Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Physalis angulata
(cutleaf groundcherry)

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Datasheet

Physalis angulata (cutleaf groundcherry)

Summary

  • Last modified
  • 21 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Host Plant
  • Preferred Scientific Name
  • Physalis angulata
  • Preferred Common Name
  • cutleaf groundcherry
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • P. angulata is a herbaceous annual species of American origin which has been very widely introduced across many tropical, subtropical and warmer temperate regions. It is often characterized as a pantropical invasive weed of crops, gardens...

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Pictures

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PictureTitleCaptionCopyright
Physalis angulata (cutleaf groundcherry, husk tomato); habit at Mokuauia, Oahu. February, 2005.
TitleHabit
CaptionPhysalis angulata (cutleaf groundcherry, husk tomato); habit at Mokuauia, Oahu. February, 2005.
Copyright©Forest Starr & Kim Starr-2005 - CC BY-3.0
Physalis angulata (cutleaf groundcherry, husk tomato); habit at Mokuauia, Oahu. February, 2005.
HabitPhysalis angulata (cutleaf groundcherry, husk tomato); habit at Mokuauia, Oahu. February, 2005.©Forest Starr & Kim Starr-2005 - CC BY-3.0
Physalis angulata (cutleaf groundcherry, husk tomato); foliage and flowers. Georgia, USA. August 2009.
TitleFoliage and flowers
CaptionPhysalis angulata (cutleaf groundcherry, husk tomato); foliage and flowers. Georgia, USA. August 2009.
Copyright©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-3.0
Physalis angulata (cutleaf groundcherry, husk tomato); foliage and flowers. Georgia, USA. August 2009.
Foliage and flowersPhysalis angulata (cutleaf groundcherry, husk tomato); foliage and flowers. Georgia, USA. August 2009.©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-3.0
Physalis angulata (cutleaf groundcherry, husk tomato); flower and fruit. Mokuauia, Oahu. February, 2005.
TitleFlower and fruit
CaptionPhysalis angulata (cutleaf groundcherry, husk tomato); flower and fruit. Mokuauia, Oahu. February, 2005.
Copyright©Forest Starr & Kim Starr-2005 - CC BY-3.0
Physalis angulata (cutleaf groundcherry, husk tomato); flower and fruit. Mokuauia, Oahu. February, 2005.
Flower and fruitPhysalis angulata (cutleaf groundcherry, husk tomato); flower and fruit. Mokuauia, Oahu. February, 2005.©Forest Starr & Kim Starr-2005 - CC BY-3.0
Physalis angulata (cutleaf groundcherry, husk tomato); close-up of flower. Georgia, USA. August 2009.
TitleFlower
CaptionPhysalis angulata (cutleaf groundcherry, husk tomato); close-up of flower. Georgia, USA. August 2009.
Copyright©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-3.0
Physalis angulata (cutleaf groundcherry, husk tomato); close-up of flower. Georgia, USA. August 2009.
FlowerPhysalis angulata (cutleaf groundcherry, husk tomato); close-up of flower. Georgia, USA. August 2009.©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-3.0
Physalis angulata (cutleaf groundcherry, husk tomato); fruit capsule. Georgia, USA. August 2009.
TitleFruit capsule
CaptionPhysalis angulata (cutleaf groundcherry, husk tomato); fruit capsule. Georgia, USA. August 2009.
Copyright©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Physalis angulata (cutleaf groundcherry, husk tomato); fruit capsule. Georgia, USA. August 2009.
Fruit capsulePhysalis angulata (cutleaf groundcherry, husk tomato); fruit capsule. Georgia, USA. August 2009.©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Physalis angulata (cutleaf groundcherry, husk tomato); fruit. Georgia, USA. August 2009.
TitleFruit
CaptionPhysalis angulata (cutleaf groundcherry, husk tomato); fruit. Georgia, USA. August 2009.
Copyright©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Physalis angulata (cutleaf groundcherry, husk tomato); fruit. Georgia, USA. August 2009.
FruitPhysalis angulata (cutleaf groundcherry, husk tomato); fruit. Georgia, USA. August 2009.©Rebekah D. Wallace/University of Georgia/Bugwood.org - CC BY-NC 3.0 US
Physalis angulata (cutleaf groundcherry); habit, at high density in a maize crop (Zea mays). Vonitsa region, Greece. August, 2013.
TitleHabit
CaptionPhysalis angulata (cutleaf groundcherry); habit, at high density in a maize crop (Zea mays). Vonitsa region, Greece. August, 2013.
Copyright©Ilias S. Travlos
Physalis angulata (cutleaf groundcherry); habit, at high density in a maize crop (Zea mays). Vonitsa region, Greece. August, 2013.
HabitPhysalis angulata (cutleaf groundcherry); habit, at high density in a maize crop (Zea mays). Vonitsa region, Greece. August, 2013.©Ilias S. Travlos
Physalis angulata (cutleaf groundcherry); habit, at high density in a maize crop (Zea mays). Vonitsa region, Greece. August, 2013.
TitleHabit
CaptionPhysalis angulata (cutleaf groundcherry); habit, at high density in a maize crop (Zea mays). Vonitsa region, Greece. August, 2013.
Copyright©Ilias S. Travlos
Physalis angulata (cutleaf groundcherry); habit, at high density in a maize crop (Zea mays). Vonitsa region, Greece. August, 2013.
HabitPhysalis angulata (cutleaf groundcherry); habit, at high density in a maize crop (Zea mays). Vonitsa region, Greece. August, 2013.©Ilias S. Travlos
Physalis angulata (cutleaf groundcherry); habit, in a soybean crop (Glycine max). Vonitsa region, Greece. September, 2010.
TitleHabit
CaptionPhysalis angulata (cutleaf groundcherry); habit, in a soybean crop (Glycine max). Vonitsa region, Greece. September, 2010.
Copyright©Ilias S. Travlos-2010
Physalis angulata (cutleaf groundcherry); habit, in a soybean crop (Glycine max). Vonitsa region, Greece. September, 2010.
HabitPhysalis angulata (cutleaf groundcherry); habit, in a soybean crop (Glycine max). Vonitsa region, Greece. September, 2010.©Ilias S. Travlos-2010

Identity

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Preferred Scientific Name

  • Physalis angulata L.

Preferred Common Name

  • cutleaf groundcherry

Other Scientific Names

  • Physalis indica Lam.
  • Physalis lanceifolia Nees
  • Physalis minima L.
  • Physalis parviflora R. Br.

International Common Names

  • English: angular winter cherry; annual ground cherry; annual groundcherry; balloon cherry; bladder cherry; bladderberry; bush tomato; cape gooseberry; gooseberry; ground cherry; wild cape gooseberry; wild capegooseberry; wild gooseberry; winter cherry
  • Spanish: capuli cimarron; huevo de tortuga; tomatillo
  • French: coqueret
  • Portuguese: barrilheiro; bucho-de-ra

Local Common Names

  • Chinese: ku zhi
  • Argentina: alkekenje
  • Australia: annual groundcherry
  • Brazil: bucho de ra; bucho-de-ra'; camapu; camapu; ju'a de Capote; mata-fome
  • Chile: tomatillos de Brihuega
  • Cook Islands: tamaru; tupera tamaru-ariki; tupere
  • Cuba: huevo de gato
  • Dominican Republic: tope tope
  • Ecuador: popoja
  • Fiji: cevucevu; kospeli; mborosousou; mborosousou ni vavalangi; mbotembote yandra; mburasu; mokoai; thevuthevu
  • France: coqueret; coqueret anguleux
  • French Polynesia: tamanufairi; tamaru haari
  • Germany: Kantige Blasenkirsche
  • Guatemala: meltonate
  • Indonesia: cecendet; tjeplukan
  • Japan: sennari-houzuki; sennarihozuki; sennari-hozuki
  • Kiribati: te baraki; te bin
  • Marshall Islands: kaoror
  • Micronesia, Federated states of: tuhke wah pwonopwon duwehte tehn sele (Pohnpeian)
  • Nauru: oatamo; watamo
  • Nigeria: koropo
  • Niue: manini
  • Northern Mariana Islands: tomate chaca (Chamorro); tomates caputi (Chamorro)
  • Palau: bubeubedul; pohaa
  • Peru: bolsa mullaca
  • Philippines: putok putokan; tino tino
  • Puerto Rico: sababyche
  • Samoa: magalo; magalo; vi vao
  • Spain: bomba; huevillo; meltonate; mullaca; popoja; tomatillo; tomatillos de Brihuega; vejigon
  • Sweden: kantig lyktört
  • Tonga: polo pa
  • Tuvalu: pini; te peen
  • Venezuela: hueva de sapo; topotopo

EPPO code

  • PHYAN (Physalis angulata)

Summary of Invasiveness

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P. angulata is a herbaceous annual species of American origin which has been very widely introduced across many tropical, subtropical and warmer temperate regions. It is often characterized as a pantropical invasive weed of crops, gardens and plantations, although in many regions it has naturalized (such as Central America, Africa, India and Pacific islands) (Raju et al., 2007). It is a host of the causal agent of tomato bacterial spot Xanthomonas campestris pv. vesicatoria, as well as viruses found in tobacco, potato, okra, capsicum pepper, lucerne, beans and several other crops, physalis mottle virus (PhyMV), and also several root-knot nematodes (Meloidogyne spp.). It is used in the traditional treatment of a wide variety of disease and consequently can be an economically beneficial plant. No detailed analysis of its impacts has been conducted.

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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The identity of Physalis angulata L. as a distinct species is now well-established (Nicolson et alet al., 1988; Reddy et al., 1999), although it has been known by several synonyms. Two of the more commonly quoted are P. minima and P. lanceifolia, but neither is in current use.

Morphological variation in P. angulata was noted even by Linnaeus. P. lanceifolia and P. pendula were recorded by Kartesz (1994) as synonyms of P. angulata. Bean (2006) recognized two forms within P. angulata: a glabrous typical form and an antrorsely hairy form with unlobed or singly lobed leaves. The existence of numerous varieties of P. angulata suggests that there is considerable genetic variation within this species, something which has been confirmed by several genetic studies. This suggests there is potential for further selection.

The genus name Physalis, a Greek word, means bladder and refers to the inflated calyx, while the Latin species name angulata means angled and refers to the stems or the angled nature of calyx (Hall et al., 1991).

Description

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P. angulata is an annual herbaceous plant, branched erect, with tap-roots, angled and hollow stems growing up to 1-2 m in height, although there are reports of plants growing only up to 50 cm tall (South Australia, 2012). It is usually hairless (glabrous); however, occasional plants have short hairs, especially on the younger parts (Hall et al., 1991).

As stated by Bean (2006), in some Australian states (such as Queensland and New South Wales) the typical form of P. angulata is almost completely glabrous, and frequently the leaves have several lobes. Specimens from Top End in Northern Territory (Australia), Kimberley in Western Australia and Malesia also appear glabrous to the naked eye, but often have numerous antrorse hairs (up to 0.15 mm long) on the stems and pedicels. The leaves of this form are often entire or repand, or have at most a single pair of lobes or teeth and the calyx is moderately hairy (Bean, 2006).

The leaves are ovate to ovate-oblong leaves (or laceolate), obtuse, acute or rounded (somewhat asymmetric) at base, acute-acuminate, often repand-dentate, rather dark dull green, very short-puberulent on both sides on the nerves, 1-2 per node, usually irregularly toothed but sometimes smooth, 3-15 cm long and 2-10 cm wide. Leaf bases are unequal. The petioles are usually 3-4 cm long or sometimes longer, up to 10 cm (Stone, 1970; PIER, 2011; South Australia, 2012). A single plant can have up to 200 leaves, depending on the weed biotypes and several factors such as water and nutrient availability (Travlos, 2012).

The flowers are borne on stalks 5-40 mm in length. The flowering time is June to October in Greece and Turkey (Gönen et al., 2001; Travlos et al., 2010). The corolla is yellowish-green, usually without spots although occasionally with distinct spots, 4-12 mm long and 6-12 mm wide. The anthers are bluish or violet, up to 2.5 mm long and are borne on stalks up to 5 mm long (style 4-5 mm long). The green outer layer is 4-7 mm long with triangular lobes about as long as the tube. The fruit is an orange-coloured round berry, 1-1.8 cm long and is enclosed in the outer layer. This outer layer (a balloon-like calyx of 5 lobes with a small apical opening) is 2-6 mm long; it grows around and encloses the fruit. The fruiting calyx is pale green (pale brown when dry), inflated, 10-angled or ribbed, 23-35 mm long and 15-25 mm wide; it is borne on a stalk 1-4 cm long (Hall et al., 1991; South Australia, 2012). Seeds are disc-shaped to broadly reniform, 1-2 mm long, flat, pale yellow (Stone, 1970; Gönen et al., 2000; PIER, 2011).

Cotyledons are ovate with reddish petioles and without a distinct midvein. The first leaves are similar in shape to the cotyledons, but with an acute apex and evident venation.

Plant Type

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Annual
Broadleaved
Herbaceous
Seed propagated
Shrub

Distribution

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P. angulata is a widely-distributed weed of tropical, subtropical and warmer temperate regions. It is generally considered to be native to the Americas from the USA to Argentina, though its native range is not entire clear (USDA-ARS, 2013). It may also be native to Australia, although it is possible there was a pre–1788 introduction into Australia from the earliest European-Pacific-Central American contact. It is widespread in Asia and Africa and less widely distributed in Europe.

It is now considered to be a widespread weed of disturbed areas of Australia, where it is frequently found near rivers (South Australia, 2012). It is not certain that it has naturalized in all the countries it is recorded in (see distribution table), but has already done so in India, eastern Africa and some oceanic islands with warm climates. In Turkey it is one of the most commonly distributed weeds found in summer crops (Gönen et al., 2000); similarly in Greece, wherever found, it also grows in very high densities (Travlos et al., 2010).

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: 09 Jun 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AngolaPresent
BeninPresentIntroduced
BotswanaPresentIntroduced
Burkina FasoPresentIntroduced
Cabo VerdePresentIntroduced
CameroonPresentIntroduced
Central African RepublicPresentIntroduced
ChadPresentIntroduced
ComorosPresentIntroduced
Congo, Democratic Republic of thePresentIntroduced
Congo, Republic of thePresentIntroduced
Côte d'IvoirePresentIntroduced
EgyptPresentIntroduced
Equatorial GuineaPresentIntroduced
EswatiniPresentIntroduced
EthiopiaPresentIntroducedOriginal citation: Hedberg and Edwards (1989)
GabonPresentIntroduced
GambiaPresentIntroduced
GhanaPresentIntroduced
GuineaPresentIntroducedInvasivePapua New Guinea (eastern New Guinea Island)
KenyaPresentIntroduced
LiberiaPresentIntroduced
MadagascarPresentIntroduced
MaliPresentIntroduced
MauritaniaPresentIntroduced
MauritiusPresentIntroduced
MozambiquePresentIntroduced
NamibiaPresentIntroduced
NigerPresentIntroduced
NigeriaPresentIntroduced
RéunionPresentIntroduced
RwandaPresentIntroduced
SenegalPresentIntroduced
SeychellesPresentIntroduced
Sierra LeonePresentIntroduced
SomaliaPresentIntroduced
South AfricaPresentIntroducedEastern Cape, Free State, Gauteng, KwaZulu-Natal, Limpopo, Mpoumalanga, North West, Northern Cape
SudanPresentIntroduced
TanzaniaPresentIntroduced
-Zanzibar IslandPresentIntroduced
TogoPresentIntroduced
UgandaPresentIntroducedInvasive
ZambiaPresentIntroduced
ZimbabwePresentIntroduced

Asia

CambodiaPresentIntroduced
ChinaPresent
-AnhuiPresent
-FujianPresent
-GansuPresent
-GuangdongPresent
-HainanPresent
-HenanPresent
-HubeiPresent
-HunanPresent
-JiangxiPresent
-ZhejiangPresent
Hong KongPresent
IndiaPresent
-Andhra PradeshPresentWarangal District
IndonesiaPresentIntroduced
JapanPresentIntroducedInvasive
KuwaitPresentIntroducedAl-Abdali
LaosPresentIntroduced
MalaysiaPresentIntroduced
MyanmarPresentIntroduced
NepalPresentIntroduced
PakistanPresent
PhilippinesPresent
South KoreaPresentIntroducedInvasive
Sri LankaPresentIntroducedInvasive
TaiwanPresentIntroducedInvasive
ThailandPresentIntroducedInvasive
TurkeyPresentIntroducedSanliurfa, Cucurova region (Adana and Icel)
VietnamPresentIntroduced
YemenPresentIntroduced

Europe

FrancePresent
GreecePresent, Localized2012IntroducedInvasiveIt is mainly located in the western part of the country

North America

Antigua and BarbudaPresentNative
BahamasPresentNative
BarbadosPresentNative
BelizePresent
BermudaPresentNative
British Virgin IslandsPresent, WidespreadNative
Costa RicaPresent
CubaPresentNative
Dominican RepublicPresentNative
El SalvadorPresentNative
GrenadaPresentNative
GuadeloupePresentNative
GuatemalaPresentNative
HaitiPresentNative
HondurasPresentNative
JamaicaPresentNative
MartiniquePresentNative
MexicoPresentNativeBaja California, Campeche, Chiapas, Jalisco, Michoacan, Sinaloa, Sonora, Tabasco and Veracruz
NicaraguaPresentNative
PanamaPresentNative
Puerto RicoPresent, WidespreadNative
Saint Kitts and NevisPresentNative
Saint Vincent and the GrenadinesPresentNative
Trinidad and TobagoPresentNativeInvasive
U.S. Virgin IslandsPresentNativeSt Croix and St Thomas
United StatesPresentPresent based on regional distribution.
-AlabamaPresent, WidespreadNative
-ArizonaPresent, WidespreadNative
-ArkansasPresent, WidespreadNative
-CaliforniaPresent, WidespreadNative
-ConnecticutPresent, WidespreadNative
-DelawarePresent, WidespreadNative
-FloridaPresent, WidespreadNative
-GeorgiaPresent, WidespreadNative
-HawaiiPresentIntroducedInvasiveHawaii (Big), Kauai, Molokai and Oahu islands
-IllinoisPresent, WidespreadNative
-KansasPresent, WidespreadNative
-KentuckyPresent, WidespreadNative
-LouisianaPresent, WidespreadNative
-MainePresent, WidespreadNative
-MassachusettsPresentNative
-MississippiPresent, WidespreadNative
-MissouriPresent, WidespreadNative
-NevadaPresent, WidespreadNative
-New JerseyPresent, WidespreadNative
-New MexicoPresent, WidespreadNative
-North CarolinaPresent, WidespreadNative
-OklahomaPresent, WidespreadNative
-South CarolinaPresent, WidespreadNative
-TennesseePresent, WidespreadNative
-TexasPresent, WidespreadNative
-VirginiaPresent, WidespreadNative

Oceania

American SamoaPresentIntroducedInvasiveTa’u, Swain’s and Tutuila islands
AustraliaPresentPresent based on regional distribution.
-New South WalesPresentIntroducedInvasive
-QueenslandPresentIntroducedInvasive
Cook IslandsPresentSuwarrow (Taka) Atoll, 'Atiu, Mangaia and Ma'uke islands
Federated States of MicronesiaPresentIntroducedInvasiveInvasive in Fais, Kosrae and Pohnpei island. Also introduced in Chuuk islands, Pis, Weno, Ant Atoll, Faraulep, Ulithi Atoll, Yap island.
FijiPresentIntroducedInvasiveTaveuni, Vanua Levu and Viti Levu island
French PolynesiaPresentIntroducedInvasiveMarquesas, Society islands, Tuamotu Archipelago, Austral Islands
GuamPresentIntroducedInvasive
KiribatiPresentIntroducedInvasivePhoenix, Tungaru
Marshall IslandsPresentIntroducedInvasiveRalik Chain, Ratak Chain
NauruPresentIntroducedInvasiveNauru Island
New CaledoniaPresentIntroducedInvasiveIncluding Ile Grande Terre
New ZealandPresent
NiuePresentIntroducedInvasive
Northern Mariana IslandsPresentIntroducedInvasive on Anatahan Island. Also present on Farallon de Medinilla, Pagan, Rota, Saipan, Sarigan and Tinian islands.
PalauPresentIntroducedInvasiveAngaur, Babeldaob, Koror, Malakal, Ngerkebesang, Peleliou, Pulo Ana Island
Papua New GuineaPresentIntroducedInvasiveEastern New Guinea island
SamoaPresent
Solomon IslandsPresentIntroduced
TongaPresentIntroducedInvasiveTongatapu, Tofua, Eua and Vava'u Island
TuvaluPresentIntroducedInvasiveNui Atoll, Tuvalu Islands
VanuatuPresent
Wallis and FutunaPresentIntroducedInvasive

South America

ArgentinaPresentJujuy Province
BoliviaPresentSanta Cruz and Tarija
BrazilPresent, WidespreadNative
-Espirito SantoPresent
-MaranhaoPresent
-Sao PauloPresent
ColombiaPresentNative
EcuadorPresentNativeGalapagos Islands (Espanola, Floreana, Osborn, Campeon, Isabela), Guayas, Los Rios, Napo, Pastaza, Sucumbios
-Galapagos IslandsPresentNativeEspanola, Floreana, Pinta, Edeen, Santa Cruz, Seymour Norte, Santa Fe and Santiago islands; Oeste, Osborn, Campeon, Mosquera, Sombrero Chino, Punta Bowditch Norte and Gardner por Espanola islets; Volcan Sierra Negra
French GuianaPresent
GuyanaPresent
PeruPresentNativeAyacucho, Cusco, Huanuco, Junin, La Libertad, Lima, Loreta, Piura, San Martin
SurinamePresent
VenezuelaPresentNativeAmazonas, Bolivar, Delta Amacuro

History of Introduction and Spread

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The first reports of introduced P. angulata come from tropical regions of India during the 1700s (Rydberg, 1896; Stone, 1970). It was introduced to Australia before 1788 (Bean, 2006; South Australia, 2012), and apparently aboriginally introduced to Polynesia (Whistler, 1988). It is now widespread throughout tropical, subtropical and warmer temperate regions (Raju et al., 2007).

In 2001 there was only one record of P. angulata in Greece, in a rural site in Ilia prefecture, in the Peloponnese (Greuter and Rhaus, 2001), but by 2009 it was found in summer crops (maize, cotton and soyabean) and it is now considered to be a widely distributed invader in corn fields in Greece (Travlos et al., 2010; Travlos, 2012).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Australia USA Before 1788 Yes No Bean (2006) Collected by R. Brown in 1802
Greece Before 2001 Yes No Greuter and Raus (2001); Travlos et al. (2010) Specimens collected by Travlos
India   No No Rydberg (1896)
Turkey Before 1996 Yes No Gönen et al. (2000) Specimens collected by Gönen et al.

Risk of Introduction

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There are many countries where P. angulata has not yet been introduced, but the risk of further introductions remains high, given the cosmopolitan character of the species, its adaptability in a wide range of conditions and the lack of any wide-ranging prohibitions of its movement.

Habitat

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P. angulata is a plant of urban open spaces, disturbed areas, roadsides, croplands, orchards, nurseries and fallow land. It grows best in moist, fertile soils, prefers full sun conditions but it is also tolerant of partial shade. In general, across its wide distribution, P. angulata is considered a plant of tropical, sub-tropical and warm temperate conditions. It can occasionally be a hydrophyte but also occurs in uplands (USDA-NRCS, 2013). Whistler (1988) indicated the habitat of P. angulata to be croplands and disturbed places. It infests primarily annual crops but also occurs in orchards, nurseries, fallow land, roadsides and waste areas.

Throughout Florida to eastern Texas and northward to Pennsylvania, it occurs in fields, pastures, roadsides and open woodlands, preferring disturbed sites (Hall et al., 1991).

In Hawaii, Wagner et al. (1999) characterized it as ‘sparingly naturalized in disturbed sites’, and in Fiji it is ‘often locally abundant as a weed in villages, waste places, pastures, plantations, and canefields, along roads, on open slopes, and in forest along creeks from near sea level to about 600 m’ (Smith, 1991). In Niue, it is ‘a fairly common to very common weed of plantations in many areas, particularly the Basin’ (Sykes, 1970). In New Guinea, it occurs in gardens, waste land and cultivated fields, from low altitudes up to 1000 m (Henty and Pritchard, 1975). In Tonga it is ‘an occasional plantation weed’ (Yuncker, 1959; PIER, 2011). In India, P. angulata is an adventive weed, growing in nutrient-rich and garbage soils (Raju et al., 2007). In Australia, it is frequently found near rivers (South Australia, 2012). In Greece, it is mainly found amongst several summer crops (maize, cotton and soybean), especially in the western (more humid) part of the country (Travlos et al., 2010).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial ManagedCultivated / agricultural land Principal habitat
Terrestrial ManagedManaged forests, plantations and orchards Secondary/tolerated habitat
Terrestrial ManagedDisturbed areas Secondary/tolerated habitat
Terrestrial ManagedRail / roadsides Secondary/tolerated habitat
Terrestrial ManagedUrban / peri-urban areas Secondary/tolerated habitat
Terrestrial Natural / Semi-naturalNatural grasslands Secondary/tolerated habitat
Terrestrial Natural / Semi-naturalRiverbanks Secondary/tolerated habitat
Terrestrial Natural / Semi-naturalWetlands Secondary/tolerated habitat

Hosts/Species Affected

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P. angulata has been found growing in maize, cotton and soybean fields and is now considered to be a widely-distributed invader in corn fields in Greece (Travlos et al., 2010; Travlos, 2012).

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
Glycine max (soyabean)FabaceaeMain
Gossypium (cotton)MalvaceaeMain
Zea mays (maize)PoaceaeMain

Growth Stages

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Flowering stage, Vegetative growing stage

Biology and Ecology

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Genetics

Most sources suggest a basic chromosome number of n = 12, although Raju et al. (2007) and Mairura (2008) refer to it being a tetraploid species (2n = 48). Bean (2006) commented on the high genetic diversity in Australia, including two forms within P. angulata: a glabrous typical form with leaves with several lobes and an antrorsely hairy form with unlobed or singly lobed leaves. Worldwide, there are numerous varieties of P. angulata, which also suggests a high genetic diversity.

Reproductive Biology

P. angulata reproduces naturally by seed. In field conditions or under cultivation plants may yield up to about 200 fruit and more than 4000 seeds (Travlos et al., 2010; Travlos, 2012). Hall et al. (1991) reports that maximum germination occurred at 21°C with 10 hours of alternating temperatures and 30°C with 14 hours of alternating temperatures. Planting depth directly affected emergence, with a decrease from 89% to 0% germination with a corresponding increase of depth from 0 to 10 cm. Moreover, de Souza et al. (2013) found that osmotic seed priming improved seed germination and emergence of P. angulata.

Environmental Requirements

In general, across its wide distribution, P. angulata is considered a plant of tropical, sub-tropical and warm temperate conditions. P. angulata is hardy to USDA hardiness zones 9 to 11. It is a plant of frost-free areas; it is damaged by frost but may recover from short exposures down to -7 oC. It needs full sunlight, although it can tolerate partial shade. P. angulata grows best in moist, well-drained sandy loam soil.

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 Preferred > 430mm and < 860mm annual precipitation
BW - Desert climate Preferred < 430mm 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)
Ds - Continental climate with dry summer Preferred Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)
Dw - Continental climate with dry winter Preferred Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
40 35

Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • neutral

Soil texture

  • light
  • medium

Special soil tolerances

  • infertile

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Heliothis subflexa Plants|Inflorescence
Physalis mottle virus No

Notes on Natural Enemies

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The inflated calyx of P. angulata provides the moth Heliothis subflexa with a refuge from parasites (Sisterson and Gould, 1999). P. angulata is sensitive to many fungal diseases and is a host of the causal agent of tomato bacterial spot Xanthomonas campestris pv. vesicatoria. P. angulata hosts viruses found in tobacco, potato, okra, capsicum pepper, beans and several other crops, as well as physalis mottle virus (PhyMV), and several root-knot nematodes (Meloidogyne spp.) (Mairura, 2008).

Means of Movement and Dispersal

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Natural Dispersal and Vector Transmission (Biotic)

Natural dispersal of P. angulata by water is likely (Travlos et al., 2010). There is presumed to be dispersal by birds and mammals eating the fruit, but no documentation exists (Travlos, personal observations).

Accidental Introduction

The seeds of P. angulata can be carried short distances on people’s clothing or footwear (Travlos et al., 2010).

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessionsVery often with their seeds Yes Travlos et al. (2010)
Water Yes

Impact Summary

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CategoryImpact
Cultural/amenity Negative

Economic Impact

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P. angulata has rarely been reported as an invasive weed of crops and data on crop losses caused by it are only common at the local level (e.g. Travlos et al., 2010). Conversely, it has been, and continues to be, an economically useful plant, mainly because of its use in traditional treatment of malaria, asthma, hepatitis, dermatitis, rheumatism, fever and many other diseases (Chiang et al., 1992; Lin et al., 1992; Caceres et al., 1995; Freiburghaus et al., 1996; Ankrah et al., 2003; Choi and Hwang, 2003). Moreover, P.angulata extracts act against human cancer cells (Lee et al., 2009), roots have shown anti-inflammatory and immunomodulatory properties (Bastos et al., 2008) and the stems have demonstrated anti-neoplastic activity (Ribeiro et al., 2002).

In larger quantities the plant is poisonous to cattle and sheep, and it gives the meat a musk-like smell (Mairura, 2008).

Environmental Impact

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P. angulata is a weed of cropland, gardens and plantations. It is a host of the causal agent of tomato bacterial spot Xanthomonas campestris pv. Vesicatoria, as well as viruses found in tobacco, potato, okra, capsicum pepper, lucerne, beans and several other crops, physalis mottle virus (PhyMV), and also several root-knot nematodes (Meloidogyne spp.) (Mairura, 2008).

Although P. angulata is reported as naturalized and invasive in a number of countries, especially in India and the Pacific region, no detailed analysis of its impacts has been conducted.

Social Impact

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In tropical Africa the fruit is eaten as a snack, but eating too much of the fruit may cause dizziness. The leaves are eaten in salads, although they taste bitter. In larger quantities the plant is poisonous to cattle and sheep, and it gives the meat a musk-like smell (Mairura, 2008).

Garcia et al. (2006) have shown that that physalin B from P. angulata is a potent immunomodulatory substance for the bloodsucking insect Rhodnius prolixus and its parasitism by Trypanosoma rangeli, a protozoan parasite of triatomines and mammals. Moreover, physalin B extracted by P. angulata seems an excellent compound to reduce parasite transmission and control Chagas disease by reducing parasitism of R. prolixus with T. cruzi (Castro et al., 2012). It is already used in several countries by local people against asthma, bacterial infections, skin diseases, fever, inflammation, kidney and liver problems, malaria and cancer (Bastos et al., 2006; Hsieh et al., 2006; Lee et al., 2009; Hseu et al., 2011; Hsu et al., 2012). P.angulata extracts act against human cancer cells (Lee et al., 2009), roots have shown anti-inflammatory and immunomodulatory properties (Bastos et al., 2008) and the stems have demonstrated anti-neoplastic activity (Ribeiro et al., 2002).

Risk and Impact Factors

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Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Tolerant of shade
  • Fast growing
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Has high genetic variability
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Negatively impacts agriculture
  • Negatively impacts animal health
  • Reduced native biodiversity
  • Threat to/ loss of native species
  • Negatively impacts animal/plant collections
Impact mechanisms
  • Competition - monopolizing resources
  • Poisoning
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally

Uses

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The many-seeded fruit is edible when ripe (Hall et al., 1991). Its berries are often used as food by Aborigines in several regions of Australia (South Australia, 2012). In tropical Africa the fruit is eaten as a snack, but eating too much of the fruit may cause dizziness. The leaves are eaten in salads, although they taste bitter.

Extracts or infusions from this plant have been used in various countries in popular medicine as a treatment for a variety of illnesses, such as malaria, asthma, hepatitis, dermatitis and rheumatism (Chiang et al., 1992; Lin et al., 1992; Santos et al., 2003; Soares et al., 2003; Mairura, 2008). In Brazil, the juice of P. angulata is considered to be a sedative and to relieve rheumatism and earache. The leaves are sometimes used against inflammations of the bladder, spleen and liver. The whole plant cooked is recommended in baths for inflammatory processes, such as rheumatism (Lorenzi, 1982; Bastos et al., 2006).

P.angulata extracts act against human cancer cells (Lee et al., 2009), its roots have shown anti-inflammatory and immunomodulatory properties (Bastos et al., 2008) and the stems have demonstrated anti-neoplastic activity (Ribeiro et al., 2002).

It has been demonstrated that some of the extracts or active componants obtained from P. angulata have a broad range of biological activities, including antibacterial, molluscicidal, antiprotozoal and cytotoxic activities (Kastelein and Camargo, 1990; Lee et al., 1991; Chiang et al., 1992; Lin et al., 1992; Cáceres et al., 1995; Freiburghaus et al., 1996; Pietro et al., 2000; Ismail and Alam, 2001; Januario et al., 2002; Santos et al., 2003; Soares et al., 2003; Lee et al., 2009).

Phytochemical studies of P. angulata have demonstrated the presence of alkaloids, steroids known as physalins (D, I, G, K, B, F, E), physagulins (E, F and G), anolides and flavonoids (Row et al., 1978, 1980; Lee et al., 1991; Chiang et al., 1992; Shingu et al., 1992; Ismail and Alam, 2001; Damu et al., 2007).

Garcia et al. (2006) have shown that that physalin B from P. angulata is a potent immunomodulatory substance for the bloodsucking insect Rhodnius prolixus and its parasitism by Trypanosoma rangeli, a protozoan parasite of triatomines and mammals and the etiological agent of Chagas disease. Moreover, physalin B extracted by P. angulata seems an excellent compound to reduce parasite transmission and control Chagas disease by reducing parasitism of R. prolixus with T. cruzi (Castro et al., 2012). It is already used in several countries by local people against bacterial infections, skin diseases, fever and kidney and liver problems (Bastos et al., 2006; Hsieh et al., 2006; Lee et al., 2009; Hseu et al., 2011; Hsu et al., 2012).

Uses List

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Environmental

  • Host of pest

Human food and beverage

  • Fruits
  • Vegetable

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Similarities to Other Species/Conditions

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P. angulata and P. minima are distinguished from its highly invasive relative P. peruviana (the cape gooseberry) as they are annual, not perennial, and have anthers less than 3 mm long (Flora of China, 2012). The corollas of P. angulata, P. pubescens and P. minima are less than 1 cm diameter, whereas the corolla of P. peruviana is more than 1.3 cm in diameter (Ganapathji et al., 1991). P. viscosa (sticky Cape gooseberry), P. alkekengi (Chinese lantern) and P. virginiana (= P. longifolia) can all be distinguished by their fully rhizomatous rootstock (South Australia, 2012). P. angulata may be also confused with P. ixocarpa Brot. ex Hornem, but can be distinguished by its longer pedicels, creamy-yellow corolla and angular fruiting calyx (South Australia, 2012). P. philadelphia has anthers 3-4 mm long but is annual, has bright yellow flowers, fruits up to 3 cm diameter and is only sparsely pubescent.

In India, there are some distinguishing morphological characters of the tetraploid taxon P. angulata and the diploid taxon P. lagascae: P. angulata plants are erect, robust, smooth, with bigger flowers (>7 mm across), bluish anthers and purple fruiting calyces, whereas P. lagascae plants are diffuse to erect, relatively smaller, with smaller flowers <6 mm across), yellow anthers and green fruiting calyces (Raju et al., 2007). Raju et al. (2007) reported that the tetraploid P. minima so reported by Ganapathi et al. (1991) and Sudhakaran and Ganapathi (2001), and earlier by Bhaduri (Menzel, 1951) and others are likely to be variants of P. angulata.

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.

Physical/Mechanical Control

P. angulata can be readily uprooted or hoed out when young but requires deeper digging once established.

Biological Control

No attempts at biological control have been reported.

Chemical Control

Glyphosate is considered as a suitable herbicide for the control of P. angulata (at young growth stages), but it is likely to damage surrounding crops or vegetation (Travlos et al., 2010). P. angulata (like other Physalis species) is resistant to dinitroalanine herbicides and 2,4-D. However, mesotrione, sulcotrione and nicosulfuron are used and registered in some countries for the control of this plant.

Gaps in Knowledge/Research Needs

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There is a particular need for information on the susceptibility of P. angulata to herbicides and the degree of competition between P. angulata and other weeds and crops. Additionally, the beneficial effects of this plant need to be further studied, in order to reveal any potential interest for further use and cultivation. More information on the impacts of P. angulata are also needed.

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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.
Plantwise Knowledge Bankhttp://www.plantwise.org/KnowledgeBank/Datasheet.aspx?dsid=40711
PROTAhttp://www.prota4u.org/search.asp
South Australiahttp://www.flora.sa.gov.au/efsa/lucid/Solanaceae/Solanaceae%20species/key/Australian%20Solanaceae%20species/Media/Html/Physalis_
USDA-NRCAhttp://plants.usda.gov/core/profile?symbol=phan5

Contributors

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09/09/13 Original text by:

Ilias Travlos, Agricultural University of Athens, Greece

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