Invasive Species Compendium

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Datasheet

Solanum viarum
(tropical soda apple)

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Datasheet

Solanum viarum (tropical soda apple)

Summary

  • Last modified
  • 24 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Host Plant
  • Preferred Scientific Name
  • Solanum viarum
  • Preferred Common Name
  • tropical soda apple
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • S. viarum is a fast-growing herb and a very aggressive invader. It produces thousands of small seeds (40,000 to 50,000 seeds per plant) that can be easily dispersed by birds, raccoons, cattle and by human activ...

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Pictures

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PictureTitleCaptionCopyright
Solanum viarum (tropical soda apple); infestation of well grown plants. USA
TitleInvasive habit
CaptionSolanum viarum (tropical soda apple); infestation of well grown plants. USA
Copyright©Charles T. Bryson/USDA Agricultural Research Service/Bugwood.org: CC BY 3.0 US.
Solanum viarum (tropical soda apple); infestation of well grown plants. USA
Invasive habitSolanum viarum (tropical soda apple); infestation of well grown plants. USA©Charles T. Bryson/USDA Agricultural Research Service/Bugwood.org: CC BY 3.0 US.
Typical habit, showing foliage
TitleHabit, showing foliage
CaptionTypical habit, showing foliage
Copyright©J. Jeffrey Mullahey/University of Florida/Bugwood.org: CC BY-NC 3.0 US
Typical habit, showing foliage
Habit, showing foliageTypical habit, showing foliage ©J. Jeffrey Mullahey/University of Florida/Bugwood.org: CC BY-NC 3.0 US
Solanum viarum (tropical soda apple); fruits. Thorny nightshade from Argentina, first appeared in the USA in pastures and rangelands in Glades County, Florida, in 1988. Mottled green fruits that look like small watermelons are a distinguishing feature .
TitleFruits
CaptionSolanum viarum (tropical soda apple); fruits. Thorny nightshade from Argentina, first appeared in the USA in pastures and rangelands in Glades County, Florida, in 1988. Mottled green fruits that look like small watermelons are a distinguishing feature .
Copyright©J. Jeffrey Mullahey, University of Florida, Bugwood.org: CC BY-NC 3.0 US
Solanum viarum (tropical soda apple); fruits. Thorny nightshade from Argentina, first appeared in the USA in pastures and rangelands in Glades County, Florida, in 1988. Mottled green fruits that look like small watermelons are a distinguishing feature .
FruitsSolanum viarum (tropical soda apple); fruits. Thorny nightshade from Argentina, first appeared in the USA in pastures and rangelands in Glades County, Florida, in 1988. Mottled green fruits that look like small watermelons are a distinguishing feature .©J. Jeffrey Mullahey, University of Florida, Bugwood.org: CC BY-NC 3.0 US
Solanum viarum (tropical soda apple); fruiting plant in November. USA
TitleMaturing fruits
CaptionSolanum viarum (tropical soda apple); fruiting plant in November. USA
Copyright©John W. Everest/Auburn University/Bugwood.org. CC BY 3.0 US.
Solanum viarum (tropical soda apple); fruiting plant in November. USA
Maturing fruitsSolanum viarum (tropical soda apple); fruiting plant in November. USA©John W. Everest/Auburn University/Bugwood.org. CC BY 3.0 US.
Frank Krainin, PPQ, studies a Solanum viarum plant after frost. An early observation that TSA roots could survive the winter in Georgia, USA.
TitleInvasive habit
CaptionFrank Krainin, PPQ, studies a Solanum viarum plant after frost. An early observation that TSA roots could survive the winter in Georgia, USA.
Copyright©Arthur E. Miller, USDA APHIS PPQ, Bugwood.org. CC BY 3.0 US.
Frank Krainin, PPQ, studies a Solanum viarum plant after frost. An early observation that TSA roots could survive the winter in Georgia, USA.
Invasive habitFrank Krainin, PPQ, studies a Solanum viarum plant after frost. An early observation that TSA roots could survive the winter in Georgia, USA.©Arthur E. Miller, USDA APHIS PPQ, Bugwood.org. CC BY 3.0 US.
Young plant of Solanum viarum. Bhutan.
TitleHabit
CaptionYoung plant of Solanum viarum. Bhutan.
Copyright©Chris Parker/Bristol, UK
Young plant of Solanum viarum. Bhutan.
HabitYoung plant of Solanum viarum. Bhutan.©Chris Parker/Bristol, UK

Identity

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Preferred Scientific Name

  • Solanum viarum Dunal (1852)

Preferred Common Name

  • tropical soda apple

Other Scientific Names

  • Solanum chloranthum DC.
  • Solanum khasianum var. chatterjeeanum Sengupta & Sengupta
  • Solanum reflexum Schrank (1819)
  • Solanum viridiflorum Schltdl.

International Common Names

  • English: sodom apple
  • Spanish: calabacilla; manzana de soda; manzana tropical

Local Common Names

  • Argentina: tutía de vibora
  • Bhutan: kachera kanra
  • Brazil: arrebenta-cavalo; joa-bravo; joão-bravo; juá; juá-bravo
  • China: mao guo qie
  • India: kantha kari
  • Puerto Rico: berenjena

EPPO code

  • SOLVI (Solanum viarum)

Summary of Invasiveness

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S. viarum is a fast-growing herb and a very aggressive invader. It produces thousands of small seeds (40,000 to 50,000 seeds per plant) that can be easily dispersed by birds, raccoons, cattle and by human activity (seed-contaminated grass, manure, mud, and farming machinery; Medal et al., 2012). Once established in new areas, S. viraum is able to grow forming monocultures that occur in patches of 4 ha or more (Waggy, 2009). S. viarum is classified as a weed by the Global Compendium of Weeds (Randall, 2012) and it also was declared a noxious weed by the US Department of Agriculture (USDA-Aphis) for the US territories of Alabama, Arizona, California, Florida, Georgia, Massachusetts, Minnesota, Mississippi, North and South Carolina, Oregon, Tennessee, Texas, Vermont and Puerto Rico (USDA-NRCS, 2012).

S. viarum thrives in overgrazed or drought-affected pastures, and invades plantation crops and natural habitats including forests and river banks. Although animals do not eat the foliage, the fruits are readily eaten and distributed by cattle and other mammals.

 

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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This species has previously been known as S. khasianum, and there is some continuing confusion over the status of  the two subspecies S. khasianum subsp. chatterjeeanum an accepted synonym for S. viarum, and S. khasianum subsp. khasianum, also known as S. aculeatissimum Jacquin). The latter differs in having glabrous rather than pubescent prickles. The name Solanum reflexum Schrank according to GBIF (2012) and Catalogue of Life (2012), equates with S. aculatissimum. According to The Plant List (2012) it is a synonym for the ‘accepted’ name S. viarum, while Missouri Botanical Garden (2012) indicates that S. reflexum is the ‘accepted’ name for S. viarum. For purposes of this datasheet, the name S. viarum is retained. Some relevant data for ‘S. reflexum’ and S. aculatissimum may have been overlooked.

Description

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S. viarum is an erect perennial, 50-150 cm high, with shortly pubescent stems and branches with recurved prickles up to 5 mm long, pubescent at their base. There are also longer, straight spines up to 2 cm long on the petioles and the veins of upper and lower surfaces of the leaves. The leaves are broadly ovate up to 20 cm long and 15 cm wide, bluntly lobed with markedly undulate edges, generally dark green, glossy above, duller below. The flowers are white, 1.5 cm across in clusters of 1-5 on pedicles about 1 cm long, the more distal flowers are often male only. Sepals about 3-5 mm long, corolla of white, somewhat narrow, reflexed petals, anthers pale yellow. The fruit is a globose berry, mottled green when young, maturing yellow, 2-3 cm across, containing up to 400 brown, flattened, discoid seeds, 2-3 mm in diameter (from Mill, 2001; Weber, 2003). Roots have buds which will regenerate new shoots. The root system can be extensive, with feeder roots 1-2 cm in diameter located a few cm below ground extending 1-2 m from the crown of the plant (Ferrell and Mullahey, 2006).

Plant Type

Top of page Broadleaved
Herbaceous
Perennial
Seed propagated
Shrub
Vegetatively propagated

Distribution

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S. viarum is native to Argentina, southern Brazil, Paraguay, and Uruguay (Nee, 1999). It has been introduced into the southeastern United States, Mexico, Honduras, Puerto Rico, India, Nepal, South Africa and some other parts of Asia (Mullahey et al., 1993; 1996), where it has become a major concern to agriculture and cattle production (Bryson et al., 2009; Medal et al,. 2012). It is recorded as occurring ‘throughout India’ (GISIN, 2008) or ‘widely distributed from the Himalayan foot-hills in the North to the Nilgiris in the South of India’ (Singh et al., 1998), though it is confirmed in only a few of the states. In Bhutan, it occurs mainly below 2000m (Parker, 1993). Considering invasiveness-related traits of this species, it is expected to spread to other tropical and subtropical areas outside its current range (Nee, 1991). 

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

Africa

Central African RepublicPresentIntroduced1985USA, Missouri Botanical Garden (2008)
EswatiniPresentIntroducedISSG (2012)
South AfricaPresentIntroduced2000InvasiveWelman (2003); Kandari et al. (2011)Weed

Asia

BhutanPresent, LocalizedIntroducedMill (2001)
ChinaPresent, LocalizedIntroducedUSA, Missouri Botanical Garden (2008)
-TibetPresentIntroducedUSA, Missouri Botanical Garden (2008); Flora of China (1994)
-YunnanPresentIntroducedUSA, Missouri Botanical Garden (2008); Flora of China (1994)
IndiaPresentGISIN (2008); Singh et al. (2012); USDA-ARS (2012)'throughout India'
-Arunachal PradeshPresentIntroducedInvasiveChandra (2012)
-AssamPresentIntroducedDeka and Deka (2007)
-Himachal PradeshPresentIntroducedInvasiveChandra (2012)
-Jammu and KashmirPresentIntroducedInvasiveChandra (2012)
-KarnatakaPresentIntroducedDasar et al. (2005)
-KeralaPresentMuraleedharan et al. (1999)
-Madhya PradeshPresentIntroducedVerma (1993)
-MaharashtraPresentIntroducedSuryawansi et al. (2001)
-ManipurPresentIntroducedInvasiveChandra (2012)
-MeghalayaPresentIntroducedInvasiveChandra (2012)
-NagalandPresentIntroducedInvasiveChandra (2012)
-SikkimPresentIntroducedInvasiveChandra (2012)
-TripuraPresentIntroducedInvasiveChandra (2012)
-Uttar PradeshPresentIntroducedInvasiveChandra (2012)
-UttarakhandPresentIntroducedInvasiveChandra (2012)
-West BengalPresentIntroducedMill (2001)
MyanmarPresentIntroducedInvasiveMill (2001); USDA-ARS (2012)
NepalPresentIntroducedInvasiveMill (2001); USDA-ARS (2012)
TaiwanPresentIntroducedTalekar et al. (1999)
ThailandPresentIntroducedGBIF (2008)
VietnamPresentIntroducedInvasiveUSDA-ARS (2012)

Europe

BelgiumPresent, Few occurrencesIntroducedVerloove (2006); DAISIE (2012)
SpainPresentIntroducedInvasiveMAGRAMA (2012)

North America

Costa RicaPresentGBIF (2008)
HondurasPresentIntroducedInvasiveBryson and Byrd (2007); USDA-ARS (2012)
MexicoPresentIntroducedInvasiveUSDA-ARS (2012)
Puerto RicoPresentIntroducedInvasiveKairo et al. (2003); Bryson and Byrd (2007); Axelrod (2011); Más and Lugo (2013)
United StatesPresent, LocalizedIntroduced1988InvasiveUSDA-NRCS (2008)
-AlabamaPresent, LocalizedIntroducedInvasiveUSDA-NRCS (2008); USDA-NRCS (2012)
-CaliforniaPresent, LocalizedIntroducedInvasiveEDDMapS (2013)Has quarantine status
-FloridaPresent, WidespreadIntroduced1987InvasiveUSDA-NRCS (2008); Florida Exotic Pest Plant Council (2011)
-GeorgiaPresent, LocalizedIntroducedInvasiveUSDA-NRCS (2008)
-LouisianaPresent, LocalizedIntroducedInvasiveUSDA-NRCS (2008)
-MississippiPresent, LocalizedIntroduced1994InvasiveUSDA-NRCS (2008); USDA-ARS (2012)
-North CarolinaPresent, LocalizedIntroducedInvasiveUSDA-NRCS (2008); USDA-ARS (2012)
-OklahomaPresent, LocalizedIntroducedInvasiveEDDMapS (2013)
-PennsylvaniaPresent, LocalizedIntroduced1996USDA-NRCS (2008)
-South CarolinaPresent, LocalizedIntroducedInvasiveUSDA-NRCS (2008); USDA-NRCS (2012)
-TennesseePresent, LocalizedIntroducedInvasiveUSDA-NRCS (2008); USDA-NRCS (2012)
-TexasPresent, LocalizedIntroducedMedal et al. (2007); USDA-NRCS (2012)

Oceania

AustraliaPresentCABI (Undated)Present based on regional distribution.
-New South WalesPresentIntroducedInvasiveUSDA-ARS (2012)
-QueenslandPresent, Few occurrencesIntroducedAustralian Weeds Committee (2012)

South America

ArgentinaPresentNativeGRIN (2008); Nee (1999)
BrazilPresentNativeGRIN (2008)
-AcrePresentNativeForzza RC et al. (2012)
-AmazonasPresentNativeGRIN (2008)
-BahiaPresentNativeForzza RC et al. (2012)
-Espirito SantoPresentLorenzi (1982); Forzza RC et al. (2012)
-GoiasPresentNativeGRIN (2008); Forzza RC et al. (2012)
-Mato GrossoPresentNativeForzza RC et al. (2012)
-Mato Grosso do SulPresentLorenzi (1982); Forzza RC et al. (2012)
-Minas GeraisPresentNativeUSDA-ARS (2008); Forzza RC et al. (2012)
-ParanaPresentNativeUSDA-ARS (2008); Forzza RC et al. (2012)
-Rio de JaneiroPresentLorenzi (1982); Forzza RC et al. (2012)
-Rio Grande do SulPresentNativeUSDA-ARS (2008); Forzza RC et al. (2012)
-RondoniaPresentNativeForzza RC et al. (2012)
-Santa CatarinaPresentNativeUSDA-ARS (2008); Forzza RC et al. (2012)
-Sao PauloPresentNativeUSDA-ARS (2008); Forzza RC et al. (2012)
ParaguayPresentNativeUSDA-ARS (2008); Nee (1999)
UruguayPresentNativeUSDA-ARS (2008); Nee (1999)

History of Introduction and Spread

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Originating in South America, S. viarum has found its way into northern America and into Africa and Asia. It has apparently been in India for a long time, as it has been widely cultivated and specially bred for medicinal purposes. In the Americas it has spread from its native range into other parts of South and Central America (including Mexico and Honduras) probably by seeds adhering to peoples shoes, mud, and contaminated grass seeds (Medal et al., 2008).

In Florida, S. viarum was first collected in 1988. The species was most likely introduced accidentally to Florida in cattle carrying undigested seeds that were imported from Brazil (ca. 1985) and from here it has spread to other southeastern and mid-Atlantic states (Medal et al., 2012). It is now also found in a small area of California (EDDMapS, 2013), and is on quarantine or noxious weed lists for a number of other states in the USA other than in its recorded range (USDA-NRCS, 2012). It has become a major concern of agriculture and a serious threat to natural plant communities (Cuda et al., 2002; Medal et al., 2008; ISSG, 2012) with over 400,000 ha of pasture land infested by this species in Florida alone (Medal et al., 2012).

S. viarum was first detected in Puerto Rico around 2006 in western areas (at Mayagüez). More recently, the species has been collected in various localities, including Puerto Rico’s east coast, suggesting a rapid spread throughout the island (Más and Lugo, 2013). In Australia, S. viarum was first recorded in the Kempsey area on the coast of New South Wales in August 2010; however it is thought to have been present there for some time. More recently it has been found in Southern Queensland and it is likely that this species is already present in other parts of eastern Australia (Australian Weeds Committee, 2012). In South Africa it was first reported in the early 2000s. 

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Florida 1987 Yes No Bryson and Byrd (2007)
Mississippi 1994 No No Bryson and Byrd (2007)
Pennsylvania 1996 Yes No
South Africa 2000 Yes No Welman (2003) First reported 2003

Risk of Introduction

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According to different risk assessments performed on this species (Lehtonen, 1994; Gordon et al., 2008) the risk of introduction of S. viarum is high. This species is an aggressive invasive and each plant is able to produce thousands of seeds that can be dispersed by birds, cattle, and by human activities. It is also able to regenerate from its root system (Medal et al., 2008, 2012).

The risk of deliberate introduction is relatively high, given the popularity of the species as a medicinal herb and its importance as a source of pharmaceutical compounds, especially solasodine.

S. viarum is listed on the USA Federal Noxious Weed List, and hence its possession, movement and release is prohibited in the USA (USDA-NRCS, 2008). It is not yet recorded in Pacific or Indian Ocean islands (PIER, 2008), and it may appear imperative that introduction to these or any other new areas is prevented, noting its ability to spread very rapidly as has recently occurred in the south-eastern USA. 

Habitat

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Where native, S. viarum can be found growing in grassland, thickets, and disturbed places such as roadsides and river banks. Outside its native range, this species grows as a common weed in agricultural fields, pastures, and natural areas. In the southeastern USA it has been observed in pastures, ditch banks, citrus plantations, sugarcane fields, vegetable fields, and native areas like oak hammocks, pine forests, riparian habitat, and native grasslands (Medal et al., 2012). S. viarum thrives in disturbed areas associated with human activities.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
 
Terrestrial – ManagedCultivated / agricultural land Secondary/tolerated habitat Harmful (pest or invasive)
Cultivated / agricultural land Secondary/tolerated habitat Productive/non-natural
Managed forests, plantations and orchards Secondary/tolerated habitat Harmful (pest or invasive)
Managed grasslands (grazing systems) Principal habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Natural
Rail / roadsides Principal habitat Harmful (pest or invasive)
Rail / roadsides Principal habitat Natural
Urban / peri-urban areas Principal habitat Harmful (pest or invasive)
Urban / peri-urban areas Principal habitat Natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Secondary/tolerated habitat Harmful (pest or invasive)
Natural grasslands Principal habitat Harmful (pest or invasive)
Natural grasslands Principal habitat Natural
Riverbanks Principal habitat Harmful (pest or invasive)
Riverbanks Principal habitat Natural
Wetlands Secondary/tolerated habitat Harmful (pest or invasive)
Wetlands Secondary/tolerated habitat Natural
Scrub / shrublands Secondary/tolerated habitat Harmful (pest or invasive)
Scrub / shrublands Secondary/tolerated habitat Natural

Hosts/Species Affected

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S. viarum has become a major concern of agriculture and cattle farming. It invades improved pastures where it reduces livestock carrying capacity and it is also a reservoir for at least six crop viruses (potato leaf-roll virus, potato virus Y, tomato mosaic virus, tomato mottle virus, tobacco etch virus, and cucumber mosaic virus) and the potato fungus Alternaria solani (McGovern et al., 1994a, 1994b, 1996; Medal et al., 2012). Several insect pests also utilize this species as an alternate host.

Host Plants and Other Plants Affected

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Plant nameFamilyContext
CitrusRutaceaeOther
Paspalum notatum (Bahia grass)PoaceaeMain
Saccharum officinarum (sugarcane)PoaceaeOther

Biology and Ecology

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Genetics

The normal chromosome number is 2n=24 (Kumaraswamy and Krishnan, 1987; Chiarini, 2006; Missouri Botanical Garden, 2008) but autotetraploids have been deliberately created for cultivation in India (Srinivasappa et al., 1999). The variety Arka Sanjeevini is diploid whereas the variety Arka Mahima is tetraploid with a lower overall yield but higher content of solasodine (Paturde et al., 2002). 

Reproductive Biology

All Solanum species have poricidally dehiscent anthers that make this genus an example of the buzz pollination syndrome, found in about 200 plant genera (Buchmann, 1983). Solanum flowers are mainly hermaphroditic, nectar is absent and pollen is the exclusive floral reward. Pollination in these flowers is performed by insects (mainly bees). In Florida, the orchid bee Euglossa viridissima was observed collecting pollen from S. viarum and may serve as a pollinator (Pemberton and Wheeler, 2006).

The plant can recover from the rootstock after the shoots are killed by frost, and there is some degree of vegetative reproduction with shoots developing from spreading roots. Seed production is high, and seeds survive in the soil for up to one year (Mullahey et al., 1998). 

Physiology and Phenology

Flowering and fruit production occur throughout the year in S. viarum, but in Florida, reproductive activity is concentrated from September through May (Lehtonen, 1994). Throughout the year, plant of S. viarum may have both immature and mature fruit present, ensuring production of large numbers of viable seeds (approximately 40,000 or more per plant; Akanda et al., 1996). Seedling emergence primarily occurs from August through March. New plants can emerge from seed or from roots. The species has an extensive root system which may extend 1-2 metres horizontally from the crown of the plant (Medal et al., 2012).

Germination of S. viarum is moderately photoblastic with 30% germination occurring in the dark. Germination increases in response to green (545 nm) and red light (650 nm) to 75 and 66%, respectively, indicating phytochrome regulation. No germination occurs in response to blue (450 nm) or far-red light (750 nm). Germination increases from 4 to 64% between 10°C and 30°C, but no germination is found at 5°C and 40°C. Maximum germination occurs at 30°C. Seedling emergence is maximum from a depth of 3-6 cm, but no seedling emergence occurs when seeds are deeper than 12 cm. Optimum germination occurs at a depth of 5.6 cm. Mechanical and sulfuric acid scarification increases the rate of germination but not the overall percentage. Tap water or hot water pretreatments increase the rate of germination by 26%, and KNO3, GA3 or ethephon increase germination by 53%. Suryawanshi et al. (2001) recommend treatment with 12.5% nitric acid for 15 minutes followed by 1000 ppm gibberellic acid for 24 h with alternate temperatures of 20-30°C. In conclusion, seed germinates in response to variable environmental and edaphic conditions which could allow its establishment in diverse ecosystems (Akanda et al., 1996).        

Longevity

In favourable temperatures, plants flower within 60-100 days after emergence in photoperiods of 8-16 h, with flowering delayed by photoperiods less than 10 h or temperatures lower than 24/20°C (Patterson et al., 1997). Green stems persist in mild winter temperatures (Coile, 1993). Plants become less productive or may die in summer when standing in water (Mullahey and Colvin, 1993) or in winter when leaves may be severely damaged by frost (Lehtonen, 1994).

Although normally perennial, S. viarum can produce fruit in the first season of growth and thus behave as an annual. 

Associations

Within its native range, S. viarum grows in the understory of rainforest, in grasslands and in shrubby-thicket dry forests (Nee, 1991). In invaded areas, S. viarum often occurs in cultivated pastures, primarily pastures planted with bahia grass (Paspalum notatum), citrus plantations, sugarcane plantations, and vegetable fields (Medal et al., 2008; 2012; ISSG, 2012). This species is also common as a weed in sites associated with anthropogenic disturbance (waste grounds and along roadsides) and in native plant communities (Medal et al., 2012). 

Environmental Requirements

Growth and development is enhanced under sunny conditions with temperatures ranging from 20°C to 35°C and average annual rainfall from 700 mm to 2000 mm. Plants survived in 8°C nights with day temperatures of 18-36°C, but biomass and leaf area were only 3-10% of the maximum (Patterson et al., 1997).

This species does not tolerate water-logging or frosty conditions for extended periods. S. viarum prefers to grow in well drained sandy loam soils with high organic matter content (Lehtonen, 1994).

The rapid spread of S. viarum is apparently favoured by moderate drought conditions in Florida, USA, though this may be partly due to the overgrazing which occurs under those conditions (Hogue et al., 2006).

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])
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
30 40

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 15 25
Mean maximum temperature of hottest month (ºC) 34
Mean minimum temperature of coldest month (ºC) 10

Rainfall

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ParameterLower limitUpper limitDescription
Dry season duration03number of consecutive months with <40 mm rainfall
Mean annual rainfall10002000mm; lower/upper limits

Rainfall Regime

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Summer
Uniform
Winter

Soil Tolerances

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Soil drainage

  • free
  • seasonally waterlogged

Soil reaction

  • acid
  • alkaline
  • neutral

Soil texture

  • light
  • medium

Special soil tolerances

  • shallow

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Alternaria solani Pathogen Stems not specific
Amblyophallus maculatus Herbivore Leaves to genus
Anthonomus tenebrosus Herbivore Leaves to genus
Bemisia tabaci Herbivore Stems not specific
Bemisia tabaci (MEAM1) Herbivore Stems not specific
Corythaica cyathicollis Herbivore Leaves to genus
Cucumber mosaic virus Pathogen Stems not specific
Diabrotica speciosa Herbivore Leaves not specific
Gratiana boliviana Herbivore Leaves to species S. America, USA
Gratiana graminea Herbivore Leaves to genus
Helicoverpa armigera Herbivore Stems not specific
Mechanitis lysimnia Herbivore Leaves not specific
Meloidogyne arenaria Parasite Roots not specific
Meloidogyne javanica Parasite Roots not specific
Metriona elatior Herbivore Leaves
Neoleucinodes elegantalis Herbivore Fruits/pods not specific
Platyphora Herbivore Leaves not specific
Potato leafroll virus Pathogen Stems not specific
Potato virus Y Pathogen Stems not specific
Tobacco etch virus Pathogen Stems not specific
Tobacco mild green mosaic virus Pathogen not specific Charudattan and Hiebert, 2007 USA
Tomato mosaic virus Pathogen Stems not specific
Tomato mottle virus Pathogen Stems not specific
tropical soda apple mosaic virus Pathogen Stems not specific

Notes on Natural Enemies

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S. viarum has been shown to be an alternate host of a wide range of viruses, nematodes, pathogens and insects, most quite unspecific, but surveys in South America revealed some insects with potential as biocontrol agents, especially the leaf-eating coleopteran Chrysomelidae Metriona elatior, Gratiana boliviana, G. graminea and a Platyphora sp.; also a flower bud weevil, Anthonomus tenebrosus from Argentina and Brazil (Medal et al., 2000; 2002) and a leaf-root feeder Epitrix parvula (Medal et al., 2012). G. boliviana has been used for biological control in Florida (Medal et al., 2010 and references therein). Two bacteria, Ralstonia solanacearum and Erwinia carotovora subsp. carotovora have also been shown to damage S. viarum and are of interest for inundative biocontrol in the USA. Tobacco mild green mosaic virus, which has no known vector, elicits a severe hypersensitive response in S. viarum and has been developed as a bioherbicide (Charudattan and Hiebert, 2007).

Means of Movement and Dispersal

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S. viarum can be dispersed by seeds and by shoots from its extensive root system. This species is able to produce from 40,000 to 50,000 seeds per plant (Mullahey et al. 1993). Seeds can be dispersed by birds and other animals, including cattle, deer, feral pigs, and raccoons (Akanda et al. 1996; Waggy 2009; ISSG, 2012); also the tegu lizard (Tupinambis merianae) (Castro and Galetti, 2004). Animals do not eat the foliage but consume the fruits and spread the seeds in their faeces (Mullahey et al., 1998; Bryson and Byrd, 2007).

Sale of cattle that have recently been feeding on the fruits of S. viarum over long distances has resulted in spread of the plant from state to state in the USA, and the movement of hay and manure have also been implicated in accidental introductions (Mullahey et al., 2006). Other mechanisms of movement reported include transfer in turf, water, and grass seeds from contaminated pastures (Bryson and Byrd, 2007).

S. viarum is planted as a commercial medicinal crop especially in India, and thus intentional introduction is likely to have occurred at least in South Asia.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Animal productionMovement of cattle, manure within USA Yes Yes Mullahey et al., 2006
Crop productionMovement of cattle, manure, hay within USA Yes Yes Mullahey et al., 2006
Digestion and excretion Yes Mullahey et al., 2006
Foragemovement of hay, within USA Yes Yes Mullahey et al., 2006

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Land vehiclesSeeds Yes Yes Medal et al., 2012
Livestockseeds Yes Yes Mullahey et al., 2006
Soil, sand and gravelSeeds Yes Yes Medal et al., 2012

Impact Summary

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CategoryImpact
Economic/livelihood Negative
Environment (generally) Negative
Human health Positive

Economic Impact

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In Florida, USA, S. viarum occurs in pastures, citrus plantations, sugarcane and vegetable fields, turf fields and roadsides (Mullahey, 1996). It has spread extremely rapidly, especially in bahiagrass (Paspalum notatum) pastures, and especially in years of drought when its effect on available grazing is particularly serious. Foliage and stems are unpalatable to cattle (Medal et al., 2012). In addition, S. viarum grows forming dense monospecific stands that prevent cattle access to shaded areas and resulting in summer heat stress (Mullahey et al., 1998). For Florida ranchers, the control costs of S. viarum were estimated at $6.5 to 16 million annually (Thomas, 2007), and economic losses from cattle heat stress alone have been estimated at $2 million (Mullahey et al., 1998). It has also caused poisoning of goats in Florida (Porter et al., 2003).

It may act as an alternate host of a range of crop pathogens, including Cucumber mosaic virus, Potato leaf roll virus, Potato Y virus, Tomato mosaic virus and the fungus Alternaria solani (Cooke, 1997), though there are no reports of direct economic loss from this. It is also an alternate host of Meloidogyne javanica, providing a potential reservoir of this nematode in the USA (Inserra, 1994); also of Bemisia tabaci in Brazil (Lourenção and Nagai, 1994). Other insect pests detected in S. viarum (Sudbrink et al., 2000; Medal et al., 2012) include:

  • potato beetle Leptinotarsa decemlineata
  • tomato hornworm Manduca quinquemaculata
  • tobacco hornworm Manduca sexta
  • tobacco budworm Helicoverpa virescens
  • tomato pinworm Keiferia lycopersicella
  • green peach aphid Myzuz persicae
  • soybean looper Pseudoplusia includens
  • southern green stink bug Nezara viridula

In Florida, management practices principally involve herbicide applications and mowing in invaded areas which provide temporary weed suppression at an estimated cost of US $61 and $47 per ha, respectively (Thomas, 2007). Based on survey of Florida cattle producers in 2006, S. viarum control costs resulted in economic losses throughout the state of $15 million annually (Salaudeen et al., 2013).

Environmental Impact

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S. viarum has been identified as a noxious weed that can smother native plant communities where it has been introduced. It out-competes native plant species by crowding or shading them out (Waggy, 2009; Medal et al., 2012). This species also reduces biodiversity in natural forests because plants are able to dominate large areas in the understory affecting the germination and establishment of native species. In the southeastern USA it infests natural areas including state parks, nature preserves and hammocks (raised woodland above swamp land) (Mullahey, 1996). Plant prickles can also restrict wildlife grazing and create a physical barrier to animals, preventing movement through infested areas (USDA-FS, 2005). 

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • 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
  • Tolerant of shade
  • Highly mobile locally
  • Fast growing
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Has high genetic variability
Impact outcomes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Monoculture formation
  • Negatively impacts agriculture
  • Negatively impacts forestry
  • Negatively impacts livelihoods
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - shading
  • Competition - smothering
  • Competition (unspecified)
  • Pest and disease transmission
  • Poisoning
  • Rapid growth
  • Rooting
  • Produces spines, thorns or burrs
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally deliberately
  • Difficult/costly to control

Uses

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In India, S. viarum is being cultivated in many states, and collected from the wild in others, for a variety of traditional and modern uses.

S. viarum yields solasodine, a nitrogen analogue of diosgenin, whose hormonal derivatives are used in corticosteroid industry as an active ingredient of the contraceptive pill (Dasar etal., 2005; Maiti et al., 2000; Bhaskar et al., 2000; 2002). Varieties with very high yield potential for alkaloids have been developed (e.g. Glaxo, BARC, IIHR-2N, Arka-sanjivani, Arka Mahima and NBRI sel.) (Singh et al., 1998). Among these, Arka Mahima is one of the autotetrapoids which have been created, having higher levels of solasodine. Comparative histochemical studies of exotestas in diploid (2n), autotetraploid (4n) and tertiary trisomic (2n + 1) suggest that increased cell size in this layer in the autotetraploid plants probably accounts for the higher steroid content reported (Srinivas et al., 1998). However, inheritance of content is not correlated with yield. High alkaloid content is associated with low berry yield while high berry yield was correlated with medium levels of alkaloids. The weight of berries is dependent upon the number of seeds, the larger the number of seed, the higher the berry weight. Since glyco-alkaloids are located in the gelatinous layer around the seeds, increase in seed number per berry increases the alkaloid content.

The agronomy of S. viarum as a crop, grown in fallow land after rice, has been studied, especially in Karnataka, India. Solasodine content in the fruits is increased with increasing N rates as well as the fruit yield (Dasar et al., 2005). The best intercrop was found to be cluster bean (Bhaskar et al., 2000).

Stravato and Cappelli (2000) found individuals of S. viarum resistant to Fusarium oxysporum f.sp. melongenae, the causal agent of Fusarium wilt of aubergine or eggplant, and suggest the possibility of using these as a source for transferring disease resistance. Similarly, Matsui et al. (1995) found resistance to Thrips palmi in S. viarum.

Another potential use for S. viarum has been explored by Srinavasan et al.(2005), who showed that the plant contains potent attractants for Helicoverpa armigera.

S. viarum is also valued as a medicinal herb. In India, the fruit is used to treat asthma, bronchitis, coughs, colds, wounds, toothache, tooth decay, and as an abortifacient. The seeds are used as a contraceptive and for menstrual complaints (Mill, 2001). It is also given to dogs infected with Dirofilaria immitis (Chakraborty et al., 1994).

Uses List

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Genetic importance

  • Gene source

Materials

  • Chemicals
  • Pesticide

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore
  • Veterinary

Similarities to Other Species/Conditions

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S. viarum can be distinguished from other Solanum species by its straight prickles, mixture of stellate and simple hairs with and without glands, clearly petioled leaves with a velvety sheen, terminal flower clusters, and yellow berries that are dark-veined when young (Florida Exotic Pest Plant Council, 2011). This species looks very similar to Solanum capsicoides and both have white flowers, but in S. viarum fruits are yellow when ripe while in S. capsicoides fruits are bright orange (Australian Weeds Committee, 2012).

Among other close relatives of S. viarum,S. anguivi differs in having mauve flowers and tomentose underside to the leaves, while S. aculeatissimum (= S. khasianum var. khasianum) and S. torvum both have white flowers but strictly glabrous prickles on the stems. In the USA, S. carolinense differs in being smaller, and having smaller fruits and leaves but larger flowers up to 3 cm across and a deeper root system, while S. tampicense has much narrower, more deeply lobed leaves.

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.

Prevention

S. viarum is listed on the USA Federal Noxious Weed List, and hence its possession, movement and release is prohibited in the USA (USDA-NRCS, 2008). 

To avoid the risk of introducing the weed when purchasing cattle from an infested area, the livestock should be kept in a restricted area for at least 6 days by which time any ingested seeds should have been voided (Mullahey et al., 2006). Correspondingly, those selling cattle should ensure that they are not fed on contaminated pastures prior to sale, and that any hay sold off the farm is free of contamination. 

Control

S. viarum is difficult to eradicate and the best management strategy varies according to the population size. Individual plants and small populations should be pulled up and burned completely along with all fruit. Roots have to be removed entirely because this species can regenerate shoots and re-grow from the root fragments (Medal et al., 2012). Larger populations require repeated mowing and/or repeated applications of herbicides or bioherbicides. It is important that plants are not allowed to fruit in order to prevent seed dispersal (Waggy, 2009); mowing below 10 cm plant height every 60 days will prevent fruit production and result in some plant mortality (Mullahey et al., 2006). Gloves are needed when handling the cut plants. 

Biological control

A petition for release of Gratiana boliviana in the USA was approved in 2002 (Medal et al., 2007; USDA/TAG, 2008). The beetle was introduced from Argentina and Paraguay: releases began in Florida in 2003 and more than 100,000 beetles have been released in Florida, Georgia, Alabama, South Carolina and Texas. Establishment has been good, with spread of 1-10 miles (1.5-16.0 km) per year from the release sites, with 20-100% defoliation and no non-target damage has been observed (Medal et al., 2006; 2007; Medal, 2008). Studies have shown that G. boliviana is better suited for control of small infestations of S. viarum than large or remote infestations (Waggy, 2009). 

Field tests in Brazil and Argentina confirmed the specificity of Metriona elatior to S. viarum and lack of attack on related solanaceous crops (Bredow et al., 2007; Gandolfo et al., 2007), and “these data suggest that a host range expansion of M. elatior to include eggplant, potato, tomato, or bell-pepper is highly unlikely” (Bredowet al., 2007). However, other laboratory tests had shown some feeding on eggplant (Medal et al., 2002) and the release of M. elatior in the USA was rejected in 2008 (USDA/TAG, 2008). Release of Gratiana graminea was also refused at this time. A petition to release Anthonomus tenebrosus was submitted in 2007 (USDA/TAG, 2008), and work with the flower-bud weevil, Platyphora sp. is apparently still in progress. 

The bacterial pathogen Ralstonia solanacearum is effective in causing plants to wilt and die (USDA-FS, 2005) and has been applied with a wet-blade mower to control S. viarum in field trials (DeValerio et al., 2011). Another bacterium, Erwinia carotovora subsp. carotovora, has been used in conjunction with reduced doses of herbicide to increase the level of control (Roberts et al., 2002). 

A bioherbicide containing Tobacco mild green mosaic virus strain U2 has been recently registered for use against S. viarum in the USA (Charudattan and Hiebert, 2007; Charudattan, 2015).

Chemical control

S. viarum can be effectively controlled using any of several herbicides such as glyphosate, imazapyr or triclopyr, though it is suggested that fruits are also collected and destroyed to prevent reestablishment (USDA-FS, 2005). Other recommended herbicides include aminopyralid (Akanda et al., 1997; Ferrell et al., 2006), hexazinone (Mislevy and Martin, 1999), glufosinate-ammonium, picloram, clopyralid, fluroxypyr and dicamba (Dowler, 1995). Effectiveness of triclopyr and hexazinone was enhanced when they were applied 60 days after frost had damaged the foliage (Mislevy and Martin, 1999). Effectiveness of triclopyr was similarly enhanced when applied after one or two prior mowings (Miselvy et al., 1999). However, aminopyralid can be applied at any time of year and will control existing plants and germinating seedlings for over 6 months after application (Hogue et al., 2006).

Treatments containing picloram or triclopyr controlled eight-leaf, 16-leaf, and 1-yr-old S. viarum greater than 90%, 8 weeks after treatment (Call et al., 2000). Acifluorfen, clopyralid, dicamba, fluroxypyr, picloram, triclopyr, glyphosate and imazapyr all resulted in >90% weed control after 145 days, though the latter two also caused >90% damage to Paspalum notatum (Akanda et al., 1997).

Treatment of an entire pasture should consist of mowing adult plants in April-May, allowing 50-60 days for regrowth, followed by single or repeated doses of triclopyr. Follow-up treatments will be necessary to control escaped adult plants and seedlings 90-120 days after the initial treatment (Mullahey et al., 1996). 

Integrated management

Integrated weed management strategies include prevention (avoidance of contaminated hay or grass seed, control of movement of cattle), control (mechanical, chemical, biological) and monitoring (Mullahey et al., 1998). 

Integrated control combining bioherbicides with chemical herbicides have been shown to be more effective than the chemical treatments alone (Roberts et al., 2002; Ferrell et al., 2008).

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Links to Websites

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WebsiteURLComment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.

Contributors

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06/05/08 Original text by:

Chris Parker, Consultant, UK

01/02/13 Updated by:

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

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

Distribution Maps

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