Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Averrhoa carambola
(carambola)

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Datasheet

Averrhoa carambola (carambola)

Summary

  • Last modified
  • 14 November 2018
  • Datasheet Type(s)
  • Documented Species
  • Host Plant
  • Preferred Scientific Name
  • Averrhoa carambola
  • Preferred Common Name
  • carambola
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • A. carambola is a small, fruit-bearing tree possibly native to the Old World tropics that has been widely cultivated for its edible fruits. Despite being reported as ‘extinct in the wild’ (

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Pictures

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PictureTitleCaptionCopyright
Averrhoa carambola (carambola or starfruit); fruits and foliage. Maui Meadows Kihei, Maui, Hawaii, USA. August 2009.
TitleFruits and foliage
CaptionAverrhoa carambola (carambola or starfruit); fruits and foliage. Maui Meadows Kihei, Maui, Hawaii, USA. August 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Averrhoa carambola (carambola or starfruit); fruits and foliage. Maui Meadows Kihei, Maui, Hawaii, USA. August 2009.
Fruits and foliageAverrhoa carambola (carambola or starfruit); fruits and foliage. Maui Meadows Kihei, Maui, Hawaii, USA. August 2009.©Forest Starr & Kim Starr - CC BY 4.0
Averrhoa carambola (carambola or starfruit); fruiting habit. Maui Meadows Kihei, Maui, Hawaii, USA. August 2009.
TitleHabit
CaptionAverrhoa carambola (carambola or starfruit); fruiting habit. Maui Meadows Kihei, Maui, Hawaii, USA. August 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Averrhoa carambola (carambola or starfruit); fruiting habit. Maui Meadows Kihei, Maui, Hawaii, USA. August 2009.
HabitAverrhoa carambola (carambola or starfruit); fruiting habit. Maui Meadows Kihei, Maui, Hawaii, USA. August 2009.©Forest Starr & Kim Starr - CC BY 4.0
Averrhoa carambola (carambola or starfruit); leaves. Kula Ace Hardware and Nursery, Maui, Hawaii, USA. September 2007.
TitleLeaves
CaptionAverrhoa carambola (carambola or starfruit); leaves. Kula Ace Hardware and Nursery, Maui, Hawaii, USA. September 2007.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Averrhoa carambola (carambola or starfruit); leaves. Kula Ace Hardware and Nursery, Maui, Hawaii, USA. September 2007.
LeavesAverrhoa carambola (carambola or starfruit); leaves. Kula Ace Hardware and Nursery, Maui, Hawaii, USA. September 2007.©Forest Starr & Kim Starr - CC BY 4.0
Averrhoa carambola (carambola or starfruit); flowers and leaves. Napili, Maui, Hawaii, USA. July
TitleFlowers
CaptionAverrhoa carambola (carambola or starfruit); flowers and leaves. Napili, Maui, Hawaii, USA. July
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Averrhoa carambola (carambola or starfruit); flowers and leaves. Napili, Maui, Hawaii, USA. July
FlowersAverrhoa carambola (carambola or starfruit); flowers and leaves. Napili, Maui, Hawaii, USA. July ©Forest Starr & Kim Starr - CC BY 4.0

Identity

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Preferred Scientific Name

  • Averrhoa carambola L.

Preferred Common Name

  • carambola

Other Scientific Names

  • Averrhoa acutangula Stokes
  • Sarcotheca philippica (Villar) Hallier f.

International Common Names

  • English: carambola; carambola tree; carambold; Chinese gooseberry; Coromandel gooseberry; cucumber tree; five-corner; star fruit tree; starfruit
  • Spanish: carambolo
  • French: carambolier; carambolier vrai
  • Chinese: ma fen; yang-táo

Local Common Names

  • Australia: five corner
  • Brazil: camerunga; caramboleiro; limas de Cayena
  • Cambodia: spo; spu
  • Costa Rica: tiriguro
  • Dominican Republic: vinagrillo
  • El Salvador: pepino de la India
  • Finland: karambola
  • French West Indies: cornichon
  • Germany: Karambolabaum; karambole; Sternfrucht
  • Guam: bilimbin; bilimbines
  • Guyana: five fingers
  • Haiti: bimblin longue; blinblin longue; carambolier; conichon du pays; zibeline; zibeline longue; zibline; zimbline
  • India: kamaranga; kamrakh; kamruk
  • Indonesia: belimbing manis
  • Laos: fuang; nak fuang
  • Malaysia: belimbing batu; belimbing besi; belimbing manis; belimbing pessegi; belimbing saji; belimbing sayur; blinbing manis; caramba; carambola tree; kambola
  • Mexico: árbol de pepino; caramboler; carambolera
  • Myanmar: mak-hpung; zaung-yar
  • Nicaragua: melocotón
  • Pakistan: kamrak; kamranga
  • Palau: kemim; ouderteboteb
  • Panama: mamoncillo chino
  • Philippines: balimbin; balimbing; balingbing; daligan; dalihan; galangan; galuran; garahan; garulan; malimbin; sirinate
  • Puerto Rico: carambold; jalea; star pickle
  • Sri Lanka: kamaranga; kamruk
  • Suriname: blimbing legi; fransman-birambi
  • Sweden: karambola
  • Thailand: ma fu’ang; ma fueang
  • Tonga: tapanima
  • Trinidad and Tobago: coolie tamarind
  • Venezuela: tamarindo chino; tamarindo dulce
  • Vietnam: khe; khe ta

EPPO code

  • AVRCA (Averrhoa carambola)

Summary of Invasiveness

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A. carambola is a small, fruit-bearing tree possibly native to the Old World tropics that has been widely cultivated for its edible fruits. Despite being reported as ‘extinct in the wild’ (Missouri Botanical Garden, 2015), A. carambola is also listed in the Global Compendium of Weeds as “cultivation escape, naturalised, weed” (Randall, 2012), was included in a list of potential invasive species threats in the Caribbean Region (Kairo et al., 2003), and is naturalized in Puerto Rico, Belize, Micronesia, and Paraguay (Liogier and Martorell, 2000; Randall, 2012). Although currently a low risk species, further evaluation of the invasive potential of A. carambola may be necessary in the future, especially for areas surrounding fields and gardens where the species is grown. 

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Geraniales
  •                         Family: Oxalidaceae
  •                             Genus: Averrhoa
  •                                 Species: Averrhoa carambola

Notes on Taxonomy and Nomenclature

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Oxalidaceae, sometimes called the ‘wood sorrel’ family, includes over 800 species of herbs, shrubs, and trees, mostly of neotropical origin. The genus name Averrhoa is in honour of the famous twelth-century philosopher Averrhoes from Andalusia (Macfadyen, 1837). The species name carambola is its vernacular name used in most of the Old World and Neotropics. Several of the common names of the plant are associated with the shape of the fruit in cross-section, which resembles a 5-pointed star.

Description

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

Plants 3-12(-15) m tall, densely branched, young parts finely pubescent or glabrous. Leaves 7-25 cm; petiole 2-8 cm; leaflets (3-)5-13; petiolules 1-2.5 mm; leaflet blades ovate to elliptic, 3-8 × 1.5-4.5 cm, abaxially pubescent to nearly glabrous, base obliquely rounded, apex acute to acuminate. Inflorescences axillary or rameal, panicles or cymes, branches and flower buds crimson. Flowers numerous, small. Sepals narrowly elliptic, 3-5 mm, base sparingly pubescent. Petals white with purple markings or pink to red with darker markings, 6-9 × 3-4 mm. Shorter stamens sterile, occasionally 1 or 2 fertile. Ovary pubescent. Berry yellow to yellow-brown, oblong, 7-13 × 5-8 cm, deeply (3-)5(or 6)-ribbed, stellate in cross section, very fleshy. Seeds numerous, blackish brown.

Plant Type

Top of page Perennial
Seed propagated
Tree
Woody

Distribution

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A. carambola is considered native to Malesia, although it has also been speculated to be a tropical American species introduced to Asia by the Spanish galleons (Merrill, 1904; Quisumbing, 1951; Samson, 1991; Bircher and Bircher, 2000). It is now widely cultivated in both the Old World and New World tropics. 

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

CambodiaPresentMorton, 1987
ChinaPresentNativeFlora of China Editorial Committee, 2015; Flora of China Editorial Committee, 2015; Flora of Pakistan, 2015
-FujianPresent only in captivity/cultivationMorton, 1987; Flora of China Editorial Committee, 2015‘commonly grown’
-GuangdongPresentMorton, 1987; Flora of China Editorial Committee, 2015‘commonly grown’
-GuangxiPresentMorton, 1987; Flora of China Editorial Committee, 2015‘commonly grown’
-HainanPresentFlora of China Editorial Committee, 2015
-SichuanPresentFlora of China Editorial Committee, 2015
-YunnanPresentFlora of China Editorial Committee, 2015
IndiaPresentMorton, 1987; Flora of Pakistan, 2015; Missouri Botanical Garden, 2015‘commonly grown’
IndonesiaPresentNativeMissouri Botanical Garden, 2015
-JavaPresentMorton, 1987; USDA-ARS, 2015
-MoluccasPresentNativeMorton, 1987
IsraelPresent only in captivity/cultivationIntroduced1935Morton, 1987Rehovoth Research Station
MalaysiaPresentMorton, 1987; Bircher and Bircher, 2000; Flora Mesoamericana, 2014Cultivated for many centuries
MyanmarPresentKress et al., 2003; Flora of Pakistan, 2015
PakistanPresentFlora of Pakistan, 2015Cultivated
PhilippinesPresentIntroducedMerrill, 1904; Quisumbing, 1951; Samson, 1991Introduced from S. America
SingaporePresentIntroducedChong et al., 2009Casual alien
Sri LankaPresentNativeMorton, 1987; Missouri Botanical Garden, 2015
TaiwanPresent only in captivity/cultivationMorton, 1987; Flora of China Editorial Committee, 2015‘commonly grown’
ThailandPresentMorton, 1987; Hanelt et al., 2001
VietnamPresentMorton, 1987

Africa

MadagascarPresentFlora of Pakistan, 2015Cultivated
TanzaniaPresentPresent based on regional distribution.
-ZanzibarPresentMorton, 1987

North America

MexicoPresentFlora Mesoamericana, 2014Yucatan. Grown in Mesoamerica
USAPresentIntroducedRandall, 2012Naturalised
-FloridaPresentIntroducedbefore 1887Morton, 1987
-HawaiiPresentMorton, 1987

Central America and Caribbean

BelizePresentIntroducedRandall, 2012Naturalised
Costa RicaPresentBurger, 1991; Flora Mesoamericana, 2014‘widely planted’
CubaPresentIntroducedAcevedo-Rodriguez and Strong, 2012Cultivated
Dominican RepublicPresentIntroducedAcevedo-Rodriguez and Strong, 2012Cultivated
El SalvadorPresentFlora Mesoamericana, 2014
HaitiPresentIntroducedAcevedo-Rodriguez and Strong, 2012Cultivated
HondurasPresentFlora Mesoamericana, 2014
JamaicaPresentMorton, 1987; Acevedo-Rodriguez and Strong, 2012Reported food use
NicaraguaPresentFlora Mesoamericana, 2014; Flora of Nicaragua, 2015
PanamaPresentPanama Checklist, 2014Canal Area, Colón, Panamá, Veraguas, San Blas
Puerto RicoPresentIntroducedAcevedo-Rodriguez and Strong, 2012Cultivated
Trinidad and TobagoPresentEncyclopedia of Life, 2015Cultivated

South America

BoliviaPresentBolivia Checklist, 2015Beni, Santa Cruz, , Cochabamba
BrazilPresentIntroducedForzza et al., 2012Subspontaneous, not endemic. Amazônia, Caatinga, Cerrado, Mata Atlântica
-AcrePresentIntroducedForzza et al., 2012Subspontaneous, not endemic
-AlagoasPresentIntroducedForzza et al., 2012Subspontaneous, not endemic
-GoiasPresentIntroducedForzza et al., 2012Subspontaneous, not endemic
-PernambucoPresentIntroducedForzza et al., 2012Subspontaneous, not endemic
-PiauiPresentIntroducedForzza et al., 2012Subspontaneous, not endemic
-Rio Grande do NortePresentIntroducedForzza et al., 2012Subspontaneous, not endemic
-Santa CatarinaPresentIntroducedForzza et al., 2012Subspontaneous, not endemic
ColombiaPresentIntroducedVascular Plants of Antioquia, 2015Apartadó, Medellín, Turbo
EcuadorPresentIntroducedVascular Plants of Ecuador, 2015Los Ríos
GuyanaPresentMorton, 1987
ParaguayPresentIntroducedRandall, 2012Naturalised
PeruPresent only in captivity/cultivationPeru Checklist, 2015Cultivated. Loreto
SurinamePresentMorton, 1987
VenezuelaPresentMorton, 1987

Oceania

AustraliaPresentPresent based on regional distribution.
-QueenslandPresentMorton, 1987‘popular’
French PolynesiaPresentMorton, 1987; Wagner and Lorence, 2015Tahiti
GuamPresentWagner et al., 2015
Micronesia, Federated states ofPresentIntroducedRandall, 2012; Wagner et al., 2015Naturalised
New CaledoniaPresentMorton, 1987
Northern Mariana IslandsPresentWagner et al., 2015Pagan, Rota, Tinian
TongaPresentHanelt et al., 2001

History of Introduction and Spread

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A. carambola, like A. bilimbi, is generally thought to have originated in tropical Southeast Asia, perhaps Sri Lanka and the Moluccas (Morton, 1987). It is also speculated by some to be native to the Neotropics, which has led to uncertainty over its true origin, especially since it has been cultivated across the Old World tropics for centuries. In the Philippines, for example, Merrill (1904) and Quisumbing (1951) report both A. bilimbi and A. carambola as having been introduced via the galleon trade to the Philippines from tropical America during the Spanish colonial period.

The species is reported to have been introduced into southern Florida, USA, before 1887 (Morton, 1987). Largely viewed as “a curiosity” initially, a variety selected in Florida from seeds introduced from Hawaii was vegetatively propagated during the 1940's and 1950's and, in late 1965, was officially released under the name 'Golden Star' and distributed to growers” (Morton, 1987).

Date of introduction to the West Indies is uncertain. While its close relation A. bilimbi was listed as present in Jamaica by the 1830’s (Macfadyen, 1837), A. carambola was reportedly present by 1851 (Britton and Wilson, 1924) but was not included in Bello’s (1881; 1883) work on Puerto Rico, Britton’s (1918) flora of Bermuda, or Britton’s (1920) work on the Bahamas, suggesting it was not considered a major species during this time. It was, however, in Britton and Wilson’s (1924) flora of Puerto Rico and the Virgin Islands, in which it was reportedly already present in St. Thomas by 1851 and seen by the authors in Puerto Rico in 1915. The species was introduced to Puerto Rico for its fruit and has since become naturalized here, as well as in many other tropical and subtropical regions (Liogier and Martorell, 2000).

Risk of Introduction

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Risk of introduction for this species is low, but further evaluation may be required. As of 2010 the species was considered a non-invasive fruit tree recommended for gardens in Hawaii (Clifford and Kobayashi, 2010), but it was more recently listed in the Global Compendium of Weeds as a weed and cultivation escape (Randall, 2012) and was also on a 2003 list of potential invasive species threats in the Caribbean Region (Kairo et al., 2003). The species reported to be naturalized in Puerto Rico, Belize, Micronesia, and Paraguay (Liogier and Martorell, 2000; Randall, 2012). Its invasiveness is restricted by its slow growth rate, its inability to self-fertilize, limited seed production and limited viability of its seeds (Clifford and Kobayashi, 2010). However, its close relation A. bilimbi is currently flagged for further evaluation by a PIER risk assessment (2015), and it may be possible that A. carambola will need the same treatment in the future.

Habitat

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A. carambola occurs in humid premontane forests and humid-to-very humid tropical forests of Colombia (Vascular Plants of Antioquia, 2015), rainforests as well as savannas in Bolivia (Bolivia Checklist, 2015), in disturbed areas of Peru (Peru Checklist, 2015), and in coastal areas of Ecuador (Vascular Plants of Ecuador, 2015). In China, the species occurs in cultivated areas and, as a cultivation escape, along roadsides and in secondary forests (Flora of China Editorial Committee, 2015).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
 
Terrestrial – ManagedCultivated / agricultural land Present, no further details Productive/non-natural
Managed forests, plantations and orchards Present, no further details Productive/non-natural
Disturbed areas Present, no further details Natural
Disturbed areas Present, no further details Productive/non-natural
Rail / roadsides Present, no further details Natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Present, no further details Natural
Natural forests Present, no further details Productive/non-natural
Natural grasslands Present, no further details
Littoral
Coastal areas Present, no further details Natural
Coastal areas Present, no further details Productive/non-natural

Biology and Ecology

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Reproductive Biology

The heterostylus self-incompatibility (SI) system is a feature of both A. bilimbi and A. carambola. Members of these species have various style lengths such that the stigmas in each morph more or less correspond to the positions of the anthers in the other morphs; this requires multiple morphs of the species, and ensures that trees are not inbred (Knight, 1965; Ganders, 1979; Wong et al., 1994). A. carambola is distylous, with two morphs, and A. bilimbi is tristylous, with three morphs, so when cultivating either species all morphs of that species should be planted to ensure fruit yield (Ganders, 1979; Staples and Herbst, 2005).

Environmental Requirements

Both Averrhoa species - A. bilimbi and A. carambola - prefer a climate with a dry season, but can also adapt to hot humid conditions and wetter climates (FAO EcoCrop, 2015). A. carambola has a wider climate range than fellow genus member A. bilimbi; it is capable of growing within the latitudinal range from 32°N-30°S and can withstand growing in both hot humid tropics and subtropical countries including Egypt and Israel, and can tolerate short periods of freezing temperatures as low as -3 C (Samson, 1991; Bircher and Bircher, 2000). A. carambola prefers well-drained soils ideally between pH 5.5-6.5 but can tolerate pH between 5 and 8.5 (FAO EcoCrop, 2015). It cannot tolerate drought, flooding, or salinity (Samson, 1991; Bircher and Bircher, 2000; FAO EcoCrop, 2015).

The species generally occurs at low elevations. It has been reported in Nicaragua between 0-100 m (Flora of Nicaragua, 2015), up to 500 m in Peru (Peru Checklist, 2015), between 0-500 and 1000-1500 m in Colombia (Vascular Plants of Antioquia, 2015), and at 0-1000 m in Bolivia (Bolivia Checklist, 2015).

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]))
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 Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
32 30

Air Temperature

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Parameter Lower limit Upper limit
Absolute minimum temperature (ºC) -3

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall6003000mm; lower/upper limits

Soil Tolerances

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Soil reaction

  • acid
  • alkaline
  • neutral

Soil texture

  • heavy
  • light
  • medium

Means of Movement and Dispersal

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Natural Dispersal

It is possible that the seeds of A. carambola could be dispersed by water, as the species is known to grow in the coastal areas of Ecuador (Vascular Plants of Ecuador, 2015), but there are no confirmed reports of seed dispersal through this vector.

Vector Transmission (Biotic)

Seeds might be carried or eaten by animals who feed on the fleshy fruit encapsulating the seeds, but there are no confirmed reports of seed dispersal through this vector.

Intentional Introduction

A. carambola has been intentionally introduced in tropical regions around the world for human cultivation, as it is used in food and medicine. It has been the subject of breeding programs and has been proposed as a potential agroforestry species (Morton, 1987; FAO EcoCrop, 2015).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Breeding and propagationHas been the subject of breeding programs and has been proposed as a potential agroforestry species Yes Yes FAO EcoCrop, 2015; Morton, 1987; Samson, 1991
Escape from confinement or garden escape Yes Yes Flora of China Editorial Committee, 2015; Liogier and Martorell, 2000; Randall, 2012
FoodFruits are sold in local markets. Yes Yes Morton, 1987; Quisumbing, 1951; Samson, 1991
Garden waste disposal Yes Flora of China Editorial Committee, 2015; Liogier and Martorell, 2000; Randall, 2012
Medicinal use Yes Yes Morton, 1987; Quisumbing, 1951; Samson, 1991
Ornamental purposes Yes Yes Burger, 1991

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Debris and waste associated with human activitiesSeeds are encapsulated in fleshy, edible fruit which is a food crop Yes Yes FAO EcoCrop, 2015; Samson, 1991

Impact Summary

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CategoryImpact
Environment (generally) Negative
Human health Positive and negative

Impact: Environmental

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A. carambola is known to be a cultivation escape, naturalized, and weedy (Liogier and Martorell, 2000; Randall, 2012), but the extent of its potential environmental damage requires further evaluation.

Impact: Social

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Outside of cultivation, increased populations of A. carambola could positively impact livelihoods of communities, particularly those who rely on local foraging and can harvest fruits of this species for food and sale. However, there is mounting clinical evidence for the toxicity of consuming the fruits, as they contain high levels of oxalic acid which can cause renal failure in humans (Neto et al., 2003).

Risk and Impact Factors

Top of page Invasiveness
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Tolerant of shade
Impact outcomes
  • Negatively impacts human health
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately

Uses

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Starfruit is normally consumed fresh as a dessert fruit. Dessert fruit and minimally processed slices should preferably have a high sugar to acid ratio. Tart fruit are preferred in Taiwan for processing into juice. Starfruit can be consumed as jams, preserves, pickles, candy, juice and liquor. Green fruit are sometimes consumed as a vegetable. Sliced fruit can be added to salads (Janick & Paull, 2008).

Fruits of acid cultivars have been used to clean and polish brass, and the juice used to bleach rust stains from cloth. The wood of A. carambola is white, though older wood is reddish. It is close grained and medium hard, and has been used for furniture and in construction (Morton, 1987).

Uses List

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Human food and beverage

  • Beverage base
  • Food additive
  • Fruits

Materials

  • Dyestuffs

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

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

Bello D, 1883. [English title not available]. (Apuntes para la flora de Puerto Rico. Segunda parte. Monoclamídeas.) Anales de la Sociedad Española de Historia Natural, 12:103-130.

Bello Espinosa D, 1881. [English title not available]. (Apuntes para la flora de Puerto Rico. Primera parte.) Anal. Soc. Española de Hist. Nat, 10:231-304.

Bircher AG, Bircher WH, 2000. Encyclopedia of fruit trees and edible flowering plants in Egypt and the subtropics. Cairo, Egypt: American University in Cairo Press, 596 pp.

Bolivia Checklist, 2015. Catalogue of the Vascular Plants of Bolivia, Tropicos website. St. Louis, MO, USA: Missouri Botanical Garden. http://tropicos.org/Project/BC

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

Britton NL, 1924. Botany of Porto Rico and Virgin Islands. Scientific Survey of Porto Rico and Virgin Islands. New York, USA: New York Academy of Sciences, 200 pp.

Burger WC, 1991. Flora Costaricensis. New Series 28, Publication 1428. Chicago, IL, USA: Field Museum of Natural History. http://archive.org/stream/floracostaricens28fiel/floracostaricens28fiel_djvu.txt

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

Clifford P, Kobayashi K, 2010. Non-Invasive Fruit Trees for Gardens in Hawai'i., USA: College of Tropical Agriculture and Human Resources, University of Hawai'i at Manoa, 10 pp. [Fruit and Nuts F_N-17.] http://www.ctahr.hawaii.edu/oc/freepubs/pdf/F_N-17.pdf

Cook OC, Collins GN, 1903. Economic Plants of Porto Rico. Contributions from the US National Herbarium, Volume 8 Part 2.

Duke J, 2015. Dr. Duke's Phytochemical and Ethnobotanical Databases online resource. Beltsville, USA: Germplasm Resources Information Network (GRIN), National Germplasm Resources Laboratory. http://www.ars-grin.gov/duke/

Encyclopedia of Life, 2015. Encyclopedia of Life. www.eol.org

FAO EcoCrop, 2015. Averrhoa carambola. Eco-Crop Online Database. Land and Water Development Division, Food and Agricultural Organization of the UN (FAO). http://ecocrop.fao.org/ecocrop/srv/en/cropView?id=483

Flora Mesoamericana, 2014. Flora Mesoamericana. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/Project/FM

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

Flora of Nicaragua, 2015. Flora of Nicaragua, Tropicos website. St. Louis, MO, USA: Missouri Botanical Garden. http://tropicos.org/Project/FN

Flora of Pakistan, 2015. Flora of Pakistan/Pakistan Plant Database (PPD). Tropicos website. USA: St. Louis, Missouri and Cambridge, Massachusetts. http://www.tropicos.org/Project/Pakistan

Forzza RC, Leitman PM, Costa AF, Carvalho Jr AA, et al. , 2012. List of species of the Flora of Brazil (Lista de espécies Flora do Brasil). Rio de Janeiro, Brazil: Rio de Janeiro Botanic Garden. http://floradobrasil.jbrj.gov.br/2012/

Ganders FR, 1979. The biology of heterostyly. New Zealand Journal of Botany, 17(4):607-635.

Hanelt P, Buttner R, Mansfeld R, 2001. Mansfeld's Encyclopedia of Agricultural and Horticultural Crops (except Ornamentals). Berlin, Germany: Springer.

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17/03/2015 Original text by:

Marianne Jennifer Datiles, Department of Botany-Smithsonian NMNH, Washington DC, USA

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