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Overview

Brief Summary

Naranjilla or Lulo (Solanum quitoense) is a sub-shrubby perennial with orange fruits slightly larger than a golf ball. The greenish flesh yields a juice with a flavor that has been described as a blend of orange, pineapple, and tomato. The plant is around 2 m tall with purplish stem and leaves. Two varieties of this species have been recognized, quitoense, a spineless form found in southern Colombia and Ecuador, and septentrionale, a form with spines found in central Colombia, Panama, and Costa Rica that is also grown in French Polynesia. Naranjillo is cultivated widely within its range, but also has characteristics of a "weed" and is often considered a species in the process of being domesticated.

Solanum is an enormous genus that includes many well known plants of agricultural importance (e.g., Potato [S. tuberosum], Tomato [S. lycopersicum], and Eggplant [S. melongena]). Within the Solanum subgenus Leptostemonum, Naranjilla belongs to the section Lasiocarpa, a clade that includes around 13 species of perennial shrubs or small trees with its center of distribution in northwestern South America. Several species in this section produce edible fruits and two of them, Naranjilla and Cocona (S. sessiliflorum), are economically important fruit crops in Latin America. Naranjilla has been introduced to Panama, Costa Rica, and Guatemala and is now naturalized in Central America and cultivated in French Polynesia (where there is some concern it could become an invasive weed).

Naranjillo has often been identified as having potential for development as a premium crop for international markets, but some practical hurdles must be overcome, notably the production of varieties with better pathogen resistance. Wild relatives of Naranjilla are a potential source of desirable traits that could be exploited for genetic improvement. For example, crosses of Naranjilla with S. hirtum have reportedly yielded varieties with the same taste as Naranjilla, but with resistance to root knot nematodes.

(Heiser 1985; Bohs 2004; Levin et al. 2006; Bedoya-Reina and Barrero 2010 and references therein)

  • Bedoya-Reina, O.C. and L.S. Barrero. 2010. Preliminary assessment of COSII gene diversity in lulo and a relative species: Initial identification of genes potentially associated with domestication. Gene 458: 27-36.
  • Bohs, L. 2004. A Chloroplast DNA Phylogeny of Solanum Section Lasiocarpa. Systematic Botany 29(1): 177-187.
  • Heiser, C.B., Jr. 1985. Ethnobotany of the Naranjilla (Solanum quitoense) and Its Relatives. Economic Botany 39(1): 4-11.
  • Levin, R.A., N.R. Myers, and L. Bohs. 2006. Phylogenetic relationships among the "spiny Solanums" (Solanum subgenus Leptostemonum, Solanaceae). American Journal of Botany 93(1): 157-169.
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Distribution

Localities documented in Tropicos sources

Solanum quitoense var. septentrionale R.E. Schult. & Cuatrec.:
Colombia (South America)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.
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© Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO 63110 USA

Source: Missouri Botanical Garden

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Localities documented in Tropicos sources

Solanum quitoense Lam.:
Colombia (South America)
Costa Rica (Mesoamerica)
Ecuador (South America)
Honduras (Mesoamerica)
Panama (Mesoamerica)
Peru (South America)
Venezuela (South America)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.
Creative Commons Attribution Non Commercial Share Alike 3.0 (CC BY-NC-SA 3.0)

© Missouri Botanical Garden, 4344 Shaw Boulevard, St. Louis, MO 63110 USA

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Cultivated between 1000 and 2400 m in mountains of Colombia and Ecuador; recently introduced and becoming a successful weed in montane regions of Costa Rica and Panama.

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Physical Description

Diagnostic Description

Formal Description

Habit

Erect or divergently branched, lignescent perennials 1-3 m tall. Stems stellate-woolly, densely pubescent with a mixture of stipitate to essentially sessile stellate trichomes, the multiseriate stalks of stipitate cauline stellae of variable length, the longest 1-4 mm, the lateral rays 6-9, 0.6-1.5 mm long, the midpoints about equaling rays or somewhat shorter, the hairs often purplish; unarmed or prickly, the prickles, when present, small, 1-3 (-5) mm long, 0.5-1.5 mm broad at base, usually somewhat recurved.

Sympodial Structure

Sympodial units 2-foliate, geminate.

Leaves

Leaves simple, the blades 13-50 x 11-40 cm, about as long as wide, elliptic to ovate, thin-textured, felty-pubescent adaxially with sessile stellae, the dominant form with prolonged midpoints 1-2 mm long and 4-6 short, basally inserted lateral rays 0.1-0.4 mm long, also with a distinct class of smaller stellae often scattered among the larger ones, with rays and midpoints 0.1-0.3 mm long, densely pubescent abaxially with interwoven stalked and sessile stellae, the foliar pubescence in vivo with purplish cast, especially along principal veins; unarmed or sparsely prickly on major veins; major lateral veins 5-6 on each side; base truncate or shallowly cordate; margin coarsely dentate with 5-6 deltoid or depressed-deltoid, acute- or obtuse-tipped lobes at lateral vein terminations, the interlobal sinuses entire or occasionally with 1 or 2 smaller, tooth-like lobes; apex acute; petioles 5-15 cm, typically 1/4-1/2 the length of the blades, densely pubescent with hairs like those of the stem, unarmed or sparsely prickly.

Inflorescences

Inflorescences up to 1 cm long, leaf-opposed, unbranched, with 1-24 flowers, the distal flowers mostly female-sterile, the axes densely stellate-pubescent, unarmed or minutely prickly; peduncle 0-0.5 cm; rachis ca. 0.5 cm; pedicels 5-15 mm in flower, 5-15 mm in fruit, closely spaced 2-3 mm apart, articulated at the base.

Flowers

Flowers with the calyx broadly campanulate, 14-20 mm long, the tube 5-8 mm, 8-13 mm wide, the lobes 7-13 x 5-8 mm, ovate to broadly ovate, commonly somewhat petaloid and white, stellate-woolly abaxially; fruiting calyx somewhat accrescent but not completely covering the fruit. Corolla 3-5 (-7) cm in diameter, 20-25 mm long, stellate, the tube 4-6 mm, thin-textured, white, the lobes 10-15 x 5-7 mm, ovate-lanceolate, spreading, acute at apices, densely stellate-pubescent abaxially, glabrous or sparsely pubescent adaxially. Stamens with filaments 0-1 mm; anthers 7-15 x 2-3.5 mm at base, lanceolate, attenuate, more or less connivent or divergent at tips, yellow, the pores small and directed distally. Ovary densely pubescent; style ca. 10 x 0.5-1mm, cylindrical, straight, glabrous; stigma capitate.

Fruits

Fruits 1-4 per inflorescence, 3-6.5 cm in diameter, globose, orange when ripe, with green flesh, spreading-hirsute when young with sessile stellate hairs with with elongate midpoints 3.5-4.5 mm long and 5-15 short, basally inserted lateral rays ca. 0.2 mm long, most trichomes finally deciduous.

Seeds

Seeds numerous, 2.5-4 x 2.5-4 mm, yellow or light tan, lenticular, broadly ovate, the surfaces minutely pitted.

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Type Information

Isotype for Solanum nollanum Britton
Catalog Number: US 1414942
Collection: Smithsonian Institution, National Museum of Natural History, Department of Botany
Verification Degree: Card file verified by examination of original publication
Preparation: Pressed specimen
Collector(s): J. Nolla
Year Collected: 1930
Locality: Rio Piedras., Greater Antilles, Puerto Rico, West Indies
  • Isotype: Britton, N. L. 1930. Sci. Surv. Porto Rico & Virgin Isl. 6: 561.
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© Smithsonian Institution, National Museum of Natural History, Department of Botany

Source: National Museum of Natural History Collections

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Evolution and Systematics

Systematics or Phylogenetics

Phylogeny

Solanum quitoense belongs to the Leptostemonum clade of Solanum (Bohs, 2005). Within Leptostemonum, it belongs to the Lasiocarpa clade, a monophyletic group that includes most of the species traditionally recognized in Solanum section Lasiocarpa Dunal (Whalen et al., 1981; the S. quitoense species group of Whalen, 1984; Levin et al., 2006). Within this clade, chloroplast sequences from the trnT-F region indicate that S. quitoense belongs to a clade that also includes S. candidum, S. hyporhodium, S. lasiocarpum, S. felinum, S. pseudolulo, S. repandum, and S. vestissimum (Bohs, 2004). Resolution is poor to non-existent among the species of this latter clade.

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Molecular Biology and Genetics

Molecular Biology

Molecular Data

nuclear ITS sequence: GenBank AY263460 (Jardín Botanico de Bogotá, Colombia, Olmstead, no voucher) nuclear waxy (GBSSI) sequence: GenBank AY562965 (voucher: Bohs 2873, UT) chloroplast trnS-G sequence: GenBank AY555471 (voucher: Bohs 2873, UT) chloroplast trnT-F sequence: GenBank AY266243 (voucher: Heiser s.n.; Bohs DNA extract 996; plant from Quito, Ecuador, market) http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=33355744 chloroplast trnT-F sequence: GenBank AY266228 (voucher: Bohs 2873, UT) http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=33355729

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Relevance to Humans and Ecosystems

Benefits

Common Names and Uses

Common names: Colombia; lulo, naranjilla. Ecuador: naranjilla. Uses: One of the most esteemed fruit crops in Andean South America; the raw fruits are sweetened to make a delicious juice. The tart yet sweet flavor is very refreshing, and many travelers look forward to partaking of “jugo de lulo.” However, nowadays (in 2005) it is rare, at least in highland Ecuador, to be offered unadulterated lulo juice. Solanum quitoense is susceptible to attack by nematodes and other pests, limiting its cultivation. In recent years its has been hybridized with various species, especially with S. sessiliflorum, with the result that most “lulo” grown and served in Ecuador comes from this hybrid (Heiser et al., 2005). The hybrid can be distinguished from the true lulo by the color of its fruit pulp: pure Solanum quitoense has bright green fruit pulp, whereas the hybrids most often have yellowish or, at most, light greenish fruit pulp.

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Wikipedia

Solanum quitoense

Whole and transversely-cut fruit

Solanum quitoense, known as naranjilla (Spanish pronunciation: [naɾaŋˈxiʎa], "little orange") in Ecuador and Panama and as lulo ([ˈlulo], from Quechua) in Colombia, is a subtropical perennial plant from northwestern South America. The specific name for this species of nightshade means "from Quito."[3]

The naranjilla plant is attractive, with large elongated heart- or oval-shaped leaves up to 45 cm in length. The leaves and stems of the plant are covered in short purple hairs. Naranjilla are delicate plants and must be protected from strong winds and direct sunlight. They grow best in partial shade.

The fruit has a citrus flavour, sometimes described as a combination of rhubarb and lime. The juice of the naranjilla is green and is often used as a drink.

Classification[edit]

Within the genus Solanum, S. quitoense is a part of the subgenus Leptostemonum. Within this clade, S. quitoense belongs to the section Lasiocarpa. Other species within Lasiocarpa include: S. candidum, S. hyporhodium, S. lasiocarpum, S. felinum, S. psudolulo, S. repandum and S. vestissimum.[1]

Naranjilla new leaf

Other plants bear morphological similarity to S. quitoense, but they may or may not be closely related. Some of these plants are: S. hirtum, S. myiacanthum, S. pectinatum, S. sessiliflorum and, S. verrogeneum. Many of these plants, related or not, can be confused with S. quitoense. Furthermore, Solanum quitoense's physical traits vary from plant to plant, making identification challenging: at least three varietals (with spines, without spines, or a third variety known as baquicha, which features red-ripening fruits and smooth leaves) are known to occur. One characteristic that is unique to S. quitoense is the ring of green flesh within the ripe fruit.[1] The only related fruit to have green flesh is a cultivated variant of S. lasiocarpum.

The new growth of this plant is densely covered in protective trichomes. Coloration in the plant's trichomes around the new growth and flowers varies from purple to white. Identification can be difficult for this reason.

The flower of Solanum quitoense.


Agriculture[edit]

The naranjilla has been proposed as a new flavoring for the global food industry,[4] but it fares poorly in large-scale cultivation, presenting an obstacle to its wider use.[1] Its fruit, like tomatoes, is easily damaged when ripe, so is usually harvested unripe.[4] The fruits are found at markets. It is common for locals to make beverages by adding sugar and water to the freshly squeezed fruits.[4]

Rural Costa Rican farmers prepare fruit with salt.

Pests & diseases[edit]

S. quitoense has limited potential in agriculture due to the plant's extreme vulnerability to pests and diseases when grown as a crop. One common type of infection is caused by the root-knot nematode. The ripe fruit is very delicate, and is frequently attacked by fungus, especially when mechanically damaged, so it is often picked unripe to avoid rotting.[4]

Hybrids are an increasingly popular solution to the nematode pest problem. S. quitoense has been hybridized with other plants, most commonly with S. sessiliflorum, a plant with similar phenotypic traits. The leaves, flowers and fruits of S. sessiliflorum are similar in form to S. quitoense, but has much larger fruits that are yellow; the resulting hybrids have fruits with yellowish fruit pulp.[1]

Nutrition[edit]

Unripe fruit flesh.
Ripe fruit flesh.

Contents of the fruit varies from region to region. These statistics are based on Costa Rican fruit:[4]

fruit nutrientspercent contained in fruit
Water90%
Protein1%
Fatless than .0001%
Carbohydrates3.8%
Fiber1.4%
Sugar3%
Calories(kcal/100g) 18
Vitamin C2.6%

These statistics are based on fruits found in Colombia and Ecuador:[5]

Fruit nutrientsmg per 100g of nutrients.
Calcium5.9-12.4 mg
Phosphorus12.0-43.7 mg
Iron0.34-0.64 mg
Carotene0.071-0.0232 mg
Thiamine0.04-0.094
Riboflavin0.03-0.047
Niacin1.19-1.76

References[edit]

  1. ^ a b c d e Solanaceae Source (2005): Solanum quitoense. Version of December 2005. Retrieved 2008-SEP-25.
  2. ^ Tropicos
  3. ^ Lamarck, Jean Baptiste Antoine Pierre de Monnet de. Tableau Encyclopédique et Methodique ... Botanique 2: 16. 1794.
  4. ^ a b c d e Óscar Acosta, Ana M. Pérez, Fabrice Vaillant (2009) Chemical characterization, antioxidant properties, and volatile constituents of naranjilla (Solanum quitoense Lam.) cultivated in Costa Rica. Archivos Latinoamericana de Nutrición 59(1): 88-94
  5. ^ Naranjilla
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References and More Information

Commentary

Commentary taken from Whalen et al. (1981):

Solanum quitoense is cultivated for its edible fruit in highlands of Colombia and Ecuador. It is used predominantly for juice and in preparation of refrescos with sugar and ice. The lulo is one of the most delightful of tropical American fruits, with a flavor reminiscent of citrus, and is rich in Vitamin A and ascorbic acid (Romero-Castañeda, 1961). Cultivation of S. quitoense is restricted by climatic requirements and susceptibility to root-knot nematode infection (Munier, 1962). It is most commonly grown in small plantings on newly cleared land.

Wild occurrences of S. quitoense in Colombia and Ecuador are generally attributed to escape from cultivation. Volunteer plants are seen in disturbances and around habitations in regions where the species is cultivated. For reasons we do not understand, S. quitoense has been most successful as a weed in Costa Rica and Panama, where it was introduced into cultivation in this century. A few collections from montane forests in Colombia may represent native populations (Cuatrecasas et al. 12111 from Norte de Santander and Cuatrecasas 15031 and 22694 from the Cordillera Occidental in Valle). The regions in which these plants were found should be explored more fully as potentially important sources of germ plasm.

Heiser (1969) speculates that S. quitoense was domesticated in highlands of central Colombia and transported by man to Ecuador and Peru. The basis for the argument is the existence of prickly forms and close relatives in the former region. The opposite direction of introduction, into Colombia from Ecuador, is postulated by Patiño (1962) and Schultes and Romero-Casteñeda (1962). Morphological data from Colombian and Ecuadorean plants point to a relative lack of variability in Ecuadorean S. quitoense. The pattern observed may be due to founder effect associated with human dispersal of S. quitoense southward in the Andes from a region of initial domestication in central Colombia as proposed by Heiser.

Heiser (1972) pointed out the close relationship of S. quitoense to S. candidum, a wide ranging species extending from Mexico to Peru. The morphological similarity between these taxa is great, and the differences between them are primarily in characters that would likely have been influenced by human selection during the process of domestication. In particular, S. quitoense is unarmed or only sparsely prickly and has larger fruits with more readily deciduous hairs than S. candidum. An additional difference is the green flesh of S. quitoense fruits, an attribute of that species not found in any other members of section Lasiocarpa.

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