Invasive Species Compendium

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Solanum quitoense
(naranjilla)

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Datasheet

Solanum quitoense (naranjilla)

Summary

  • Last modified
  • 22 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Solanum quitoense
  • Preferred Common Name
  • naranjilla
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • Solanum quitoense is a perennial shrub, native to the montane forests of Colombia and Ecuador. It is widely cultivated for its fruit in areas within and outside its native distribution range. It has escaped fro...

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Pictures

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PictureTitleCaptionCopyright
Solanum quitoense (Narangillo); habit and leaves. Pali o Waipio, Maui, Hawaii, USA. November 2012.
TitleHabit
CaptionSolanum quitoense (Narangillo); habit and leaves. Pali o Waipio, Maui, Hawaii, USA. November 2012.
Copyright©Forest & Kim Starr - CC BY 4.0
Solanum quitoense (Narangillo); habit and leaves. Pali o Waipio, Maui, Hawaii, USA. November 2012.
HabitSolanum quitoense (Narangillo); habit and leaves. Pali o Waipio, Maui, Hawaii, USA. November 2012.©Forest & Kim Starr - CC BY 4.0
Solanum quitoense (Narangillo); flower. Pali o Waipio, Maui, Hawaii, USA. November 2012.
TitleFlower
CaptionSolanum quitoense (Narangillo); flower. Pali o Waipio, Maui, Hawaii, USA. November 2012.
Copyright©Forest & Kim Starr - CC BY 4.0
Solanum quitoense (Narangillo); flower. Pali o Waipio, Maui, Hawaii, USA. November 2012.
FlowerSolanum quitoense (Narangillo); flower. Pali o Waipio, Maui, Hawaii, USA. November 2012.©Forest & Kim Starr - CC BY 4.0
Solanum quitoense (Narangillo); unripe fruit. Pali o Waipio, Maui, Hawaii, USA. November 2012.
TitleFruit
CaptionSolanum quitoense (Narangillo); unripe fruit. Pali o Waipio, Maui, Hawaii, USA. November 2012.
Copyright©Forest & Kim Starr - CC BY 4.0
Solanum quitoense (Narangillo); unripe fruit. Pali o Waipio, Maui, Hawaii, USA. November 2012.
FruitSolanum quitoense (Narangillo); unripe fruit. Pali o Waipio, Maui, Hawaii, USA. November 2012.©Forest & Kim Starr - CC BY 4.0
Solanum quitoense (Narangillo); ripe fruits, whole and sectioned. Note scale.
TitleFruits
CaptionSolanum quitoense (Narangillo); ripe fruits, whole and sectioned. Note scale.
Copyright©Fibonacci/via wikipedia - CC BY-SA 3.0
Solanum quitoense (Narangillo); ripe fruits, whole and sectioned. Note scale.
FruitsSolanum quitoense (Narangillo); ripe fruits, whole and sectioned. Note scale.©Fibonacci/via wikipedia - CC BY-SA 3.0

Identity

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Preferred Scientific Name

  • Solanum quitoense Lam.

Preferred Common Name

  • naranjilla

Other Scientific Names

  • Solanum angulatum Ruiz & Pav.
  • Solanum quitense Kunth
  • Solanum quitoense f. septentrionale (R.E. Schultes & Cuatrec.) D'Arcy

International Common Names

  • English: golden fruit of the Andes; Quito orange
  • Spanish: lulo; lulo de castilla; lulo de perro; naranjilla de Quito
  • French: morelle de Quito; naranjille; orange de Quito

Local Common Names

  • Bolivia: naranjina
  • Colombia: toronjita
  • Dominican Republic: lulito
  • Indonesia: terong kuning
  • Netherlands: gele torong
  • Peru: lulum; naranjita; nuqui

EPPO code

  • SOLQU (Solanum quitoense)

Summary of Invasiveness

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Solanum quitoense is a perennial shrub, native to the montane forests of Colombia and Ecuador. It is widely cultivated for its fruit in areas within and outside its native distribution range. It has escaped from cultivation to establish weedy populations mainly in disturbed open sites, secondary forests and along roadsides and trails. Once established, this species grows as a weed and has the potential to rapidly colonize and dominate large areas in the understory affecting the germination and establishment of native vegetation. Currently it is listed as invasive in the Galapagos Islands and in the Dominican Republic. There is some concern that this species could also become an invasive weed in other tropical areas where it has been introduced.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Solanales
  •                         Family: Solanaceae
  •                             Genus: Solanum
  •                                 Species: Solanum quitoense

Notes on Taxonomy and Nomenclature

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Naranjilla, Solanum quitoense Lam. (syn. Solanum angulatum R. & P.), belongs to the Solanaceae family. Its name means ‘little orange’ and is derived from its round shape and orange colour. In Spanish-speaking countries it is called lulo (Colombia, Peru), naranjilla (Ecuador and most other counties), naranjilla de Quito and naranjilla de Castilla. In English, it is called naranjilla or Quito’s orange; in French, morelle de Quito; and in Dutch, gele terong.

Two varieties of Solanum quitoense are recognized: Solanum quitoense var. quitoense, a spineless form found in southern Colombia and Ecuador and Solanum quitoense var. septentrionale, a form with spines found in central Colombia, Panama and Costa Rica (Heiser, 1972; Solanaceae Source, 2017, The Plant List, 2017).

Related species include the cocona (Solanum sessiliflorum, formerly S. topiro Humb. & Bonpl. and mistakenly called S. hyporhodium) (Martin et al., 1987); the pepino or melon shrub (S. muricatum Alt.); and the tree tomato, named tamarillo in New Zealand to make it more appealing to the export markets (S. betaceum, formerly Cyphomandra betacea Sendt.), which is a native of the Andean highlands, producing in the higher tropics and warm subtropics. Physalis peruviana, the cape gooseberry, is another member of this family that bears edible fruit and has become popular (Paull and Duarte, 2012).

Description

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Naranjilla is a spreading herbaceous shrub of up to 2-3 m. The stems are thick, cylindrical and pubescent, and become woody with age. The alternate leaves are oblong–ovate and up to 60 cm long and 35-45 cm wide, woolly and soft. Spines are present in the stems, petioles, midrib and lateral veins, as well as in the upper and lower parts of the leaf blade. In some types, however, the leaves are spineless. Young leaves, stems and petioles have a coat of purple stellate hairs. The primary root (being of sexual origin) is pivotal but the secondary root system is fairly superficial, not going below 50-60 cm.

The pentamerous fragrant flowers (3-4 cm in diameter) appear in short axillary clusters of five to 10 (sometimes 12) flowers at each axil that open sequentially. The five sepals form a greenish persistent calyx. The five petals are creamy to white on the upper surface and purple and hairy on the lower surface, and form a star-like figure. The flower has five prominent yellow stamens and the stigma is also yellow. Flower buds are covered with purple hairs before opening. Once the plants starts to flower it will do so almost continuously with certain fluctuations, which means that harvesting of cultivated S. quitoense will follow the same pattern. A plant can produce around 1000 flowers during its lifetime, with 5–10% of them setting fruit.  

The fruit is a globose to ovoid berry that is produced in clusters of three to six (Martin et al., 1987). It can measure 4-10 cm in diameter. When mature, the external colour can be yellowish-orange to deep orange and the peel is covered by a brown hairy coat that protects the fruit until fully ripe. These hairs can be fairly hard and are rubbed off after harvest to show the bright orange colour of the smooth, leathery and thick peel. The five-pointed calyx is persistent. Internally the fruit resembles a tomato, with four compartments separated by membranous partitions. These are filled with a translucent, sticky, juicy green or yellowish-green pulp, which has a very pleasant acid flavour and contains numerous (around 1000) pale-buff, thin, flat, greenish-yellow seeds, measuring about 3 mm. Because of its sequential and continuous flowering habit, the plant will simultaneously have flowers and small, medium and ripening fruit. Fruit mature in 50-60 days (Martin et al., 1987). When cultivated fruits are processed, the whole pulp is used leaving only the peel. This differs from cocona, which has a hard ‘hoof ’ and partitions enclosing the juicy pulp with the seeds and has a different flavour and texture (Paull and Duarte, 2012).

The following description is from the Flora of Panama (2017) and Solonaceae Source (2017):

Large herb or small shrub to 3 m tall; twigs stout, tomentose with long-armed, multangulate hairs on long, multiseriate stalks and armed with short or long, stout, flattened, yellowish spines. Leaves large, often exceeding 30 cm long, broadly ovate, shallowly sinuate-lobed, the lobes acuminate, the tips often glabrous above and mucronate, above with a mixture of sessile, porrect, pauci- radiate hairs with long midpoints and long, multiseriate, stalked, multangulate hairs, these latter often deciduous, beneath more densely clothed with mostly stalked stellae and suffused with purple, at least when young, armed or not on the major veins; petioles to 15 cm long, stout, tomentose and often armed. Inflorescence a congested fascicle or sub-umbellate raceme on a short, stout peduncle; pedicels to 10 mm long, the tomentum in part infused with purple. Calyx 10 cm long, lobed 1/2 the way down, the lobes deltoid, densely purplish tomentose outside and, except for the villous tips, glabrous within; corolla white, unequally lobed almost to the base, 5 cm across, densely tomentose outside, glabrous within; anthers subsessile, very stout at the base and narrowly abruptly in the upper 1/3, the slender tips bent outwards to allow the terminal pores to dehisce extrorsely; ovary densely tomentose, style glabrous. Fruits large, to 6 cm across, a yellow-orange, juicy berry, glabrescent but scabridulous until maturity, the pericarp thin, 5-10 mm thick, enclosing large, many-seeded locules; seeds lenticular-discoid, scrobiculate, with narrow but distinct wings, 2.5-4 mm across overall, yellow or light tan.

Plant Type

Top of page Herbaceous
Perennial
Seed propagated
Shrub

Distribution

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S. quitoense is recorded as native in montane forest areas of Colombia and Ecuador (Catalogue of the Vascular Plants of Ecuador, 2017; Solonaceae Source, 2017; USDA-ARS, 2017). However, there is some doubt as to whether the majority of wild occurrences in these countries are truly native or are escapes from cultivation (Solonaceae Source, 2017) The species has been widely introduced for cultivation  in South America, Central America, the Caribbean and South East Asia and has now become naturalized in many places (Jansen et al., 1991; Acevedo-Rodríguez and Strong, 2012; Bolivia Catalogue, 2015; USDA-ARS, 2017). It is a particularly successful weed in Costa Rica and Panama (Solonaceae Source, 2017) and is also cultivated in French Polynesia and Guam (PIER, 2017). It is recorded as invasive in the Dominican Republic and on the Galapagos Islands (Mir, 2012; PIER, 2017).

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.

Last updated: 10 Jan 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Asia

IndonesiaPresentIntroducedJansen et al. (1991)Cultivated
PhilippinesPresentIntroducedJansen et al. (1991)Cultivated

North America

Costa RicaPresentIntroducedNaturalizedUSDA-ARS (2017)Cultivated and naturalized
CubaPresentIntroducedAcevedo-Rodríguez and Strong (2012)Cultivated
Dominican RepublicPresentIntroducedInvasiveMir (2012)
GuatemalaPresentIntroducedUSDA-ARS (2017)Cultivated
HondurasPresentIntroducedMolina R. (1975)Cultivated
NicaraguaPresentIntroducedMissouri Botanical Garden (2017)Cultivated
PanamaPresentIntroducedNaturalizedUSDA-ARS (2017)Cultivated and naturalized
Puerto RicoPresentIntroducedAcevedo-Rodríguez and Strong (2012)Escaped from cultivation

Oceania

French PolynesiaPresentIntroducedPIER (2017)Cultivated
GuamPresentIntroducedPIER (2017)

South America

BoliviaPresentIntroducedBolivia Catalogue (2015)Cultivated
ColombiaPresentNativeNaturalizedUSDA-ARS (2017)Also cultivated and naturalized
EcuadorPresentNativeCABI (Undated); USDA-ARS (2017)Bolívar, Carchi, Chimborazo, Esmeraldas, Galapagos, Imbabura, Loja, Napo, Pichincha, Tungurahua; also cultivated; Original citation: Catalogue of the Vascular Plants of Ecuador (2017)
-Galapagos IslandsPresentIntroducedInvasivePIER (2017); Jaramillo Díaz et al. (2018)
PeruPresentIntroducedUSDA-ARS (2017)Cultivated
VenezuelaPresentIntroducedCABI (Undated)Cultivated; Original citation: Hokche et al. (2008)

History of Introduction and Spread

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S. quitoense has been introduced to tropical and subtropical regions of the world as a fruit crop. In the 1950s it was introduced to Panama and Costa Rica and in 1963 to Guatemala (Morton, 1987). According to Morton (1987), in the early 1950s, plantings were made in Puerto Rico, Jamaica, Hawaii, Queensland and Costa Rica where one of several growers set out 70,000 plants. In recent years, it has been introduced to South East Asia, mainly in Indonesia and the Philippines (Jansen et al., 1991). On the Galapagos Islands, S. quitoense was probably introduced by settlers and now it can be found naturalized in the wild and is spreading (Charles Darwin Foundation, 2008).

Risk of Introduction

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The risk of new introductions of S. quitoense is high. This species is one of the most appreciated fruit crops in the Andean region of South America. The plant is cultivated for its fruit across South America and, in recent years, its cultivation has spread to many other tropical and subtropical areas including Central America, the Caribbean and South East Asia (Jansen et al., 1991; FAO, 2017).

Habitat

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Within its native distribution range in the Andes and Amazonian regions of Ecuador and Colombia, S. quitoense grows in montane forests at elevations up to 2500 m (Catalogue of the Vascular Plants of Ecuador, 2017). In Bolivia, it is cultivated in the Andean region at elevations ranging from 1500 m to 2000 m (Bolivia Catalogue, 2017). In the Galapagos Islands, it has escaped from cultivation and is now naturalized along trails and in coastal areas (Charles Darwin Foundation, 2008). In Central America it has become naturalized in montane forests, humid hills, rainforests, cloud forests, along roadsides and in disturbed sites near villages at elevations between 950 m and 1650 m (Bohs, 2015; Missouri Botanical Gardens, 2017). In Puerto Rico, it is naturalized in the montane forest of Toro Negro (Liogier and Martorell, 2000).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Present, no further details Productive/non-natural
Disturbed areas Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Natural
Disturbed areas Present, no further details Productive/non-natural
Rail / roadsides Present, no further details Harmful (pest or invasive)
Rail / roadsides Present, no further details Natural
Rail / roadsides Present, no further details Productive/non-natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Present, no further details Harmful (pest or invasive)
Natural forests Present, no further details Natural
Natural forests Present, no further details Productive/non-natural
Scrub / shrublands Present, no further details Harmful (pest or invasive)
Scrub / shrublands Present, no further details Natural
Scrub / shrublands Present, no further details Productive/non-natural
Littoral
Coastal areas Present, no further details Harmful (pest or invasive)
Coastal areas Present, no further details Natural
Coastal areas Present, no further details Productive/non-natural

Biology and Ecology

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Soil

Naranjilla requires light, fertile, well-drained soils (Martin et al., 1987) that are rich in organic matter (ideally >10%). It will grow well in loamy soils or sandy and clay loams.

Climate

TEMPERATURE In the oriental parts of the Andes this crop does well at elevations of 1600-2400 m with the optimum being 1800 m; the acid type can be grown above 2000 m. At higher latitudes the altitude can be lower. The temperature range for this crop is 16-24°C, with an optimal temperature of 17-18°C. It therefore does well on the central coast of Peru, which has a subtropical climate with cool winters and not excessively hot summers. This species requires cooler locations than cocona, although both can be grown in areas where the climatic conditions for both species overlap.

LIGHT AND PHOTOPERIOD Naranjilla tolerates certain amounts shade. In many cases, especially in the Amazonia, it is planted under partial shade given by trees that are left after clearing the forest. The plant can also grow and adequately produce under full sun. No specific information is available about photoperiodic requirements, but since it flowers almost continuously there doesn't appear to be any.

WIND There is not much information on the wind tolerance of naranjilla. Being a rather small plant with a well-anchored root system, it should not be very susceptible to uprooting by winds. However, its large and soft leaves will probably be severely damaged. 

Genetics

The chromosome number reported for S. quitoense is 2n = 24 (Bernardello et al., 1994). Hybrids of S. quitoense with S. hirtum, S. tequilense and S. vestissimum have been developed (Heiser, 1972) and currently most ‘granadilla fruit’ grown and commercialized in Ecuador comes from hybridization between S. quitoense × S. sessiliflorum (Heiser et al., 2005). Hybrids can be distinguished from the wild form of S. quitoense by the colour of their fruit pulp. S. quitoense has bright green fruit pulp, whereas hybrids most often have yellow or light green fruit pulp (Heiser et al., 2005).

Reproductive biology

All Solanum species have poricidally dehiscent anthers that result in buzz pollination syndrome, where pollen can only be released by particular bee species. This pollination syndrome can be found in about 200 genera of flowering plants (Buchmann, 1983). Naranjilla flowers are hermaphrodite and can be self-fertile (Martin et al., 1987) or show allogamy, depending on insects for pollination. Flowers can have short, medium or long styles, and the last two types can be cross-pollinated. Pollination is performed by insects such as honey-bees and bumblebees (Paull and Duarte, 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]))
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])

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
25 25

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 17 25
Mean maximum temperature of hottest month (ºC) 30

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall1500>2500mm; lower/upper limits

Rainfall Regime

Top of page Bimodal
Uniform

Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • alkaline
  • neutral

Soil texture

  • light
  • medium

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Meloidogyne Pathogen Other/All Stages not specific
Pseudaulacaspis pentagona Pathogen Other/All Stages not specific

Notes on Natural Enemies

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S. quitoense is susceptible to attack by root knot nematodes (Meloidogyne species) and by the white scale Pseudaulacaspis pentagona (Morton, 1987).

Means of Movement and Dispersal

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S. quitoense spreads by seed. Seeds can be dispersed by birds and mammals (e.g. cattle, deer, feral pigs etc.). In cultivation, it can be propagated by air-layering or by cuttings (Morton, 1987; FAO, 2017).

Intentional introduction

S. quitoense has been widely introduced for cultivation (Morton, 1987; FAO, 2017).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Crop productionFruit crop Yes Yes USDA-ARS, 2017
Disturbance Yes Yes Heiser, 1972
Escape from confinement or garden escape Yes Yes Charles Darwin Foundation, 2008
FoodFruit crop Yes Yes USDA-ARS, 2017

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Debris and waste associated with human activitiesAssociated with cultivation Yes Yes USDA-ARS, 2017

Environmental Impact

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S. quitoense is an invasive species with the potential to outcompete native plant communities. It grows as a weed and may dominate large areas in the understory affecting the germination and establishment of native species. Currently, it is listed as invasive in the Galapagos Islands and Dominican Republic. There is concern that this species could become an invasive weed in areas where it is now cultivated, such as other islands in the Caribbean, French Polynesia and areas in Central and South America (Heiser, 1972; Liogier and Martorell, 2000; Charles Darwin Foundation, 2008; Mir, 2012; PIER, 2017).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Long lived
  • Fast growing
  • Gregarious
  • Reproduces asexually
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Monoculture formation
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - shading
  • Rapid growth
  • Rooting
  • Produces spines, thorns or burrs
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately

Uses

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S. quitoense is cultivated and commercialized for its fruit, which is used mainly for preparing refreshing drinks. The ripe fruit is usually placed in boiling water for a short time to soften the pulp, after which it is split into two by cutting at the equatorial line and squeezed so that only the peel remains. The balance is blended with chilled water and sugar, strained to eliminate the seeds and served. The peel and seeds constitute around 50% of the total fruit weight. The juice oxidizes readily and turns brown. Even canned juice with 0.1% ascorbic acid added and pasteurized at 92°C for 75 seconds will eventually lose its characteristic colour and flavour. Naranjilla pulp is used in yoghurt and ice cream and is used as an instant soluble powder for preparing juice. Frozen pulp is exported, especially to the USA, as concentrate with 34% total soluble solids. It is quickly frozen after using a rotary evaporator at 35°C. Prepared juice is exported in this form as it retains its appearance for a long time (Paull and Duarte, 2012).

Uses List

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Human food and beverage

  • Beverage base
  • Food additive
  • Fruits

Prevention and Control

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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.

There is no information available regarding the control of S. quitoense, however, for other Solanum species that have become invasive (e.g. S. viarum and S. tampicense) herbicides such as triclopyr, 2,4-D, aminopyralid and picloram have been recommended and repeated applications of herbicides are needed to kill any re-growth and/or new seedlings that emerge (Mullahey et al., 1996; Call et al., 2000).

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. Washington DC, USA: Smithsonian Institution.1192 pp. http://botany.si.edu/Antilles/WestIndies/catalog.htm

Acosta, Ó., Pérez, A. M., Vaillant, F., 2009. Chemical characterization, antioxidant properties, and volatile constituents of naranjilla (Solanum quitoense Lam.) cultivated in Costa Rica. Archivos Latinoamericanos de Nutrición, 59(1), 88-94.

Barbeau, G., 1990. Dirección General de Técnicas Agropecuarias, MIDINRA, Editorial Ciencias Sociales, Managua, Nicaragua, pp. 155–159

Bernardello, L. M., Heiser, C. B., Piazzano, M., 1994. Karyotypic studies in Solanum section Lasiocarpa (Solanaceae). American Journal of Botany, 81(1), 95-103. doi: 10.2307/2445568

Bohs LA, 2015. Solanaceae. In: Manual de Plantas de Costa Rica. Vol. VIII. Monographs in Systematic Botany from the Missouri Botanical Garden , 131 [ed. by Hammel BE, Grayum MH, Herrera C, Zamora N]. 205-336.

Bolivia Catalogue, 2015. Bolivia Catalogue. St. Louis, Missouri, USA: Missouri Botanical Garden.http://www.tropicos.org/Project/BC

Buchmann SL, 1983. Buzz pollination in angiosperms. In: Handbook of experimental pollination biology, [ed. by Jones CE, Little RJ]. New York, USA: Van Nostrand Reinhold Company. 73-133.

Call, N. M., Coble, H. D., Perez-Fernandez, T., 2000. Tropical soda apple (Solanum viarum) herbicide susceptibility and competitiveness in tall fescue (Festuca arundinacea). Weed Technology, 14(2), 252-260. doi: 10.1614/0890-037X(2000)014[0252:TSASVH]2.0.CO;2

Casierra-Posada, F., García, E. J., Lüdders, P., 2004. Determining the proper time for picking lulo fruit (Solanum quitoense Lam. var. quitoense and septentrionale). Agronomía Colombiana, 22(1), 32-39.

Catalogue of Vascular Plants of Ecuador, 2017. Catalogue of Vascular Plants of Ecuador. St. Louis, Missouri, USA: Missouri Botanical Garden.http://tropicos.org/Project/CE

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

Diaz, J. G., Manzano, J. E., 2002. Quality of lulo (Solanum quitoense L.) stored at different temperatures. Proceedings of the Interamerican Society for Tropical Horticulture, 46, 27-28.

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

Flora of Panama, 2017. Flora of Panama (WFO), Tropicos website. St. Louis, MO and Cambridge, MA, USA: Missouri Botanical Garden and Harvard University Herbaria.http://www.tropicos.org/Project/FOPWFO

Gallozzi, R., Duarte, O., 2007. Naranjilla (lulo) y cocona. Guía práctica de manejo agronómico, cosecha, poscosecha y procesamiento. Como parte de: Cultivos de Diversifi cación para pequeños productores de frijol y maíz en América Central, Producido por el IICA y COSUDE. San José, Costa Rica

Heiser CB, 1972. The relationships of the naranjilla, Solanum quitoense. Biotropica, 4, 77-84.

Heiser, C., Soria, J., Miller, C., Anderson, G., 2005. A new synthetic allopolyploid naranjilla, Solanum indianense (Solanaceae). Novon, 15(2), 290-292.

Hokche O, Berry PE, Huber O, 2008. Nuevo Catálogo de la Flora Vascular de Venezuela. [ed. by Hokche O, Berry PE, Huber O]. Caracas, Venezuela: Fundación Instituto Botánico de Venezuela.589 pp.

Janick, J., Paull, R. E., 2008. The encyclopedia of fruit & nuts, CABI.xviii + 954 pp.

Jansen PCM, Jukema J, Oyen LPA, van Lingen TG, 1991. Solanum quitoense Lam. In: PROSEA Plant Resources of South-East Asia Vol. 2, [ed. by Verheij EWM, Coronel RE]. Wageningen, Netherlands: Pudoc. 446 pp.

Jaramillo Díaz P, Guézou A, Mauchamp A, Tye A, 2018. CDF Checklist of Galapagos Flowering Plants. (FCD Lista de especies de Plantas con flores Galápagos). In: Charles Darwin Foundation Galapagos Species Checklist (Lista de Especies de Galápagos de la Fundación Charles Darwin) [ed. by Bungartz F, Herrera H, Jaramillo P, Tirado N, Jiménez-Uzcátegui G, Ruiz D, Guézou A, Ziemmeck F]. Puerto Ayora, Galapagos, Ecuador: Charles Darwin Foundation.

Liogier HA, Martorell LF, 2000. San Juan, Puerto Rico: La Editorial, University of Puerto Rico.382 pp.

Martin, F. W., Campbell, C. W., Ruberté, R. M., 1987. Agriculture Handbook, USDA, (No.642) . 247pp.

Mir C, 2012. Estrategia Nacional de Especies Exóticas Invasoras Realizado en el marco del Proyecto “Mitigando las amenazas de las especies exóticas invasoras en el Caribe Insular”. Dominican Republic: Ministerio de Medio Ambiente y Recursos Naturales Santo Domingo.

Missouri Botanical Garden, 2017. Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden.http://www.tropicos.org/

Molina R., A., 1975. A list of the plants of Honduras. (Enumeracion de las plantas de Honduras). Ceiba, 19(1), 1-118.

Morton, J. F., 1987. J.F. Morton.517 pp.

Mullahey, J. J., Mislevy, P., Brown, W. F., Kline, W. N., 1996. Tropical soda apple, an exotic weed threatening agriculture and natural systems. Down to Earth (Midland), 51(1), 10-17.

Paull, R. E., Duarte, O., 2012. Tropical fruits, Volume 2, (Ed.2) : CABI.ix + 371 pp. http://www.cabi.org/cabebooks/ebook/20123357661 doi:10.1079/9781845937898.0000

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

Rodriguez , V. Arce, Soria Idrovo, N., Heiser, C., 1994. Hybrid 'Palora', a new option as a naranjilla (Solanum quitoense) cultivar in Ecuador. Proceedings of the Interamerican Society for Tropical Horticulture, 38, 194-196.

Solonaceae Source, 2017. http://solanaceaesource.org/

The Plant List, 2017. London, UK: Royal Botanic Gardens, Kew.http://www.theplantlist.org

USDA-ARS, 2017. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, USA: National Germplasm Resources Laboratory.http://www.ars-grin.gov/cgi-bin/npgs/html/tax_search.pl

Villachica, H., Carvalho, J. E. U., Muller, C. H., Diaz, S. C., Almanza, M., 1996. In: Frutales and Hortalizas Promisorios de la Amazonía. Tratado de Cooperación Amazónica, Lima, Peru, pp. 203–207

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

Bolivia Catalogue, 2015. Bolivia Catalogue., St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/Project/BC

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

Jansen PCM, Jukema J, Oyen LPA, van Lingen TG, 1991. Solanum quitoense Lam. In: PROSEA Plant Resources of South-East Asia Vol. 2, [ed. by Verheij EWM, Coronel RE]. Wageningen, Netherlands: Pudoc. 446 pp.

Jaramillo Díaz P, Guézou A, Mauchamp A, Tye A, 2018. CDF Checklist of Galapagos Flowering Plants. (FCD Lista de especies de Plantas con flores Galápagos). In: Charles Darwin Foundation Galapagos Species Checklist (Lista de Especies de Galápagos de la Fundación Charles Darwin), [ed. by Bungartz F, Herrera H, Jaramillo P, Tirado N, Jiménez-Uzcátegui G, Ruiz D, Guézou A, Ziemmeck F]. Puerto Ayora, Galapagos, Ecuador: Charles Darwin Foundation.

Mir C, 2012. [English title not available]. (Estrategia Nacional de especies exóticas invasoras realizado en el marco del Proyecto “Mitigando las amenazas de las especies exóticas invasoras en el Caribe Insular”)., Dominican Republic: Ministerio de Medio Ambiente y Recursos Naturales Santo Domingo.

Missouri Botanical Garden, 2017. Tropicos database. In: Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

Molina R A, 1975. A list of the plants of Honduras. (Enumeracion de las plantas de Honduras.). Ceiba. 19 (1), 1-118.

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

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

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16/06/17 Updated by:

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

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