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Datasheet

Sicyos angulatus (burcucumber)

Summary

  • Last modified
  • 13 July 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Host Plant
  • Preferred Scientific Name
  • Sicyos angulatus
  • Preferred Common Name
  • burcucumber
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • S. angulatus is characteristically a weed of maize, soyabean and sorghum crops (Anon., 2010). This is the case in its native range (it is officially classified as a noxious weed in Delaware and Indiana, and as a...

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Pictures

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PictureTitleCaptionCopyright
Oneseed burr cucumber (Sicyos angulatus) climbing trees adjacent to a roadside.
TitleInvasive habit
CaptionOneseed burr cucumber (Sicyos angulatus) climbing trees adjacent to a roadside.
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus) climbing trees adjacent to a roadside.
Invasive habitOneseed burr cucumber (Sicyos angulatus) climbing trees adjacent to a roadside.Shunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); infested soybean field.
TitleInvasive habit
CaptionOneseed burr cucumber (Sicyos angulatus); infested soybean field.
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); infested soybean field.
Invasive habitOneseed burr cucumber (Sicyos angulatus); infested soybean field.Shunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); riverside population showing invasive nature.
TitleInvasive habit
CaptionOneseed burr cucumber (Sicyos angulatus); riverside population showing invasive nature.
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); riverside population showing invasive nature.
Invasive habitOneseed burr cucumber (Sicyos angulatus); riverside population showing invasive nature.Shunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); abandoned field infested.
TitleInvasive habit
CaptionOneseed burr cucumber (Sicyos angulatus); abandoned field infested.
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); abandoned field infested.
Invasive habitOneseed burr cucumber (Sicyos angulatus); abandoned field infested.Shunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); infested maize field.
TitleInvasive habit
CaptionOneseed burr cucumber (Sicyos angulatus); infested maize field.
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); infested maize field.
Invasive habitOneseed burr cucumber (Sicyos angulatus); infested maize field.Shunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); open flowers and buds on stem.
TitleFlower
CaptionOneseed burr cucumber (Sicyos angulatus); open flowers and buds on stem.
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); open flowers and buds on stem.
FlowerOneseed burr cucumber (Sicyos angulatus); open flowers and buds on stem.Shunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); developing fruits.
TitleFruit
CaptionOneseed burr cucumber (Sicyos angulatus); developing fruits.
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); developing fruits.
FruitOneseed burr cucumber (Sicyos angulatus); developing fruits.Shunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); cotyledon stage
TitleSeedling
CaptionOneseed burr cucumber (Sicyos angulatus); cotyledon stage
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); cotyledon stage
SeedlingOneseed burr cucumber (Sicyos angulatus); cotyledon stageShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); immature plant.
TitleJuvenile stage
CaptionOneseed burr cucumber (Sicyos angulatus); immature plant.
CopyrightShunji Kurokawa
Oneseed burr cucumber (Sicyos angulatus); immature plant.
Juvenile stageOneseed burr cucumber (Sicyos angulatus); immature plant.Shunji Kurokawa

Identity

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Preferred Scientific Name

  • Sicyos angulatus Linnaeus, 1753

Preferred Common Name

  • burcucumber

International Common Names

  • English: one seed burcucumber; oneseed burr cucumber
  • Spanish: calabacilla; chayotillo
  • French: concombre anguleux

Local Common Names

  • Germany: Kantenblatt- Haargurke
  • Japan: arechiuri
  • Korea, Republic of: gasibak
  • Netherlands: Stekelaugurk
  • Spain: calabacilla

EPPO code

  • SIYAN (Sicyos angulatus)

Summary of Invasiveness

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S. angulatus is characteristically a weed of maize, soyabean and sorghum crops (Anon., 2010). This is the case in its native range (it is officially classified as a noxious weed in Delaware and Indiana, and as a weed in Kentucky and Nebraska) (Anon., 2010). In Europe, it is mainly found in irrigated maize fields, but it has not been found in soyabean and sorghum fields. Densities of soyabean plants are lower than in maize, impeding S. angulatus climbing over the plant, and sorghum is not irrigated (Anon., 2010). The most problematic characteristic of S. angulatus is its rapid growing long vines. It can reach 5 to 8 m, often entangling and covering crops or natural vegetation. The dispersal of spiny fruits in flowing water may aid rapid local expansion of S. angulatus (Kil et al., 2006). S. angulatus is on the EPPO List of Invasive Alien Plants. In Japan, S. angulatus was also designated as an invasive alien species, based on the parameters of the Invasive Alien Species Act on February 1, 2006. It grows in fields of maize and of sorghum for tillage, in Japan (Anon., 2010). It is not a strong competitor for light and nutrients, so does not reduce yields by direct competition; however, it pulls maize or soybean plants to the ground, making them impossible to harvest (Anon., 2010). Maize stems are broken by the traction and weight of the invasive plant (Anon., 2010). In Korea, it is regulated under the Wildlife Protection Act.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Violales
  •                         Family: Cucurbitaceae
  •                             Genus: Sicyos
  •                                 Species: Sicyos angulatus

Notes on Taxonomy and Nomenclature

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The name burcucumber is derived from the appearance of the dried fruit (Mann et al., 1981). The common name in Japan “arechiuri” means “ruderal Cucurbitaceae plant”. The Spanish name “calabacilla” means “gourd”, and the Korean name “gasibak” means “spiny Cucurbitaceae plant”.

Description

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The leaves of S. angulatus are thin, 5-lobed, up to 25 cm across and borne on stout, pubescent petioles 2.5 to 10 cm long (Mann et al., 1981). They are alternate, broadly heart-shaped and finely toothed (Anon., 2010). Stems are hairy and form a creeping vine up to 6 m long, with numerous branched tendrils (Anon., 2010). New vines can form by growth from axillary buds (Mann et al., 1981). The root system consists of a shallow branched taproot (Anon., 2010).

S. angulatus is monoecious with 5-petalled, green and white flowers (Mann et al., 1981). The male flowers appear in a corymbose raceme on a very long peduncle and the female flowers appear in a capitate cluster on a short peduncle (Torrey and Gray, 1969). The calyx is green, five-toothed, and pubescent. The corollas of both sexes are white with green striations, and consist of five petals fused at the base into an open bowl, and free and spread at the tips. Staminate flowers form on either paniculate or racemose inflorescences. The anthers unite to form a central column. Pistillate flowers are borne on a compact cyme, in a globose cluster of 8-20 flowers. The pistil consists of a superior ovary, a slender style,and 3 stigmas (Medley and Burnham, 2011).

The bur-like fruits are small and spiny, 1.0-1.5 cm long, one-seeded, produced in clusters of 3-20, initially green, turning brown, indehiscent, containing a single brown flattened seed (Anon., 2010). Each fruit contains one seed, and 3 to 15 fruits are borne in a cluster (Mann et al., 1981). The seeds are large (15 mm by 10 mm), dark brown to black, compressed, smooth, and covered with a crustaceous pericarp (Britton and Brown, 1947). The pericarp is actually covered with four types of “hairs” consisting of barbed prickles (8 to 10 mm long), long jointed hairs, short pointed conical hairs, and unicellular conical hairs (Barber, 1909).

Plant Type

Top of page Annual
Broadleaved
Herbaceous
Seed propagated
Vine / climber

Distribution

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S. angulatus in its native range is distributed throughout the eastern part of North America. Only by three USA state governments (Delaware, Indiana and Kentucky), is this species recognized as a noxious weed (USDA-NRCS, 2009). However, Esbenshade et al. (2001a,b,c) reported that S. angulatus has become a more serious problem in agronomic crops throughout much of the northeastern USA. In its introduced range, S. angulatus is distributed in eastern Asia, western Asia, and Europe. Its most northerly point is Norway (59°N) and most southerly point is Martinique (14°N). There is no report of their distribution in the southern hemisphere.

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

ChinaPresentEPPO, 2014
Georgia (Republic of)PresentIntroducedMikeladze et al., 2015
IndiaPresentIntroducedPresent based on regional record
-Himachal PradeshPresentIntroducedThakur,, 2016
JapanPresentEPPO, 2014
-HonshuPresentIntroduced1952 Invasive Shimizu et al., 2001; Watanabe et al., 2002; Kurokawa et al., 2009Shimizu port in Shizuoka
-ShikokuPresentIntroducedYamamoto et al., 1982
Korea, Republic ofPresentIntroduced Invasive Kil et al., 2006; EPPO, 2014
TurkeyPresentIntroducedTerzioglu and Ansin, 1999; EPPO, 2014

North America

CanadaRestricted distributionEPPO, 2014
-OntarioPresentNative Not invasive Britton and Brown, 1947; USDA-NRCS, 2009; EPPO, 2014
-QuebecPresentNative Not invasive Britton and Brown, 1947; USDA-NRCS, 2009; EPPO, 2014
MexicoPresentEPPO, 2014
USARestricted distributionEPPO, 2014
-AlabamaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-ArkansasPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-ConnecticutPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-DelawarePresentNative Invasive Webb and Johnston, 1981; USDA-NRCS, 2009; EPPO, 2014
-FloridaPresentNative Not invasive Britton and Brown, 1947; USDA-NRCS, 2009; EPPO, 2014
-GeorgiaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-IllinoisPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-IndianaPresentNative Invasive Gibson et al., 2005; EPPO, 2014South central and south east districts
-IowaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-KansasPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-KentuckyPresentNative Invasive USDA-NRCS, 2009; EPPO, 2014
-LouisianaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-MainePresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-MarylandPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-MassachusettsPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-MichiganPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-MinnesotaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-MississippiPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-MissouriPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-NebraskaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-New HampshirePresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-New JerseyPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-New YorkPresentNative Not invasive USDA-NRCS, 2009; Anon., 2010
-North CarolinaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-North DakotaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-OhioPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-OklahomaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-PennsylvaniaPresentNative Invasive Esbenshade et al., 2001a; Esbenshade et al., 2001b; Esbenshade et al., 2001c; USDA-NRCS, 2009; EPPO, 2014
-Rhode IslandPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-South CarolinaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-South DakotaPresentNative Not invasive Britton and Brown, 1947; USDA-NRCS, 2009; EPPO, 2014
-TennesseePresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-TexasPresentNative Not invasive Britton and Brown, 1947; USDA-NRCS, 2009; EPPO, 2014
-VermontPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-VirginiaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-West VirginiaPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014
-WisconsinPresentNative Not invasive USDA-NRCS, 2009; EPPO, 2014

Central America and Caribbean

GuadeloupePresentEPPO, 2014
MartiniquePresentEPPO, 2014

Europe

AustriaPresentEPPO, 2014
BulgariaPresentIntroducedTzonev, 2005; Anon., 2010; EPPO, 2014Belene (Persina) Island
CroatiaPresentIntroduced1869 Invasive Hulina, 1996; EPPO, 2014Granesina
Czech RepublicPresent, few occurrencesEPPO, 2014
FinlandAbsent, formerly presentIntroducedFlora Nordica, 2009
FranceRestricted distributionIntroduced1981 Invasive Larché, 2004; Chauvel et al., 2005; EPPO, 2014
GermanyPresentEPPO, 2014
GreecePresentIntroduced2002 Invasive Anagnou-Veroniki et al., 2008; EPPO, 2014First record is in Kavala (northern Greece) in 2002
HungaryPresentAnon., 2010; EPPO, 2014
ItalyPresentIntroduced Invasive Pignatti, 1982; Rapparini, 1999; Fabbi et al., 2006; EPPO, 2014
MoldovaPresentEPPO, 2014
MontenegroPresentIntroducedStesevic and Jovovic, 2005In tomato crops in Jasenova village
NorwayAbsent, formerly presentOuren, 1987; EPPO, 2014
PolandPresentEPPO, 2014
RomaniaPresentEPPO, 2014
Russian FederationRestricted distributionEPPO, 2014
SerbiaPresentEPPO, 2014
SloveniaPresentEPPO, 2014
SpainPresentAnon., 2010; EPPO, 2014
SwedenAbsent, unreliable recordEPPO, 2014
UKPresentEPPO, 2014
UkrainePresentEPPO, 2014
Yugoslavia (former)PresentIntroducedTrinajstic and Dubravec, 1975

History of Introduction and Spread

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S. angulatus is native to the northeastern USA. It has been intentionally or unintentionally introduced to Asia and Europe.

It was never intentionally introduced to Japan. Nevertheless, herbarium records show a rapid expansion of its distribution throughout Japan, except for Hokkaido and Okinawa, after it was found in Shizuoka for the first time in 1952 (Shimizu et al., 2001). Soybean imported from the USA is suspected as the source of unintentional introduction (Shimizu et al., 2001). Its occurrence in the southwestern region of Japan is very rare, in the central and northeastern regions, the problems caused by this species are serious (Kurokawa et al., 2009). Inter-simple sequence repeat (ISSR) genotyping suggested multiple introductions from the same gene pools into both central and northeastern regions of Japan (Kurokawa et al., 2009). In Korea, it has been rapidly spreading and has achieved nationwide distribution in 15 years (Kil et al., 2006).

It was intentionally introduced into Europe for ornamental purposes (Hulina, 1996) in the 19th century (Anon., 2010). It was listed in the Flora of Croatia in 1869 (Hulina, 1996). It entered the UK as a contaminant of bird seed (Hanson and Mason, 1985). The first report of its presence in Greece was in 2002 (Anagnou-Veroniki et al., 2008). In France, it first appeared on irrigated maize crop plots in the French Basque country in 1981 (Larché, 2004). In France, it appears to have been introduced as a result of successive imports of maize from the USA (Larché, 2004). It has spread to the Landes, Gironde, Lot et Garonne and Dordogne regions (Larché, 2004). In Norway, it was registered as an adventive weed in soybean in the 1970s and 1980s (Ouren, 1987). S. angulatus is thought to have been introduced in Spain first and spread east and north-east by natural means (JF Larché, personal communication, 2007 in Anon., 2010).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Europe USA   Live food or feed trade (pathway cause) Yes No Clement and Foster (1994); Hanson and Mason (1985); Stace (1997)
Europe 1800s Ornamental purposes (pathway cause) Yes No Hulina (1996)
France USA 1981 Live food or feed trade (pathway cause) Yes No Larché (2004)
Honshu USA 1952 Live food or feed trade (pathway cause) Yes No Shimizu et al. (2001)

Risk of Introduction

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Grain importation is the most likely pathway of accidental introduction of S. angulatus, considering the situation in Japan, France or Norway (Ouren, 1987; Shimizu et al., 2001; Larché, 2004). Although the distribution record for S. angulatus in Mexico is unreliable (EPPO, 2009), the possibility of it being introduced unintentionally into Mexico is high as the amount of soybean and maize imported from the USA is the second highest in the world (3.4 million and 5.8 million ton per year, respectively) (FAO, 2009).

The rapid local spread of S. angulatus may occur via dispersal in flowing water (Kil et al., 2006). Establishment of it along riversides may present a higher risk for local spread.

Habitat

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In its native range, S. angulatus grows mainly on riverbanks or in thickets. In its introduced range, it is mainly found on riverbanks or soils close to rivers (Larché, 2004; Kurokawa et al., 2009). In India, it is adventive in a forest area of Shimla (Himachal Pradesh) (Thakur, 2016). This plant grows mostly as a weed of maize crops, but can also colonize soybeans (Anon., 2010). In Japan, forage crop fields are one of the most frequently infested habitats (Watanabe et al., 2002).

Habitat List

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CategoryHabitatPresenceStatus
Terrestrial-managed
Cultivated / agricultural land Principal habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Natural
Rail / roadsides Principal habitat Natural
Terrestrial-natural/semi-natural
Natural forests Secondary/tolerated habitat
Riverbanks Principal habitat Natural

Hosts/Species Affected

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With severe infestations, S. angulatus vines can smother and pull a crop to the ground, making crop harvesting nearly impossible (Esbenshade et al., 2001c). It is a problem weed in some summer crops, such as maize, soybeans and pumpkins (Hulina, 1996; Messersmith et al., 1999; 2000; Shimizu, 1999; Esbenshade et al., 2001a; Watanabe et al., 2002; Larché, 2004; Gibson et al., 2005). In Japan, the maize yield was decreased by 80% by a population of 15-20 plants and by 90-98% with a population of 28-50 plants (Shimizu, 1999).

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Cucurbita (pumpkin)CucurbitaceaeOther
Glycine max (soyabean)FabaceaeOther
Zea mays (maize)PoaceaeMain

Growth Stages

Top of page Flowering stage, Fruiting stage, Pre-emergence, Seedling stage, Vegetative growing stage

Biology and Ecology

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Genetics

The chromosome number of S. angulatus is 2n=24 (McKay, 1930). ISSR analysis revealed that the total gene diversity (Ht) in two areas in Japan was 0.1684, although genetic diversity in its native range is unknown (Kurokawa et al., 2009). The coefficient of gene differentiation was low amongst the two geographically distinct regions (Gst =0.053) (Kurokawa et al., 2009).

Reproductive Biology

S. angulatus plants grown without competition produced 716-18,197 g dry matter and 4,500-78,000 seeds per plant (Esbenshade et al., 2001c; Smeda and Weller, 2001). Plants established in maize produced 96% less dry matter than plants in the non-competitive treatments and the competition from maize reduced S. angulatus seed production by 98% compared to plants grown without a crop (Esbenshade et al., 2001c).

Physiology and Phenology

Seeds have a water-impermeable, hard seed coat which means that they may persist in the soil seed bank until suitable conditions for germination prevail (Ozaslan et al., 2016). Intact seeds of this species are rarely germinated without removing their arils before sowing (Kang et al., 2003a). Seeds that have undergone sequential treatments of aging, NaOH, washing, combining chilling and priming and drying under red light illumination were germinated up to 80% (Kang et al., 2003a). The sequential treatments of aging, cold-stratification, and red light illumination during desiccation can highly promote percentage and speed of S. angulatus seed germination (Kang et al., 2003b). Concerning seed scarification, girdling had the highest germination percentage of 79% (Lim et al. 1994b). S. angulatus shows high variation in seed maturity and size (Jeon et al., 2003). Mature seeds showed a higher germination and seedling emergence rate than immature ones (Jeon et al., 2003). Medium and small seeds showed a higher germination rate than large ones (Jeon et al., 2003). Medium and large seeds, however, had the greatest and the least seedling emergence, respectively (Jeon et al., 2003). Seedling height, number of true leaves, areas of cotyledons and true leaves except hypocotyl length were increased with increased with seed size (Jeon et al., 2003). Cotyledon, leaf, hypocotyl, root and their total dry weights were greater in large mature seeds than larger immature ones (Jeon et al., 2003). S. angulatus germination is greatest at soil temperatures between 20 and 30°C, with germination being drastically reduced at temperatures below 10 and above 35°C (Mann et al., 1981; Curran et al., 2000). Scarified seeds germinated at osmotic potentials to -6 bars (Mann et al., 1981). Cold stratification at 4°C for 18 weeks modified seed coat permeability (Mann et al., 1981).

S. angulatus has the ability to germinate throughout the growing season (Messersmith et al., 1999). The emergence appears to correlate with rainfall (Smeda and Weller, 2001). It germinates in Kentucky from June to September (Mann et al., 1981). Under Indiana conditions, it germinates from late April to October and is stimulated by periodic rainfall (Smeda and Weller, 2001). Percent emergence of S. angulatus was highest in the 1 to 5 cm depth (Messersmith et al., 2000). Increasing the depth of planting decreases emergence with limited emergence occurring at depths of 15 cm in field (Mann et al., 1981). Fewer S. angulatus emerged from depths of 10 cm or more or when S. angulatus was placed on the soil surface (Messersmith et al., 2000). Under no-till conditions, S. angulatus seeds experienced higher mortality rates on the soil surface due to poor seed-to-soil contact, germination and subsequent desiccation, predators, and parasites (Roberts and Feast, 1972; Froud-Williams et al., 1981; Zorner et al., 1984; Reader, 1991).

Relative growth rates of S. angulatus plants were greatest up to 10 weeks after establishment and declined when flowering was initiated (Smeda and Weller, 2001). S. angulatus remains strictly vegetative under long-day conditions (Kurata, 1977; Takahashi et al., 1982). Once an optimum accumulated temperature is reached, flowering in S. angulatus is more likely due to photoperiod length (Esbenshade et al., 2001c). Its flowers bloom during August and September in Kentucky (Mann et al., 1981). S. angulatus established in maize on July 24 or later did not flower, thus eliminating their ability to reproduce (Esbenshade et al., 2001c). The flowering of this plant was found to be induced by grafting it onto the induced donor plant of the same species or by intergeneric grafting onto day-neutral cucumbers or a quantitative short-day plant Luffa (Takahashi et al., 1982).

S. angulatus showed less sensitivity to low root temperature than Cucumis ficifolius (Bulder et al., 1991). The sterol/phospholipid ratio showed a clear and consistent reaction to low temperature treatment (Bulder et al., 1991). Bioassay by the sandwich method showed that S. angulatus has strong allelopathic activity (Uraguchi et al., 2003).

Associations

The levels of cucurbitacin B and E-glucoside, which are strong feeding stimulants for the leaf beetle species, Aulacophora indica and Aulacophora lewisii, were abundant in S. angulatus (Abe and Matsuda, 2005). S. angulatus is a host for tobacco budworm (Heliothusvirescens), which can lead to further losses in some crops that are susceptible to insect feeding (Smeda and Weller, 2001). S. angulatus was first observed at Stoneville, Mississippi, in 1972, and larval infestations of H. virescens were subsequently found in 1973-75 (Laster et al., 1976). This is the first record of the noctuid infesting the plant in Mississippi (Laster et al., 1976). It was thought that S. angulatus may become important in the build-up of early season and overwintering populations of the pest (Laster et al., 1976).

S. angulatus is susceptible to cucumber mosaic virus, watermelon mosaic virus (Schmelzer and Wolf, 1971; Yamamoto et al., 1982), and cucumber green mottle mosaic virus (Horváth, 1985).

Cucumber plants grafted on S. angulatus show higher nematode resistance properties than self-rooted plants (Gu et al., 2006).

Environmental Requirements

Although S. angulatus requires adequate soil moisture, it adapts to wide range of environment conditions. As S. angulatus established in maize produced 96% less dry matter and seed than the plants in a non-competitive environment (Esbenshade et al., 2001c), it may not be tolerant to shading.

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Tolerated > 60mm precipitation per month
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Df - Continental climate, wet all year Preferred Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
14-59

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 15 35

Soil Tolerances

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Soil reaction

  • acid
  • alkaline
  • neutral

Soil texture

  • heavy
  • medium

Notes on Natural Enemies

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The cotton caterpillar (Pyralidae indica) and leaf beetle species of the genus Aulacophora show a preference for S. angulatus leaves (Choi et al., 2003; Abe and Matsuda, 2005), but it is unknown whether they act as natural enemies. Larval infestations of the tobacco budworm (Heliothis virescens) were observed in Mississippi (Laster et al., 1976). Other insects that feed on S. angulatus include: the chrysomelids Psylliodes punctulata, Systena blanda, Acalymma vittata and Diabrotica undecimpunctata, Epilachna borealis, Anasa tristis and Poecilocapsus lineatus, Aphis gossypii, Diaphania hyalinata and Diaphania nitidalis, and Melittia cucurbitae; however, none of these insects efficiently limits stands (Anon., 2010).

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

Seeds of S. angulatus spread by flooding and harvest equipment (Bradley, 1984; Kil et al., 2006). Dispersal by water has been confirmed by Kurokawa et al. (2009), who performed inter-simple sequence repeat (ISSR) analysis on the widely distributed Japanese populations of S. angulatus, to infer the genetic relationship among populations. The fruit has spines that allow it to attach to animals and people (JM Tison, personal communication, 2007 in Anon., 2010). Natural dispersal (e.g. small mammals, birds) or dispersal by agricultural practices, transports fruit over short distances. Heavy rains leading to soil erosion and floods greatly amplify seed export (Kil et al., 2006).

Accidental Introduction

Introductions of S. angulatus into France or Japan were suspected as a result of successive imports of maize or soybeans from the USA (Larché, 2004; Kurokawa et al., 2009). It is also thought that the first introduction of the species into the Tama-river (Tokyo) was thought to be derived from the contaminants of soyabean (imported) dumped by tofu factories onto the riverside (S Kurokawa, personal communication, 2007 in Anon., 2010).

Intentional Introduction

In the nineteenth century S. angulatus was introduced into Europe for ornamental purposes (Hulina, 1996). Utilization of S. angulatus as a rootstock has been reported in Asia and Europe (Manabe et al., 1978; Tachibana, 1982; Bulder et al., 1991; Lim et al., 1994a,b; Kim et al., 1997; Gu et al., 2006; Zhang et al., 2006; Huitrón et al., 2008; Zhang et al., 2008); however, there is no report of intentional introduction for rootstock purpose.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Flooding and other natural disastersIn South Korea, it initiated colonization and established a population on the riverside Yes Kil et al., 2006
Live food or feed tradeIntroduction into France and Japan from USA Yes Kurokawa et al., 2009; Larché, 2004
Ornamental purposesIn the 1800s it was introduced into Europe for ornamental purposes Yes Hulina, 1996

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Containers and packaging - non-woodSeed Yes Kurokawa et al., 2009
WaterSeed Yes Kil et al., 2006

Impact Summary

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CategoryImpact
Economic/livelihood Negative
Environment (generally) Negative

Economic Impact

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In Japan, the maize yield was decreased by 80% by a population of 15-20 plants per 10 m2 and by 90-98% with a population of 28-50 plants (Shimizu, 1999).

S. angulatus is perceived as a problematic summer weed primarily by maize and soybean growers in the south central and southeast districts in Indiana (Gibson et al., 2005).

It was reported to be the seventh most troublesome weed in Kentucky soybean (Dowler, 1995) and the sixth most troublesome weed in maize (Dowler, 1994). It is also a problem weed in maize in North Carolina and Tennessee (Dowler, 1994). Bradley (1984) estimated that in 1983, S. angulatus was distributed throughout 75 of 95 counties in Tennessee and had infested over 25,000 ha.

S. angulatus is a host of the polyphagous pest, Heliothis virescens in North America (which attacks field crops such as cotton, tobacco, legumes and vegetables). S. angulatus could contribute to the build-up of early season and overwintering populations of the pest (Pheloung et al., 1999). A similar situation could arise for Helicoverpa armigera in Europe (Anon., 2010).

Recasens et al. (2007) (in Anon., 2010) have estimated the cost of the eradication campaigns in Catalunya between 2004 and 2010 at EUR 78,320.

Environmental Impact

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It is common along watercourses and in open spaces, suppressing native vegetation in Japan (Watanabe et al., 2002). It occupies 319 ha of vegetation area in 24 rivers in Japan (Miyawaki and Washitani, 2004). As stated by Anon. (2010), in invaded riversides in the Republic of Korea, observations show that massive germination can lead to a 100% cover of the soil layer during the growing season. Average and highest seed densities at full maturation were 748 and 1128 seeds/m2, respectively. A few seedlings per 10m2 were enough to cover the whole grass mat by July (Kil et al., 2006).

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Highly mobile locally
  • Fast growing
  • Has high reproductive potential
  • Gregarious
  • 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
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
  • Negatively impacts animal/plant collections
  • Damages animal/plant products
Impact mechanisms
  • Allelopathic
  • Competition - monopolizing resources
  • Competition - shading
  • Competition - smothering
  • Pest and disease transmission
  • Rapid growth
  • Produces spines, thorns or burrs
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult/costly to control

Uses

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S. angulatus is of minor importance as a garden ornamental, it is grown in particular as a screening plant (Anon., 2010). It has been used as a rootstock for cucumbers grown under glass in Europe, but seeds of S. angulatus are not produced commercially any more and the plant is no longer used as a rootstock in regular production (M Steeghs, personal communication, 2007 in Anon., 2010).

Uses List

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Environmental

  • Graft stock

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical

Detection and Inspection

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S. angulatus can be identified by its branched tendrils, leaves with 5 angular pointed lobes, clusters of pistillate flowers and spiny clusters of fruits.

Similarities to Other Species/Conditions

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S. angulatus is easy to identify by its spiny clusters of fruits, branched tendrils, distinctive leaf shape, and clusters of pistillate flowers (Anon., 2010). It can be confused with non-invasive plants such as Bryonia spp., Cucumis melo and Cucumis sativus. It could also be confused with E. lobata, which is also considered invasive in some countries. These species can be differentiated by their fruits; only S. angulatus' fruits are non-fleshy, spiny and gathered in glomerules (Anon., 2010).

Prevention and Control

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Prevention

SPS measures

S. angulatus was listed on the EPPO A2 List (recommended for regulation as quarantine pest) in 2006. In 2009, it was included on the priority list to be managed in EPPO member countries. In Japan, S. angulatus is regulated under the Invasive Alien Species Act (2006). In Korea, it is regulated under the Wildlife Protection Act. Also in its native range, it is listed as a noxious weed in Delaware, Indiana, and Kentucky (USDA-NRCS, 2009).

Eradication

The Generalitat of Catalonia made an order in 2005, declaring urgent measures for the eradication of S. angulatus (Diari Oficial de la Generalitat de Catalunya, 2009).

Control

Cultural control and sanitary measures

Tillage increased the number of emerged plants by 70-110% compared to the no-till system (Esbenshade et al., 2001a). Row spacing had no effect on S. angulatus emergence or biomass production (Esbenshade et al., 2001a). Messersmith et al. (2000) reported that in two of three field locations, late-season S. angulatus dry weight was approximately 65% less under no-tillage than in the moldboard plow treatment, whereas no difference in S. angulatus dry weight was observed between chisel plow and no-tillage treatments.

Chemical control

A pre-emergence application of atrazine, metribuzin plus chlorimuron, or linuron plus chlorimuron have been found to provide >90% visual control up to 8 weeks after treatment (Smeda and Weller, 2001). According to a field trial in the absence of competition from crops or other weeds, >80% visual control can be obtained with a post-emergence application of glyphosate or with combinations of glyphosate plus dicamba or glyphosate plus 2,4-D, chlorimuron, metribuzin plus chlorimuron, or paraquat (Smeda and Weller, 2001). Post-emergence applications of glufosinate in glufosinate-resistant maize, of chlorimuron plus thifensulfuron or nicosulfuron plus rimsulfruon plus atrazine in imidazolinone-resistant maize, and of chlorimuron plus thifensulfuron or glyphosate plus CGA-277476 (an experimental sulfonylurea herbicide) in glyphosate-resistant soybean provided good control of this species (Esbenshade et al., 2001b,c).

Post-emergence applications of CGA 152005 (proposed name, prosulfuron) and primisulfuron can both be effective for managing S. angulatus in maize (Messersmith et al., 1999).

Slife et al. (1962) found little movement of 14C-labelled 2,4-D out of the treated leaf of resistant S. angulatus after a small initial translocation, which was complete at the end of 24 hours. The apparent fixation of 2,4-D in S. angulatus led to the suggestion that S. angulatus may avoid injury from 2,4-D by immobilizing the molecule and thus preventing transport to the site(s) of metabolic activity (Dexter et al., 1971).

Gaps in Knowledge/Research Needs

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Information on natural enemies is lacking and may be important for biological control of S. angulatus. Also information on the genetic variation of this species in its native range is lacking, which may affect the estimation of its potential distribution ranges or the establishment of control measures.

References

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Links to Websites

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WebsiteURLComment
European and Mediterranean Plant Protection Organisation (EPPO)http://www.eppo.org/

Organizations

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France: EPPO European and Mediterranean Organization, OEPP/EPPO, 1 rue Le Nôtre, 75016 Paris, France, http://www.eppo.org

Contributors

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08/12/09 Original text by:

Shunji Kurokawa, Crop Production & Physiology Laboratory, Department of Forage Production, National Institute of Livestock &, Grassland Sci, National Agric Res Org., Nishinasuno, Tochigi 329-2793, Japan

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