Invasive Species Compendium

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Datasheet

Solanum carolinense
(horsenettle)

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Datasheet

Solanum carolinense (horsenettle)

Summary

  • Last modified
  • 29 March 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Host Plant
  • Preferred Scientific Name
  • Solanum carolinense
  • Preferred Common Name
  • horsenettle
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae

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Pictures

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PictureTitleCaptionCopyright
TitleS. carolinense growing in pasture
Caption
CopyrightTomoko Nishida
S. carolinense growing in pastureTomoko Nishida

Identity

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Preferred Scientific Name

  • Solanum carolinense L. (1753)

Preferred Common Name

  • horsenettle

International Common Names

  • English: apple of Sodom; ball nettle; ball nightshade; bullnettle; Carolina horsenettle; Carolina nettle; devil's potato; devil's tomato; sand brier; wild tomato
  • Spanish: ortiga de caballo
  • French: morelle de la Caroline

Local Common Names

  • Germany: Carolina-Nachtschatten
  • Japan: warunasubi

EPPO code

  • SOLCA (Solanum carolinense)

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

Top of page The chromosome number of n=12, 2n=24 has been determined for Solanum carolinense plants in Canada (Bassett and Munro, 1986) and the USA (D'Arcy, 1969). Zutshi and Kaul (1974) reported that S. carolinense is tetraploid in India, with 2n=48.

S. carolinense is divided into three varieties: var. carolinense, var. floridanum, and var. hirsutum. The distribution areas of var. floridanum and var. hirsutum are restricted to Florida and Georgia, and Alabama and Georgia, USA, respectively (USDA-NRCS, 1999).

The common name 'apple of Sodom' is also used for Solanum hermannii in New Zealand (Upritchard, 1986).

Description

Top of page S. carolinense is a perennial broadleaf weed that is 30 to 120 cm tall, erect and loosely branched. The plant has an extensive root system with taproots and creeping, horizontal roots. Kiltz (1930) showed that vertical taproots would grow to depths of 2.4 m and that horizontal roots were found in the upper 45 cm of soil. The horizontal roots (themselves sprouting shoots) can spread to several metres from the taproots.

The stems of S. carolinense are angled at the nodes and are armed with sharp, stout, white or yellowish prickles, measuring up to 12 mm long. The stems are stellate-pubescent.

The leaves are alternate, simple and ovate to oblong in outline, with unevenly-lobed, toothed or deeply-cut margins. They are 4-14 cm long and about half as wide. Both surfaces are stellate-pubescent with yellowish hairs. There are sharp, yellow prickles on the veins, midrib and petioles.

The flowers resemble those of the potato and they are produced in secund racemes. The spineless calyx has five lobes with a sparse covering of hairs on its undersurface. The corolla is violet to whitish, with five lobes, and about 2-3 cm diameter. The flowers consist of five stamens, with prominent bright yellow anthers (6-9 mm long) in a cone surrounding the pistil.

The berries are globular, pulpy, juicy and smooth, each from 8 to 20 mm diameter. Immature berries are green, often with dark streaks. The mature berries are yellow, pale or yellowish-orange. A berry generally contains 40-170 seeds.

The seeds are obovate, flattened and slightly granulose. They are about 2-3 mm diameter and 0.5 mm thick. The mature seeds are pale to dark yellow, light brown or orange.

The hypocotyls of seedlings are often purple-tinged and covered with short, stiff hairs. The cotyledons are oval to oblong, about 1.2 mm long, hairy on the margins, glossy green on the upper surface and lighter below.

Distribution

Top of page S. carolinense is native to the Gulf States, and now it has spread across almost all of the USA and into southern Ontario, Canada (Bassett and Munro, 1986; USDA-NRCS, 1999). Lorenzi and Jeffery (1987) reported occurrence in all contiguous states of the USA, except Washington, Idaho, Montana, North Dakota, South Dakota, Wyoming, Utah, Colorado and New Mexico.

It is thought to have been introduced into Japan about 100 years ago (Tsuji, 1906) and has spread over all of the islands.

S. carolinense has been recorded in New Zealand (Healy, 1982) but the weed may by now have been eradicated (Upritchard, 1986).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

BangladeshPresentHolm et al., 1979
ChinaPresentLi et al., 2006
-ZhejiangPresentLi et al., 2006
Georgia (Republic of)PresentTrapidze, 1972
IndiaPresentZutshi and Kaul, 1974
JapanPresentPresent based on regional distribution.
-HokkaidoPresentSamejima et al., 1993
-HonshuPresentNishida et al., 1999a; Takei, 1997
-KyushuPresentOsada et al., 1967
-Ryukyu ArchipelagoPresentHatsusima, 1975
-ShikokuPresentAbe, 1990
Korea, Republic ofPresentPark et al., 2001
NepalPresentHolm et al., 1979

North America

CanadaPresentPresent based on regional distribution.
-OntarioRestricted distributionBassett and Munro, 1986
USAPresentPresent based on regional distribution.
-AlabamaWidespreadUSDA, 1971
-ArizonaPresentUSDA-NRCS and, 1999
-ArkansasWidespreadUSDA, 1971
-CaliforniaPresentUSDA-NRCS and, 1999
-ColoradoPresentUSDA-NRCS and, 1999
-ConnecticutPresentUSDA-NRCS and, 1999
-DelawareWidespreadUSDA, 1971
-FloridaPresentUSDA-NRCS and, 1999
-GeorgiaWidespreadUSDA, 1971
-IdahoPresentUSDA-NRCS and, 1999
-IllinoisWidespreadUSDA, 1971
-IndianaWidespreadUSDA, 1971
-IowaPresentUSDA-NRCS and, 1999
-KansasPresentUSDA-NRCS and, 1999
-KentuckyWidespreadUSDA, 1971
-LouisianaPresentUSDA-NRCS and, 1999
-MainePresentUSDA-NRCS and, 1999
-MarylandWidespreadUSDA, 1971
-MassachusettsPresentUSDA-NRCS and, 1999
-MichiganPresentUSDA-NRCS and, 1999
-MinnesotaPresentUSDA-NRCS and, 1999
-MississippiPresentUSDA-NRCS and, 1999
-MissouriWidespreadUSDA, 1971
-NebraskaPresentUSDA-NRCS and, 1999
-New HampshirePresentUSDA-NRCS and, 1999
-New JerseyWidespreadUSDA, 1971
-New YorkPresentUSDA-NRCS and, 1999
-North CarolinaWidespreadUSDA, 1971
-OhioWidespreadUSDA, 1971
-OklahomaPresentUSDA-NRCS and, 1999
-OregonPresentUSDA-NRCS and, 1999
-PennsylvaniaWidespreadUSDA, 1971
-Rhode IslandPresentUSDA-NRCS and, 1999
-South CarolinaWidespreadUSDA, 1971
-South DakotaPresentUSDA-NRCS and, 1999
-TennesseeWidespreadUSDA, 1971
-TexasPresentUSDA-NRCS and, 1999
-UtahPresentUSDA-NRCS and, 1999
-VermontPresentUSDA-NRCS and, 1999
-VirginiaWidespreadUSDA, 1971
-WashingtonPresentUSDA-NRCS and, 1999
-West VirginiaPresentUSDA-NRCS and, 1999
-WisconsinPresentUSDA-NRCS and, 1999

Central America and Caribbean

HaitiPresentD'Arcy, 1974

South America

BrazilPresentD'Arcy, 1974

Europe

CroatiaPresentGazi Baskova & Segulja, 1978
ItalyPresentViggiani, 2008
NetherlandsPresentDirkse et al., 2007
NorwayPresentOuren, 1987

Oceania

AustraliaPresentParsons and Cuthbertson, 1992
New ZealandEradicatedUpritchard, 1986

Habitat

Top of page S. carolinense is native to the Gulf States (Bassett and Munro, 1986) and is considered troublesome in the temperate to tropical zones of North America, Europe and Asia. It grows as a weed in grain and vegetable fields, orchards, pastures and nurseries. It is also found on roadside, in waste areas, riverbanks and, occasionally, in gardens. It grows in a wide range of soil types, but thrives in sandy or gravelly soils. The plant grows rapidly during hot weather and tolerates drought (Bradbury and Aldrich, 1957).

Biology and Ecology

Top of page S. carolinense is a perennial plant that propagates by seeds, roots and root cuttings. The germinable seed rate is very high in general, and seeds are considered to play an important role in the plant's dissemination (Ilnicki et al., 1962). Freshly harvested seeds are highly dormant, and alternating temperatures from 20 to 30°C increase germination. The seeds require temperatures above 15°C to germinate (Nishida et al., 2000). Light is not necessary for seed germination (Ilnicki et al., 1962) but may be involved in breaking dormancy (Suzuki, 1975). Nitrate (Ilnicki et al., 1962) or gibberellin (Suzuki, 1975) treatment can increase the germination percentage under certain conditions. The seedlings can emerge from depths of 10 cm (Ilnicki et al., 1962) and seeds retain their viability for at least 3 years when buried at depths of 8-12 cm (Brown and Porter, 1942).

S. carolinense grows well in sunny environments (Takematsu et al., 1979). The plant appears to thrive on sandy or gravelly soils, but will grow in any type of soil (Bradbury and Aldrich, 1957). Warm temperatures (Nishida et al., 1999a) and soil fertility favour growth. The plant has an extensive root system, and is drought tolerant (Bradbury and Aldrich, 1957). Roots are apparently susceptible to freezing temperatures; hence distribution in Canada is restricted to Ontario where deeply penetrating roots remain below the frost line; shoots are also killed by frost (Bassett and Munro, 1986).

S. carolinense blooms from early summer through autumn (Bassett and Munro, 1986; Nichols et al., 1991; Nishida and Harashima, 1995; Anderson, 1999). It is pollinated by bumble bees and carpenter bees (Hardin et al., 1972). The berries and seeds begin to mature by September (Bassett and Munro, 1986).

Both the vertical and horizontal roots have adventitious buds and very small root cuttings are capable of producing shoots. Ilnicki et al. (1962) reported that 10-cm root cuttings produced shoots at a planting depth of 40 cm, whereas 5-cm root cuttings produced shoots only at shallow planting depths (5 and 10 cm). Root cuttings appear to be susceptible to freezing temperatures (Bassett and Munro, 1986; Wehtje et al., 1987). Ilnicki et al. (1962) found that no shoots were produced from root cuttings that were exposed on the soil surface for 3 days.

Solomon (1983a) studied autoallelopathy in S. carolinense seed germination. Germination of S. carolinense seeds was inhibited by the incorporation of stem, root and leaf material into soil. The potency of the inhibition decreased with time and could be reversed by washing the inhibited seeds and replanting them in fresh soil. This appears to demonstrate that autoallelopathy in this species can result in density-dependant regulation of population size.

Takahashi et al. (1995) reported that an aqueous extract of S. carolinense considerably reduced the growth of lettuce seedlings. Wiepke and Glenn (1982) reported that S. carolinense extract in ethanol negatively affected the germination of turnip, and the growth of maize and soyabean radicles.

Notes on Natural Enemies

Top of page Frumenta nundinella is a host-specific herbivore of S. carolinense (Solomon, 1980). Two generations of this moth occur in New York, USA. In the spring it infests leaves, living within capsules formed from terminal leaves, and in the summer it lives within the fruits. The leaf and fruit infestation reduces fruit production of S. carolinense (Solomon, 1983b). Leptinotarsa juncta and Epitrix fuscula are host-specific herbivores of S. carolinense in Virginia, USA (Wise and Sacchi, 1996). L. juncta feeds on leaves, flowers and developing fruits. E. fuscula feeds on roots as larvae and on leaves as adults. Infestation by the two beetles reduced fruit production in the field by more than 75%. Tildenia inconspicuella [Keiferia inconspicuella] is a leaf-miner, which feeds specifically on solanaceous plants in the mid to south-western part of the USA (Gross, 1986). The moth feeds on S. carolinense throughout the growing season, over a course of three generations; 36-61% of leaves were infested by the moth in Illinois, USA.

In Japan, adults and larvae of Epilachna vigintioctopunctata feed on S. carolinense throughout the growing season, over a course of two generations (Imura, 1999). The beetle seriously infests leaves, flowers and fruits, causing defoliation in the autumn. Adults and larvae of Acanthocoris sordidus infest stems of S. carolinense gregariously.

Means of Movement and Dispersal

Top of page Natural Dispersal (non-biotic)

S. carolinense is disseminated by seeds, roots and root cuttings. Natural dispersal on a small scale could occur via seed dissemination. Horizontal roots can extend several metres from the taproot (Kiltz, 1930) and contribute to small-scale dissemination.

Vector Transmission

The seeds can maintain viability after passing through the digestive tract of cattle (Nishida et al., 1998), horses, pigs or sheep (Muensher, 1955). Ilinicki et al. (1962) mentioned that the berries may be eaten by farm animals and the seeds subsequently scattered over large areas in animal droppings.

Agricultural Practices

Tillage of fields infested with S. carolinense promotes the dissemination of the plant by cutting the roots and dragging them elsewhere (Urakawa, 2000), provided that other favourable conditions such as good weather conditions, less competitive crop plants and ineffective weed control are available. Harvesting operations may transport mature berries to other places, which also encourages dissemination.

Movement in Trade

Large-scale dissemination can occur by the contamination of a crop or commercial seeds with S. carolinense seeds. S. carolinense is thought to have been introduced to Japan from the USA via contamination of pasture plant seeds (Tsuji, 1906; Ono, 1965) and in fodder crop (Nishida and Shimizu, 1999) but there is no definite evidence to support this theory.

Impact

Top of page S. carolinense is a troublesome weed in pastures (see, for example, Nichols et al., 1991; Nishida et al., 1999a) and in field crops such as maize (Prostoko et al., 1994) and groundnuts (Hackett et al., 1987) in the USA, Canada and Japan. It is also a problem in vegetable fields (Bassett and Munro, 1986), orchards and tree nursery stock (Frank and King, 1979; Weatherspoon and Wooten, 1981).

S. carolinense is listed as a noxious weed under the Seeds Act and Regulation administered by Agriculture Canada (Bassett and Munro, 1986). It is also listed in the Noxious Seeds Act of Manitoba.

This species is a declared noxious weed and/or noxious-weed seed in 38 states of the USA (USDA, ARS, NGRP, http://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?100938).

S. carolinense is classed as one of the 10 most troublesome pasture weeds in the south-eastern USA (Smith and Calvert, 1980).

The plant causes yield losses due to its competition with crops. The presence of S. carolinense fruits in groundnut harvests affects the grade or quality assigned to the groundnuts (Woodruff, 1966).

Frank (1990) reported that S. carolinense grown for 3 years and 1 year prior to planting snapbeans (Phaseolus vulgaris), reduced yield by 48-65% and 18-20%, respectively. Hackett et al. (1987) observed that maintaining a weed-free environment for 2 or more weeks in a field previously infested with S. carolinense permitted an increase in the yield of runner-type groundnuts.

S. carolinense contains solanin and is poisonous to cattle, horses and sheep when ingested (Kingsbury, 1964). Carlisle et al. (1980) reported that S. carolinense contained a potentially toxic level of nitrate.

S. carolinense is also an important alternate host for insect pests of crop plants such as the Colorado potato beetle (Leptinotarsa decemlineata) (McIndoo, 1935) and the pepper maggot (Zonosemata electa) (Foott, 1963). It is also host for the potato psyllid (Paratrioza cockerelli [Bactericera cockerelli]), which transmits psyllid yellow disease to potatoes and tomatoes (Wallis, 1951), tomato leafspot fungus (Septoria lycopersici) (Pritchard and Porte, 1921) and several viruses (Weinbaum and Milbrath, 1976; Ramsdell and Myers, 1978).

Uses

Top of page S. carolinense may be used as a medicine (see King's American Dispensatory http://www.ibiblio.org/herbmed/electic/kings/solanum-caro.html; herbweb.com http://squid2.laughingsquid.net/hosts/herbweb.com/herbage/A25113.htm).

Similarities to Other Species/Conditions

Top of page S. carolinense is sometimes confused with Solanum elaeagnifolium, which is most common in the south-western USA and in Mexico (Bassett and Munro, 1986; Anderson, 1999). The entire plant of S. elaeagnifolium, except for the upper surface of the leaves, appears whitish, due to a covering of dense, canescent, tomentum stellate hairs, which S. carolinense lacks. S. elaeagnifolium is generally less prickly than S. carolinense.

Solanum dimidiatum is very similar to S. carolinense, though larger (Wehtje et al., 1987). The extent of the distribution of S. dimidiatum in the USA is lower than that of S. carolinense (USDA-NRCS, 1999). In many ways, S. dimidiatum seems to be an intermediary between S. carolinense and Solanum melongena (D'Arcy, 1974). It is distinct in its showy, deep-coloured flowers (deep mauve or blue, 3-4 cm diameter) and in its large, hard, yellow fruits. Fewer than 12 seeds are produced, twisted and packed due to the constraint of the stony endcarp.

Solanum aculeatissimum is also similar to S. carolinense, but prickles are present on the calyx of S. aculeatissimum. The immature berries of this species are white and the mature berries vermillion. The seeds are winged.

Physalis subglabrata [P. longifolia var. subglabrata] and Physalis heterophylla resemble S. carolinense, but do not have the conspicuous prickles on the stems and leaves (Uva et al., 1997). The berries of P. longifolia var. subglabrata and P. heterophylla are enclosed by an inflated, papery membrane.

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.

Cultural Control

As it is very difficult to control well-entrenched infestations of S.carolinense, prevention of its establishment is the most important and efficient means of control. Animal wastes which may contain viable seeds of S. carolinense should be treated before being used as manure. S. carolinense seeds lose viability when exposed to temperatures of 55 and 60°C for 72 h and 24 h, respectively (Nishida et al., 1999b). These temperatures and durations are considered adequate if the animal waste is fermented properly.

Ilnicki et al. (1962) reported that clipping the top growth, at least in July and August, prevented S. carolinense from producing viable seeds. As S. carolinense has an extensive root system and small root cuttings can produce shoots, tillage is believed to enhance the spread of the weed (Ilnicki et al., 1962; Takematsu et al., 1979; Smith and Calvert, 1980). However, S. carolinense has been reported to have become prevalent in conservation tillage systems (Burnside, 1981; Elmore et al., 1984). Young S. carolinense plants which sprout from root cuttings may be more susceptible to suppression by crop plants, and chemical and cultural control than well established plants. Muensher (1955) mentioned that a rotation which includes a clean cultivated crop every few years may reduce the presence of the weed, if the scattered plants appearing after the cultivation are hoed or pulled out.

Regehr and Janssen (1989) observed that the S. carolinense population declined significantly in ridge-till systems of a soyabean and sorghum rotation with herbicide treatments.

Biological Control

Izhevskii et al. (1981) studied the integrated control of S. carolinense using Tobacco mosaic virus str. Alke (TMV) and herbicides. The application of TMV to S. carolinense prior to or during early bud formation gave satisfactory control of the weed in tea plantations. A prior herbicide treatment is recommended to reduce the leaf surface area of other weeds, and to ensure that the virus spray is deposited only on S. carolinense leaves.

Chemical Control

S. carolinense is susceptible to a wide range of herbicides. Some are effective in the short term, while others demonstrate long-term effectiveness.

Albert (1960) concluded that in a pasture of Bermuda grass (Cynodon dactylon), the summertime application of 2,4-D over a course of several years would be practical, although the Ontario Weed Committee lists S. carolinense as resistant to 2,4-D (Bassett and Munro, 1986). Foliar application of picloram in summer is very effective in controlling the root system and shoots of S. carolinense. However, picloram poses a greater problem with persistence and its potential dispersion in the environment. Triclopyr, which is as effective as picloram when used at higher rates, may be appropriate for use in controlling S. carolinense in pastures (Gorrell et al., 1981).

Glyphosate is most effective when applied during the fruit-bearing period (Banks et al., 1977; Nashiki et al., 1985). Whitwell et al. (1980) reported that a high temperature (32°C) resulted in more effective injury to the shoot but a low temperature (13°C) during glyphosate treatment resulted in much less regrowth.

In maize, post-emergence application of dicamba may be advisable (Prostko et al., 1994).

Talbert et al. (1982) reported that spot applications of acifluorfen were effective in controlling S. carolinensis and minimized potential damage to developing strawberry buds, although repeated applications at 2 to 3 week intervals were required.

In groundnuts, subsurface layered dinitramine or post-emergence 2,4-DB application provided acceptable levels of control (Banks and Samtelmann, 1978; Hackett et al., 1987).

S. carolinense is also suceptible to amitrole (Albert, 1960), silvex [fenoprop], terbacil (Pagano, 1975) and maleic hydrazide (Takematsu et al., 1979). The seedlings are suceptible to atrazine, cloransulam and metribuzin (Vangessel, 1999).

When herbicides are used, it is necessary to adhere to local restrictions and regulations governing herbicide use.

References

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Abe K, 1990. Tokushimakenshokubutsushi. Tokushima, Japan: Kyoikushuppancenter.

Albert WB, 1960. Control of horsenettle (Solanum carolinense) in pastures. Weed Science, 8:680-682.

Anderson WP, 1999. Perennial Weeds. Ames, USA: Iowa Sate University Press.

Banks PA; Kirby MA; Santelmann PW, 1977. Influence of postemergence and subsurface layered herbicides on horsenettle and peanuts. Weed Science, 25(1):5-8

Banks PA; Santelmann PW, 1978. Influence of subsurface layered herbicides on horsenettle and various crops. Agronomy Journal, 70(1):5-8

Bassett IJ; Munro DB, 1986. The biology of Canadian weeds. 78. Solanum carolinense L. and Solanum rostratum Dunal. Canadian Journal of Plant Science, 66(4):977-991

Bradbury HE; Aldrich RJ, 1957. Survey reveals extent of horse nettle infestation. New Jersey Agriculture, 39(4):4-7.

Brown EO; Porter RH, 1942. The viability and germination of seeds of Convolvulus arvensis L. and other perennial weeds. Iowa Agricultural Experimant Station Research Bulletin, 294:475-504.

Burnside OC, 1981. Changing weed problems with conservation tillage. In: Conference on Crop Production with Conservation in the 1980's. American Society of Agricultural Engineering Publication, 167-174.

Carlisle RJ; Watson VH; Cole AW, 1980. Canopy and chemistry of pasture weeds. Weed Science, 28(2):139-141

D'Arcy WG, 1969. Solanaceae. In: Doe J, ed. Chromosome Numbers of Phanerogams. 3. Annals of the Missouri Botanical Garden, 56:471-472.

D'Arcy WG, 1974. Solanum and its close relatives in Florida. Annals of the Missouri Botanical Garden, 61:819-867.

Dirkse G; Holverda W; Hochstenbach S; Reijerse F, 2007. Solanum carolinense L. and Pimpinella peregrina L. in The Netherlands. (Solanum carolinense L. en Pimpinella peregrina L. in Nederland.) Gorteria, 33(1):21-27. http://www.nationaalherbarium.nl/pubs/gorteriaweb/home.htm

Elmore CD; Wesley R; Cooke F; Hurst H, 1984. Effects of tillage and irrigation on weeds in a wheat-soybean double-cropping system. Proceedings, Southern Weed Science Society, 37th annual meeting, 316

Foott WH, 1963. The biology and control of the pepper maggot, Zonosemata electa (Say)(Diptera: Trypetidae) in Southwestern Ontario. Proceedings of Entomology Society of Ontario, 93:75-81.

Frank JR, 1990. Influence of horsenettle (Solanum carolinense) on snapbean (Phaseolus vulgaris). Weed Science, 38(3):220-223

Frank JR; King JA, 1979. Metolachlor and alachlor for weed control in establishing woody nursery stock. Proceedings, Northeastern Weed Science Society, 33:228-231

Gazi-Baskova V; Segulja N, 1978. The appearance of dangerous weeds of the genus Solanum on the Kvarner Island of Plavnik. Fragmenta Herbologica Jugoslavica, 6(106-115):55-59

Gorrell RM; Bingham SW; Foy CL, 1981. Control of horsenettle (Solanum carolinense) fleshy roots in pastures. Weed Science, 29(5):586-589

Gross P, 1986. Life histories and geographic distributions of two leafminers, Tildenia georgei and T. inconspicuella (Lepidoptera: Gelechiidae), on solanaceous weeds. Annals of the Entomological Society of America, 79(1):48-55

Hackett NM; Murray DS; Weeks DL, 1987. Interference of horsenettle (Solanum carolinense) with peanuts (Arachis hypogpa). Weed Science, 35(6):780-784

Hardin JW; Doerksen G; Herndon D; Hobson M; Thomas F, 1972. Pollination ecology and floral biology of four weedy genera in southern Oklahoma. Southwest Naturalist, 16:403-412.

Hatsusima S, 1975. Flora of the Ryukyus. Naha, Japan: Okinawa seibutsu kyoiku kenkyukai.

Healy AJ, ed. , 1982. Identification of weeds and clovers. Featherston, New Zealand: New Zealand Weed and Pest Control Society, Edn. 3:299 pp.

Holm LG; Pancho JV; Herberger JP; Plucknett DL, 1979. A geographical atlas of world weeds. New York, USA: John Wiley and Sons, 391 pp.

Ilnicki RD; Tisdell TF; Fertig SN; Furrer AH Jr, 1962. Life history studies as related to weed control in the Northeast. 3. Horse-nettle. University of Rhode Island Agricultural Experiment Station Bulletin, 368.

Imura O, 1999. Interactions between herbivorous arthropods and introduced plants in agro-ecosystems. In: Yano E, Matsuo K, Shiyomi M, Andow D, eds. Biological Invasions of Ecosystem by Pests and Beneficial Organisms. Tsukuba, Japan: NIAES, 152-163.

Izhevskii SS; Livshits p; Murusidze GE; Gogoladze GG, 1981. Prospects of using Alke strain in integrated control of Solanum carolinense in tea plantations. Subtropicheskie Kul'tury, No.4:60-65

Kiltz BF, 1930. Perennial weeds which spread vegetatively. Journal of the American Society of Agronomy, 22:216-234.

Kingsbury JM, 1964. Poisonous plants of the United States and Canada. Englewood Cliffs, New Jersey, USA: Prentice-Hall Inc.

Li GenYou; Jin ShuiHu; Ai JianGuo, 2006. Species, characteristics and control measures of injurious plants in Zhejiang Province. Journal of Zhejiang Forestry College, 23(6):614-624. http://zjlx.chinajournal.net.cn

Lorenzi HJ; Jeffery LS(Editors), 1987. Weeds of the United States and their control. New York, USA; Van Nostrand Reinhold Co. Ltd., 355 pp.

McIndoo NE, 1935. The relative attractiveness of certain Solanaceous plants to the Colorado potato beetle, Leptinotarsa decemlineata Say. Proceedings of Entomology Society Washington, 37(2):36-42.

Muenscher WC, 1955. Weeds, 2nd edition. New York, USA: Macmillan, 560 pp.

Nashiki M; Nomoto T; Meguro R, 1985. Glyphosate application time for sod-seeding orchardgrass establishment in pasture infested with horsenettle (Solanum carolinense L.). Weed Research, Japan, 30(2):131-136

Natsuaki KT; Kashina H; Suzuki T; Tomaru K, 1992. Tobacco mosaic virus isolated from Solanum carolinense L. in Japan. Journal of Agricultural Science, Tokyo Nogyo Daigaku, 37(3):253-259

Nichols RL; Cardina J; Gaines TP, 1991. Growth, reproduction and chemical composition of horsenettle (Solanum carolinense). Weed Technology, 5(3):513-520

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