Solanum mammosum
(nipplefruit nightshade)

Pictures

Identity

Preferred Scientific Name

• Solanum mammosum L.

Preferred Common Name

• nipplefruit nightshade

Other Scientific Names

• Solanum corniculatum Huber
• Solanum globiferum Dunal
• Solanum mammiforme C.C.Robin
• Solanum platanifolium Sendtn.

International Common Names

• English: apple of Sodom; breastberry; cow's udder; fox face; love apple; macaw bush; macawbush; Mickey Mouse plant; nipple fruit; nipple nightshade; pigs ears; pigs'-ears; tit fruit; tit plant; titty fruit; turkey berry; zombie fruit
• Spanish: berenjena de cucarachas; berenjena de gallina; berenjena de teta; berenjenade marimbo; gurito; tetilla
• French: morelle á fruit ornemental; morelle molle; poire de bachelier; pomme a chauve-souris; pomme d'amour; pomme teton; pomme zombi; tétons de jeune fille
• Chinese: huang jin guo; niu tou qie; ru qie; wu dai tong tang; wu jiao qie; wu zhi qie

Local Common Names

• : sigalu
• Bolivia: ku'bu're; pimento; popatoa; tetilla; toro torito; vaca vaquita
• Brazil: jurubeba-do-pará; peito-de-moca
• Brunei Darussalam: teron susu
• Colombia: friega platos; hoja de lun; ku-ku-na; lulo de perro; lulo de perro tetudo; lulu de teta; pepito; tetilla
• Congo Democratic Republic: babua; gbangalangala
• Costa Rica: pichichinchivo; pichichio
• Cuba: guirito; guirito espinoso; pantomina; pechito; pecho de doncella
• Dominican Republic: berenjena de gallina; berenjena de teta; calabacita; téton
• Ecuador: atallpajanbina; chucho muyo; cura gallinas; estacudo; jajapa; jarjapa de montana; mutondo puga; pinon; regargar; rejalgar; teta de vaca
• El Salvador: chichimosa; chichita
• Germany: Euter-Nachtschatten; Zitzen Nachtschatten
• Guadeloupe: pomme poison
• Guatemala: chicha; chichigua; kekchi rantu; tetereta
• Guyana: jumby bubby; tuwawa
• Haiti: amourette batarde; morelle molle; tentation neg'sotte; tetan; tété jeune fille; tétin jeune fille
• Honduras: chichigua; chichiua
• Indonesia/Sumatra: terung susu; tioeng londo
• Jamaica: bachelor's pear; young girls breast
• Japan: kanaria nasu
• Lesser Antilles: lady nipples
• Malaysia: terung asam; terung balanda; terung puyuh; terung susu
• Malaysia/Peninsular Malaysia: terung susu kambing
• Malaysia/Sabah: terong semangat
• Malaysia/Sarawak: terung tujang
• Mexico: berenjena; kan'tzu
• Nicaragua: chichigua
• Panama: moza; nipple fruit moza; tetilla; tetilla teta de China; una de gato; veninillo tetón
• Peru: cocona venenose; teta de vaca; tintuma; vaca chucho
• Philippines: berenjenita peluda; susu; tagotong; talong; utong
• Puerto Rico: berengena cimarrona; berenjena de cucarachas; berenjena de marimba; pecho de doncella
• Samoa: lapiti; lau lau faiva
• Suriname: baladona; njoenwenkebobi; njoen-wenke-bobi; nkjoeng wintje bobi; nyun wentje bobi; terong soesoe
• Sweden: karingtomat
• Turkey: sofur
• Venezuela: tuna; una de gato
• Vietnam: ca vu de

Summary of Invasiveness

S. mammosum is an annual or tender perennial plant with poisonous fruit, reported as invasive in Cuba (Oviedo Prieto et al., 2012) as well as in the Philippines (Merrill, 1923), Fiji, Tonga, and parts of Hawaii (Space and Flynn, 2002; PIER, 2014). It is listed as an ‘agricultural weed’, ‘causal alien’, ‘naturalised’ and ‘weed’ in the Global Compendium of Weeds (Randall, 2012). The species is considered native to Central and South America (Acevedo-Rodriguez and Strong, 2012) and possibly the Caribbean (Nee, 1979; PBI Solanum Project, 2014), but has been introduced to the Old World tropics for ornamental use as well as medicinal and food purposes (Lim, 2013), and has potential to escape cultivation (PBI Solanum Project, 2014). It is an annual or short-lived perennial (PBI Solanum Project, 2014) that reproduces by seeds, which may be dispersed by both biotic and abiotic factors, as they are encased in bright yellow-coloured, large, spongy fruit that can float on water (Chiarini and Barboza, 2007; 2009). The species is a known weed in Jamaica (Holm et al., 1979), Java (Randall, 2012), and Florida (Medal et al., 2009), and is spontaneous where it occurs in Brazil (Forzza et al., 2010). 

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

Solanaceae, the Nightshade family, consists of 90 genera and 3000-4000 species with great variation in habit and distribution on all continents except Antarctica, with the majority of species diversity in Central and South America (PBI Solanum Project, 2014).

Solanum is one of the largest genera of vascular plants with 1000-1500 species, around 1000 of which are speculated to be of American origin (Hunziker, 1979). Taxonomy of the genus and its seven subgenera have undergone many revisions, but the overall genus consists of herbs, shrubs, trees, or herbaceous or woody vines, usually with spines or prickles, glabrous or pubescent with simple or stellate hairs (Acevedo-Rodriguez, 1996). The Solanum genus includes the wild potato, S. tuberosum, the tomato, S. lycopersicum, and the eggplant, S. melongena, with many other members cultivated for medicinal and ornamental uses. While the etymology of the genus’ scientific name is unclear, it may be derived from the Latin word “sol”, meaning "sun," referring to its affinity for sunlight, or from the Latin word “solare”, meaning "to soothe”, the Latin word “solamen”, meaning "a comfort", or the Akkadian word “sululu”, meaning “happy”, in reference to the narcotic effects of some Solanum species after ingestion (Smith, 1971; Wiart, 2006; Quattrocchi, 2012; NZPCN, 2014).

The genus Solanum has been divided into seven subgenera, which are further divided into sections and subsections. S. mammosum belongs to the prickly Solanum subgenus Leptostemonum (Nee, 1991) and the section Acanthophora. This section includes 19 species characterized by prickly stems, lobed or dented prickly leaves with only simple hairs on the upper surface, and a chromosome number 2n=24 (22 in S. mammosum L.) (Cuda et al., 2002). Nee (1979) further divided Acanthophora into two subsections based on seed morphology, due to the unique winged seed structure of several species; however, the seeds of S. mammosum are the lenticular shape of most other Solanum species and do not have wings (Nee, 1979). Both the species name mammosum and its common name nipplefruit nightshade refer to the peculiar shape of its fruits (Smith, 1971; Wiart, 2006).

The name Solanum mammosum was also invalidly given to other nightshade species: S. mammosum as described by J.A. Pavón Jiménez based on Dunal in de Candolle refers to S. circinatum. S. mammosum as described by W. Herbert based on Dunal in de Candolle is the S. torvum of O.P. Swartz (PBI Solanum Project, 2014).

Description

Herb or shrub to 1.5 m tall; twigs viscid-villous with long simple hairs, armed with yellow or green, acicular spines which in age become flattened, recurved, woody, 2.5 cm long; stems fistulose. Leaves 6-20 cm long, broadly ovate, pin- natifidly lobed, the base truncate to deeply cordate, both sides villous with long, simple, glandular hairs, short-stalked glands and sessile, long-radiate stellae with ascending radii, armed with flattened acicular spines to 3 cm long; petioles 3-7 cm long, sparingly armed. Inflorescence lateral, several-flowered, a short raceme on a short (to 10 mm long) peduncle; pedicels 5-12 mm long, becoming stout and somewhat longer in fruit, viscid-villous to lanate, armed or not. Flowers with the calyx unarmed, deeply lobed, the lobes lanceolate, pubescent outside, 3-6 mm long; corolla violet, showy, exceeding the calyx 2-3 times in bud, 3-4 cm across, lobed about halfway; filaments very short, the anthers 10-12 mm long, linear-oblong and tapering. Fruit orange or yellow, 4-7 mm long, ovoid, often with a 2 cm long, nipple-like apex, and sometimes.bearing one or more nipple-like protrusions at the base; seeds compressed-lenticular, 5-7 mm across, not winged, finely rugose, shiny [Flora of Panama, 2014].

The fruits of this species are globulose in Colombia and Venezuela, but the more widely cultivated plants are less prickly and bear fruits shaped with peculiar nipple-like protrusions (Hanelt et al., 2001). These large fruits can reach up to 5.5 cm in diameter and possess a spongy mesocarp (Chiarini and Barboza, 2009) which may be an adaptation for dispersal by water (Nee, 1979; 1991; Chiarini and Barboza, 2007). They are usually a vibrant yellow; colour has been attributed to the dispersal syndrome and brightly coloured fruits would be more attractive to birds (Chiarini and Barboza, 2009). 

Plant Type

Distribution

S. mammosum is considered native to Mexico, Central and South America (Acevedo-Rodriguez and Strong, 2012) and possibly the Caribbean (Nee, 1979; PBI Solanum Project, 2014). According to the PBI Solanum Project (2014), “the morphological and geographical reasons for considering this species native to northern South America and possibly the Caribbean have been discussed previously (Nee, 1979). Briefly, the globose fruit form is undoubtedly the primitive condition and is nearly confined to the Llanos of Venezuela and adjacent areas where the species seems well-adapted to the seasonally dry grassland climate. It is only rare and sporadic in Brazil south of the Amazon basin, the center of diversity of the section”. In Panama, the species is both cultivated and wild (Panama Checklist, 2014). In India, the species was reportedly under experimental introduction as of 1979 (Deb, 1979), although it had already been in country since at least 1968 (Madhavadian, 1968).

The species’ reported native status differs between sources. In the Caribbean, Broome et al. (2007) and USDA-ARS (2014) report the species as native to the following places, although it is introduced according to Acevedo-Rodriguez and Strong (2012): Antigua, Barbados, Cuba, Dominica, Guadeloupe, Hispanola, Jamaica, Martiniqua, Puerto Rico, St. Lucia, St. Vincent and Grenadines. S. mammosum is reported as native to Colombia by USDA-ARS (2014) but as a naturalized species by the Vascular Plants of Antioquia (2014). It has been sporadically introduced elsewhere; rare in Africa, more common in the East Indies (PBI Solanum Project, 2014).

Distribution Table

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.

History of Introduction and Spread

According to Nee (1979), S. mammosum originated in South America but may have also been native to the Caribbean. The toxicity of the species has been used as an insecticide and to catch fish in many places including Costa Rica, Venezuela, Peru, and El Salvador, and Nee suggests the species was dispersed for these reasons with man across South and Central America (Nee, 1979; PBI Solanum Project, 2014). If the species was not native to Suriname, it was present there by 1705, as it was illustrated by Maria Merian and included in her work ‘Metamorphosis Insectorum Surinamensium’ (Stearn, 1982). The species’ native status in the Caribbean is uncertain, but it is considered an introduced species by Acevedo-Rodriguez and Strong (2012). In Trinidad it was certainly being grown in cultivation by 1870 (Prestoe, 1870), and was in Puerto Rico by 1881, as it was included in Bello’s Flora of Puerto Rico (Bello Espinosa, 1881). By 1911 at the latest the species was present in Cuba, Jamaica, Hispaniola, St. Croix, Antigua, Guadeloupe, Dominica, Martinique, St. Vincent, Barbados, and continental tropical America, as Urban reported in his work on the West Indies (Urban, 1903-1911). In 1918, Britton recorded the species growing on St. Croix in his work on the Virgin Islands (Britton, 1918). It has now been introduced from the New World tropics to Asia and Africa, and some parts of North America. 

Risk of Introduction

Risk of introduction of S. mammosum is potentially high, but more research is needed on the extent of the species’ potential damage to native ecosystems. It is not yet included on the IUCN’s Invasive Species Specialist Group (ISSG) list, and has not yet been assessed for its potential invasive risk in the Asia-Pacific region by PIER, nor for its US Invasive Species Impact Rank (I-Rank) by NatureServe (2014). However the species is known to be invasive in some places beyond its native range (Merrill, 1923; Oviedo Prieto et al., 2012; PIER, 2014) and has potential to escape cultivation (PBI Solanum Project, 2014). Despite observing none of this species on Samoa, Space and Flynn (2002) included it in their list of species that pose an invasive threat to Samoa; as they note, the species is already present in the nearby islands Fiji and Tonga. The fruits have the ability to be carried by water due to their spongy mesocarps (Nee, 1979; 1991; Chiarini and Barboza, 2007; 2009) and also attract birds with the bright yellow coloured fruits (Chiarini and Barboza, 2009). Considering its potential for intentional and unintentional dispersal by both biotic and abiotic factors, the probability of invasion may rise in areas near its cultivation.

Habitat

S. mammosum is a weedy shrub of grasslands, pastures, roadsides, waste places, secondary growth and cultivated land, and is confined entirely to the tropics with at least seasonally heavy precipitation, mostly from sea level to 100 m elevation but reaching at least 1800 m (PBI Solanum Project, 2014). The species is known to be an agricultural weed in sugarcane fields of Java (Randall, 2012). In Samoa, the species is rare in pastures on central Upolu (PIER, 2014). In the Philippines, the species occurs in thickets and waste places along roads at low altitudes (Merrill, 1923; Pelser et al., 2014). In Ecuador, it occurs along the coastal and Amazonian regions (Vascular Plants of Ecuador, 2014), while in Nicaragua, it is uncommon but found near homes and in very disturbed areas (Flora of Nicaragua, 2014). Likewise in Peru, S. mammosum is found in disturbed areas (Peru Checklist, 2014). In Venezuela, the species is found in seasonally dry grasslands (PBI Solanum Project, 2014). It is also cultivated in garden settings around the world as an ornamental.

Habitat List

Hosts/Species Affected

Although it is frequently used as an ornamental, S. mammosum is known to be poisonous to mammals (USDA-ARS, 2014) and is included in the FDA Poisonous Plant Database (US-FDA, 2014).

Biology and Ecology

Genetics

The chromosome count for S. mammosum is 2n=22 (Cuda et al., 2002).

Physiology and Phenology

This species is included on Kew’s list of ‘difficult species’; according to Kew’s Difficult Seeds project (2014), the seeds of S. mammosum may be physiologically dormant and resist germination.

Environmental Requirements

S. mammosum prefers full sun and well-drained, moist soil (Lim, 2013). It generally occurs in warm tropical areas with at least seasonally heavy precipitation, although in Venezuela, the species seems well-adapted to seasonally dry grassland climate (PBI Solanum Project, 2014). The elevation range for S. mammosum is usually from sea level to 100 m elevation, but can reach at least 1800 m (PBI Solanum Project, 2014). It grows at low altitudes in the Philippines and Fiji (Merrill, 1923; PIER, 2014) and in China within an altitude range of 200-1300 m (Flora of China Editorial Committee, 2014).

S. mammosum is found in lowland rainforests of Bolivia at altitudes of 0-500 m (Bolivia Checklist, 2014), and in Colombia, the species occurs in a variety of forest climates, from tropical dry forest to premontane humid forest, tropical humid and very humid forest, to humid submountain tropical forest. at altitudes of 0-1000 m (Vascular Plants of Antioquia, 2014). In Nicaragua it has been found to grow at altitudes up to 900 m (Flora of Nicaragua, 2014). In Peru, the species grows at altitudes up to 1000 m (Peru Checklist, 2014).

Climate

Soil Tolerances

Soil drainage

• free

Natural enemies

Means of Movement and Dispersal

S. mammosum has been intentionally introduced as an ornamental or curiosity for its uniquely shaped fruits to places including Papua New Guinea (Symon, 1986; PIER, 2014), Africa (Jaeger and Hepper, 1986) and China (An-ming, 1986). It can be expected to occur in cultivation anywhere in the Old and New tropics and has the potential to escape (PBI Solanum Project, 2014). According to Mabberly (2008) the dispersal vectors for S. mammosum are unknown, but as Chiarini and Barboza (2007) state, S. mammosum is a Solanum species bearing “big, indehiscent, spongy fruits with bulky seeds [which are] said to be dispersed by drain water after a rainstorm, the spongy tissue being an adaptation to flotation (Nee, 1979, 1991)”. Furthermore, the fruits are yellow, a colour attractive to birds that would then serve as dispersal agents (Chiarini and Barboza, 2009).

Pathway Causes

Pathway Vectors

Impact Summary

Economic Impact

S. mammosum is known to be toxic to cattle and other mammals (Torres et al., 2012), yet it continues to be used in traditional medicine and as an indigenous vegetable in Asia (AVRDC, 2003; Lim, 2013), despite the lack of pharmacological evidence (Wiart, 2006). The species does contain alkaloids which are both harmful to human health and useful in the pharmaceutical industry (Telek et al., 1977; Roddick, 1986; Hanelt et al., 2001; Lim, 2013).

S. mammosum has been reported as a secondary host of bacterial canker (Clavibacter michiganensis subsp. michiganensis), one of the most devastating bacterial pathogens to the tomato plant that occurs on all continents, destroying crops and plants through wilting, stunting, cankering and sometimes plant death (Sherf and Macnab, 1986).

Environmental Impact

S. mammosum has potential to have a negative impact on its local environment due to its prolific seed production and multiple possible biotic and abiotic vectors for seed dispersal (Nee, 1979; 1991; Chiarini and Barboza, 2007). It is known to be invasive in areas of both the Old and New tropics, and is listed as an agricultural weed in the Global Compendium of Weeds (Randall, 2012), but needs further assessment to determine the extent of its negative effect on the environment. 

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Abundant in its native range
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Pioneering in disturbed areas
• Benefits from human association (i.e. it is a human commensal)
• Has propagules that can remain viable for more than one year

• Negatively impacts agriculture
• Negatively impacts human health
• Negatively impacts animal health
• Reduced native biodiversity

• Competition
• Poisoning

• Highly likely to be transported internationally deliberately

Uses

Many Solanaceae species are known to be acutely toxic due to alkaloidal properties and have a long history of use as both human poisons and hallucinogens, especially in shaman and witch lore (PBI Solanum Project, 2014).

S. mammosum contains bioconstituents which are both highly toxic to humans and medicinally useful. It is a source of solasodine (Telek et al., 1977; Roddick, 1986; Hanelt et al., 2001; Lim, 2013), a poisonous, teratogenic, alkaloidal compound that is a precursor to pharmaceutical production of contraceptive pills and has also been the subject of recent research for its diuretic, anticancer, antifungal, cardiotonic, antispermatogenetic, antiandrogenic, immunomodulatory, and antipyretic effects on the central nervous system (Patel et al., 2013). The species has been used to treat athlete’s foot among hunter groups in Belize (Lim, 2013) and Trinidad (Lans et al., 2001), and its fruit is used to treat coughs and loss of appetite in the Philippines. An exhaustive list of medicinal uses is given by Lim (2012), including the following: “In the Dominican Republic, extracts of leaf and fruit are used for mouth infections, skin wounds and taken, to lose weight and to lower high blood pressure and high cholesterol; in Peninsular Malaysia, the plant is used to heal caterpillar rash and the Malays drink the leaf sap to treat fever; in Sarawak, fresh fruit juice is used to treat sore eyes in children; in Cambodia, Laos, and Vietnam, the plant is used to induce narcosis”.

Parts of S. mammosum are toxic and the species is reportedly used as poison for cockroaches, rats and insects (Hanelt et al., 2001; Mabberly, 2008; Flora of Nicaragua, 2014; Panama Checklist, 2014; PBI Solanum Project, 2014). Field experiments have demonstrated the potential use of the species as a natural insecticide for crops such as bananas (Segovia, 1998; Welman, 2003).

Despite its known toxicity, a recent report of the Asian Vegetable Research and Development Center listed S. mammosum as a palatable indigenous vegetable of high eating quality (AVRDC, 2003), and in the Philippines both the fruits and the leaves have reportedly been eaten as vegetables (Lim, 2013).

S. mammosum is cultivated around the world as an ornamental due to its peculiarly shaped fruits (Hanelt et al., 2001; Mabberly, 2008; Lim, 2013; Flora of Nicaragua, 2014; Panama Checklist, 2014; PBI Solanum Project, 2014). In Taiwan and Hong Kong, the decorative foliage and fruit are used in religious and festival floral arrangements, and the Chinese in Asia reportedly use the fruits during the Chinese Lunar New Year festive season, as the fruits’ golden colour is perceived to symbolize wealth (Lim, 2013).

Uses List

Drugs, stimulants, social uses

• Narcotic
• Religious

General

• Ornamental

Human food and beverage

• Vegetable

Materials

• Pesticide

Medicinal, pharmaceutical

• Source of medicine/pharmaceutical
• Traditional/folklore

Similarities to Other Species/Conditions

The following is taken from M. Nee’s description of the species on PBI Solanum Project (2014): S. mammosum can be distinguished from other species in section Acanthophora by the combination of its purple corollas, long copious pubescence, and its usually bizarre nipple-shaped fruit. It is most similar to S. palinacanthum, which also has purple corollas, but which has short glandular puberulence and exclusively globose fruits. 

Gaps in Knowledge/Research Needs

More research is needed on the current extent of this species’ invasiveness in both its native and introduced range. In areas where S. mammosum is known to be invasive, research on the damage to local ecosystems and human health would improve understanding of the species’ risk of introduction and invasiveness, especially in countries where it is more commonly used and cultivated. Additional research on the pharmacological evidence and safety of traditional use is also recommended.

References

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Contributors

24/08/2014 Original text by:

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

Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

Distribution Maps