARS, 2012Cultivated
-HawaiiPresentIntroducedPIER, 2012Cultivated
-TexasPresentIntroducedUSDA-ARS, 2012Cultivated

Central America and Caribbean

Antigua and BarbudaWidespreadIntroducedBroome et al., 2007
BahamasPresentIntroducedCorrell and Correll, 1982
BarbadosWidespreadIntroduced Invasive Kairo et al., 2003
BelizePresentIntroducedBalick et al., 2000
British Virgin IslandsPresentIntroducedAcevedo-Rodríguez and Strong, 2012Tortola
Cayman IslandsPresentIntroducedAcevedo-Rodríguez and Strong, 2012
CubaPresentIntroducedGonzález-Torres et al., 2012Potentially invasive
DominicaWidespreadIntroducedBroome et al., 2007
Dominican RepublicPresentIntroducedAcevedo-Rodríguez and Strong, 2012
El SalvadorPresentIntroducedLinares, 2003Cultivated
GrenadaWidespreadIntroducedBroome et al., 2007
GuadeloupeWidespreadIntroducedBroome et al., 2007
GuatemalaPresentIntroducedStandley and Steyermark, 1946Cultivated
HondurasPresentIntroducedMolina, 1975
MartiniqueWidespreadIntroducedBroome et al., 2007
MontserratWidespreadIntroducedBroome et al., 2007
Netherlands AntillesPresentIntroducedMori et al., 2007Saba, St. Martin, St. Eustatius
PanamaPresentIntroducedD'Arcy, 1987
Puerto RicoPresentIntroduced Invasive Liogier, 1988Weed. Also on Mona Island
Saint Kitts and NevisWidespreadIntroducedBroome et al., 2007Nevis
Saint LuciaPresentIntroducedGraveson, 2012Cultivated
Saint Vincent and the GrenadinesWidespreadIntroducedBroome et al., 2007
United States Virgin IslandsPresentIntroduced Invasive Acevedo-Rodríguez, 1996; Chakroff, 2010; Acevedo-Rodríguez and Strong, 2012St. Croix, St. Thomas, St. John

South America

French GuianaPresentIntroducedFunk et al., 2007Escaped from cultivation
GuyanaPresentIntroducedFunk et al., 2007Escaped from cultivation

Oceania

FijiPresentIntroduced Invasive PIER, 2012
GuamPresentIntroducedPIER, 2012Naturalized
Micronesia, Federated states ofPresentIntroducedPIER, 2012
Northern Mariana IslandsPresentIntroduced Invasive PIER, 2012
Papua New GuineaPresentIntroducedPIER, 2012
SamoaPresentIntroducedPIER, 2012
Solomon IslandsPresentNativePIER, 2012

History of Introduction and Spread

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T. trifolia was probably introduced into the West Indies as an ornamental plant during the nineteenth century. It appears in an 1882 collection made on the islands of St. Thomas by H.F.A. Eggers (Smithsonian Herbarium Collections). Later, in 1886 it was collected in Quebradillas, Puerto Rico by P.E.E. Sintenis (Smithsonian Herbarium Collections). By 1905, Ignaz Urban (Symbolae Antillanae, IV: p. 320, 1905) reported this species as a “culta et quasi spontanea” for Puerto Rico, Cuba, Jamaica, Cayman Islands, Hispaniola, St. Thomas, St. Croix, St. Martin, St. Bartholomew, Guadeloupe, Martinique, and St. Vincent (Urban, 1905). In 1918, N.L. Britton in his book “Flora of Bermuda” reported this species as “frequent in gardens” for the island (Britton, 1918).

In Florida, T. trifolia was also introduced as an ornamental and by 1913 it was reported as an introduced species occurring on hammocks, coastal plain, fields and cultivated grounds (Small, 1913). By 1933 this species is also reported in Texas (Small, 1933). In 1999, it was included in the Florida List of Invasive Species as an invasive Category II, which are invasive plants that have increased in abundance or frequency but have not yet altered natural plant communities (Florida Exotic Pest Plant Council, 2011). As an action plan to control T. trifolia, the Florida Exotic Pest Council requested that nursery growers, landscape professionals and garden centre retailers voluntarily stop using the plant (Florida Exotic Pest Plant Council, 2011).

T. trifolia has also been introduced on islands in the Pacific, including Hawaii, Guam, Fiji, Mariana Islands, and Micronesia, but the year of introduction on those islands is not clear, and on some islands, for example Guam and Fiji, it is considered a “common naturalized shrub” (Stone, 1970; Smith, 1985). Currently, T. trifolia is considered an invasive species that threatens ecosystems of the Pacific islands (PIER, 2012).  

Risk of Introduction

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The risk of introduction of T. trifolia is high. It is widely commercialized as ornamental, windbreak plant, and hedge plant and it is still sold in the nursery and landscape trade around the world. T. trifolia is easily dispersed by seeds and it has the capacity to develop perfectly in open areas as well as in shaded areas (Daley et al., 2012). Consequently, the probability of this species invading and colonizing new habitats remains high.

Habitat

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T. trifolia can be found growing in sunny-open areas as well as in shaded areas beneath the understory of secondary forests, hammocks, scrublands, coastal forests, mesic forests, ruderal communities, riparian areas, and disturbed areas from sea level to low elevations (Small, 1933; Stone, 1970; Smith, 1985). Because this species grows forming dense-spiny tickets and adapts well to pruning, it is commonly used to create natural hedges in urban areas (Daley et al., 2012).

Habitat List

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CategoryHabitatPresenceStatus
Littoral
Coastal areas Present, no further details Harmful (pest or invasive)
Coastal areas Present, no further details Natural
Terrestrial-managed
Disturbed areas Present, no further details Natural
Urban / peri-urban areas Present, no further details Productive/non-natural
Terrestrial-natural/semi-natural
Natural forests Present, no further details Harmful (pest or invasive)
Natural forests Present, no further details Natural
Scrub / shrublands Present, no further details Harmful (pest or invasive)
Scrub / shrublands Present, no further details Natural

Biology and Ecology

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Genetics

The chromosome number in T. trifolia is 2n = 18 (Guerra et al., 2000).

Reproductive Biology

There are no studies available on the reproductive biology of T. trifolia. However, floral traits such as white and fragrant flowers, the production of nectar, and diurnal anthesis, suggest that this is an entomophilous species (Willmer, 2011).

Physiology and Phenology

Flowering and fruiting occurs throughout the year, with concentrations during summer months (Daley et al., 2012; PIER, 2012).

Environmental Requirements

T. trifolia prefers to grow in humid tropical areas on moderately loam sandy soils with pH between 5 and 6 (Verheij and Coronel, 1991). However, this species can also grow on limestone substrate (Daley et al., 2012). T. trifolia is a shade-tolerant species but it is also adapted to growth on open areas with full sunlight. It does not tolerate water-logging or frost (Daley et al., 2012; ISSG, 2012; PIER, 2012).

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Preferred > 60mm precipitation per month
Am - Tropical monsoon climate Preferred Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall5002000mm; lower/upper limits

Rainfall Regime

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Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • alkaline
  • neutral

Soil texture

  • light
  • medium

Means of Movement and Dispersal

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T. trifolia spreads by seeds. Plants produce fruits throughout the year, and seeds can be dispersed by birds and bats (Wiles, 1987; PIER, 2012). This species is commonly planted in gardens and used as a hedge plant. Thus, seeds can be dispersed from gardens into natural forests (ISSG, 2012; PIER, 2012). As a shade-tolerant species, T. trifolia can germinate and establish in the understory of the forest (Daley et al., 2012).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Escape from confinement or garden escapeUsed as ornamental Yes Yes USDA-ARS, 2012
FoodFruits are edible Yes Yes USDA-ARS, 2012
Hedges and windbreaksUsed as hedge plant Yes USDA-ARS, 2012
Internet salesSeeds and plants sold online Yes Yes
Landscape improvementOrnamental use Yes Yes USDA-ARS, 2012
Medicinal use Yes Yes USDA-ARS, 2012
Nursery trade Yes Yes USDA-ARS, 2012
Seed tradeSeeds sold online Yes Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Host and vector organismsSeeds dispersed by birds Yes PIER, 2012
MailSeeds sold online Yes Yes

Impact Summary

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

Economic Impact

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T. trifolia is one of the hosts used by the Asian citrus psyllid Diaphorina citri. This psyllid can be one of the most serious pests of citrus if the pathogens that cause citrus greening disease are present. Citrus greening, also called Huanglongbing or yellow dragon disease, is one of the more serious diseases of citrus. Transmission of citrus greening occurs primarily via infective citrus psyllids and grafting (Halbert and Manjunath, 2004).

Environmental Impact

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Impact on Habitats

T. trifolia is commonly used as an ornamental in gardens because it attracts birds, and as a “hedge plant”. Unfortunately, birds and other animals (i.e., bats) can disperse seeds into adjacent disturbed forests as well as into native forests (Wiles, 1987; PIER, 2012). This species is shade-tolerant and thus it can establish in the understory of forests. Once established, it grows forming dense-spiny thickets altering native plant communities. In the understory of native forests, T. trifolia grows taller than native plants and consequently smothers these plants, out-competing them for light and water (Clark, 2003; Daley et al., 2012).

Impact on Biodiversity

On Mona Island (Puerto Rico), the endemic and endangered iguana Cyclura cornuta stejnegeri has been impacted by the presence of T. trifolia. On this island, sites in the sandy coastal forests which provide areas with suitable environmental conditions for nesting iguanas were colonized by T. trifolia, limiting nesting sites for the iguana (Pérez-Buitrago et al., 2010; Rojas-Sandoval, personal observation).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Cyclura stejnegeri (Mona Island iguana)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as threatened species USA ESA listing as threatened speciesPuerto RicoPérez-Buitrago et al., 2010

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Tolerant of shade
  • Highly mobile locally
  • Benefits from human association (i.e. it is a human commensal)
  • Long lived
Impact outcomes
  • Conflict
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Loss of medicinal resources
  • Monoculture formation
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - shading
  • Competition - smothering
  • Competition - strangling
  • Pest and disease transmission
  • Produces spines, thorns or burrs
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately

Uses

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T. trifolia is commonly used an ornamental in tropical and subtropical regions. It is popular among landscapers as a consequence of its fragrant flowers and bright-red fruits that attract birds to gardens. This species is also commercialized as a boundary or hedge plant, mainly because it adapts well to the frequent and severe pruning required to create a natural hedge. Plants of T. trifolia are also grown for their fruits. Fruits are acid but edible and they can be consumed in preserved, jams, and beverages (Verheij and Coronel, 1991; Burkill, 1994).

T. trifolia is also used in traditional medicine in South East Asia. Leaves exude a resinous scent when bruised, and the plant is considered antifungal and antibacterial and used to treat colic, diarrhea, and skin afflictions. Fruits are used for coughs and sore throats (Burkill, 1994).

Chemical compounds extracted from leaves and stems of T. trifolia have been evaluated for their inhibitory effects against herpes simplex virus (HSV) and human immunodeficiency virus (HIV) (Likhitwitayawuid et al., 2005; Dondon et al., 2006). A recent study evaluating the inhibitory potential of 13 coumarins from T. trifolia demonstrated that it can be targeted as a prospective species for anti-HSV and HIV drug development (Likhitwitayawuid et al., 2005).

 

Uses List

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Environmental

  • Amenity
  • Boundary, barrier or support
  • Windbreak

Human food and beverage

  • Beverage base
  • Fruits

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Ornamental

  • Seed trade

Prevention and Control

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T. trifolia is a difficult species to control. It has flexible stems that are very difficult to cut with a machete or remove by hand. Seedlings and young plants should be pulled up and removed from treated areas. Fruits and fruiting branches should be also removed from treated areas. Larger trees should be cut with special machinery, and follow-up treatments are required to control sprouts (Daley et al., 2012).

Gaps in Knowledge/Research Needs

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  1. Studies on the history of introduction of this species are highly recommended.
  2. Studies on reproductive biology and breeding systems.
  3. Studies evaluating the impact of this exotic species on native plants and natural communities are needed in order to develop appropriate management and control strategies.
  4. Recommendations for management and control in natural areas invaded by this species are also needed.

References

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Acevedo-Rodríguez P, 1996. Flora of St. John, U.S. Virgin Islands. Memoirs of the New York Botanical Garden, 78:1-581.

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

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

Breedlove DE, 1986. Flora de Chiapas. Listados Florísticos de México, 4: 1-246.

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

Burkill HM, 1994. Useful plants of the West Tropical Africa. Volumen 4, Families M-R. London, UK: Kew Publishing, Royal Botanic Gardens, 981pp.

Chakroff M, 2010. Virgin Islands Forest Resources Assessment and Strategies. A comprehensive analysis of forest-related conditions, trends, threats and opportunities. Kingshill, Virgin Islands, USA: USDA Forest Service's International Institute of Tropical Forestry, Forestry Division, VI Department of Agriculture.

Chase MW; Morton CM; Kallunki JA, 1999. Phylogenetic relationships of Rutaceae: a cladistic analysis of the subfamilies using evidence from rbcL and atpB sequence variation. American Journal of Botany, 86(8):1191-1199.

Clark D, 2003. Weeds are Still "Weeds" in Paradise. Wildland Weeds, Winter:16-17. http://www.se-eppc.org/wildlandweeds/pdf/Winter2003-Clark-pp16-17.pdf

Correll DS; Correll HB, 1982. Flora of the Bahama Archipelago. Vaduz, Germany: J. Cramer, 1692 pp.

Daley B; Valiulis J; Slatton R, 2012. Exotic Invasives; US Virgin Islands; Species Effecting Forests. [Doc. # GS-VIDA-1201.]

D'Arcy WG, 1987. Flora of Panama: Checklist and Index. Monographs in Systematic Botany from the Missouri Botanical Garden. St. Louis, Missouri, USA: Missouri Botanical Garden.

Dondon R; Bourgeois P; Fery-Forgues S, 2006. A new bicoumarin from the leaves and stems of Triphasia trifolia. Fitoterapia, 77:129-133.

Florida Exotic Pest Plant Council, 2011. Florida EPPC's 2011 Invasive Plant Species List. http://www.fleppc.org/list/11list.html

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.

González-Torres LR; Rankin R; Palmarola A (eds), 2012. Invasive plants in Cuba. (Plantas Invasoras en Cuba.) Bissea: Boletin sobre Conservacion de Plantad del Jardin Botanico Nacional, 6:1-140.

Graveson R, 2012. Plants of Saint Lucia. http://www.saintlucianplants.com

Guerra M; Santos KGBdos; Silva AEB; Ehrendorfer F, 2000. Heterochromatin banding pattern in Rutaceae-Aurantioideae: A case of parallel chromosomal evolution. American Journal of Botany, 87:735 - 747.

Halbert SE; Manjunath KL, 2004. Asian citrus psyllids (Sternorrhyncha: Psyllidae) and greening disease in citrus: a literature review and assessment of risk in Florida. Florida Entomologist, 87(3):330-353. http://www.fcla.edu/FlaEnt/

ISSG, 2012. Global Invasive Species Database (GISD). Auckland, New Zealand: University of Auckland. http://www.issg.org/database

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

Kubitzki K; Kallunki JA; Duretto M; Wilson PG, 2011. Rutaceae. In: The Families and Genera of Vacular Plants. Vol. 10 [ed. by Kubitzki, K.]. Berlin, Germany: Springer-Verlag, 276-356.

Likhitwitayawuid K; Supudompol B; Sritularak B; Lipipun V; Rapp K; Schinazi RF, 2005. Phenolics with anti-HSV and anti-HIV activities from Artocarpus gomezianus, Mallotus pallidus, and Triphasia trifolia. Pharmaceutical Biology, 43(8):651-657.

Linares JL, 2003. Commented list on the native and cultivated trees of El Salvador. (Listado comentado de los árboles nativos y cultivados en la República de El Salvador.) CEIBA, 44(2):105-268.

Liogier AH, 1988. Flora of Puerto Rico and Adjacent Islands: A Systematic Synopsis. Rio Piedras, Puerto Rico: Editorial de la Universidad de Puerto Rico.

Molina RA, 1975. Enumeration of the plants of Honduras. (Enumeración de las plantas de Honduras) Ceiba, 19(1):1-118.

Mori SA; Buck WR; Gracie CA; Tulig M, 2007. Plants and Lichens of Saba. [Virtual Herbarium of The New York Botanical Garden.] http://sweetgum.nybg.org/saba/

Pérez-Buitrago N; Sabat AM; McMillan WO, 2010. Spatial ecology of the Endangered Mona Island iguana Cyclura cornuta stejnegeri: Does territorial behavior regulate density? Herpetological Monographs, 24:86-110.

PIER, 2012. Pacific Islands Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Randall RP, 2012. A global compendium of weeds, 2. Western Australia: Department of Agriculture and Food Western Australia, 1124 pp.

Small JK, 1913. Handbook of the native and naturalized shrubs of Florida.

Small JK, 1933. Manual of the Southeastern Flora. Chapel Hill: The University of North Carolina Press, 473.

Smith AC, 1985. Flora Vitiensis nova: a new flora of Fiji. Lawai, Kauai, Hawaii, USA: National Tropical Botanic Gardens, 758 pp.

Standley PC; Steyermark JA, 1946. Rutaceae. Flora of Guatemala - Part V. Fieldiana, Botany, 24(5):398-425.

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

Stone BC, 1970. The Flora of Guam. Micronesica, 6:1-659.

Urban I, 1905. Symbolae Antillanae. Volumen IV. Berlin, Germany: Fratres Borntraeger, 771 pp.

USDA-ARS, 2012. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

Verheij EWM; Coronel RE(Editors), 1991. Plant resources of South-East Asia. No. 2. Edible fruits and nuts. Wageningen, Netherlands; Pudoc, 446 pp.

Wiles GJ, 1987. The status of fruit bats on Guam. Pacific Science, 41:148-157.

Willmer P, 2011. Pollination and floral ecology. Princeton, USA: Princeton University Press.

Wunderlin RP; Hansen BF, 2008. Atlas of Florida Vascular Plants. Tampa, Florida, USA: University of South Florida. http://www.plantatlas.usf.edu/

Links to Websites

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WebsiteURLComment
Citrus Variety Collection, Triphasia, University of California Riversidehttp://www.citrusvariety.ucr.edu/citrus/triphasia.html
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
Pacific Island Ecosystems at Risk (PIER)http://www.hear.org/Pier/index.html

Contributors

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13/11/12 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

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