Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Sida acuta
(sida)

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Datasheet

Sida acuta (sida)

Summary

  • Last modified
  • 16 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Host Plant
  • Preferred Scientific Name
  • Sida acuta
  • Preferred Common Name
  • sida
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • Originating in central America, the small perennial shrub, S. acuta has successfully invaded the tropics worldwide, largely as a contaminant in pasture seed. Its tolerance of a wide range of growing condition...

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Pictures

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PictureTitleCaptionCopyright
Leaves have toothed margins, are smooth or have sparse stellate hairs with prominent veins on the undersurface, 2-9 cm long, 0.5-4 cm wide. Flowers yellow, solitary, 1-2 cm in diameter, with five petals joined at the base and with a shallow notch at the apex. Fruit a hard, brown capsule, 3-5 mm in diameter, breaking into 5-8 triangular segments.
TitleFlower, fruit and leaf
CaptionLeaves have toothed margins, are smooth or have sparse stellate hairs with prominent veins on the undersurface, 2-9 cm long, 0.5-4 cm wide. Flowers yellow, solitary, 1-2 cm in diameter, with five petals joined at the base and with a shallow notch at the apex. Fruit a hard, brown capsule, 3-5 mm in diameter, breaking into 5-8 triangular segments.
Copyright©S.D. Sawant
Leaves have toothed margins, are smooth or have sparse stellate hairs with prominent veins on the undersurface, 2-9 cm long, 0.5-4 cm wide. Flowers yellow, solitary, 1-2 cm in diameter, with five petals joined at the base and with a shallow notch at the apex. Fruit a hard, brown capsule, 3-5 mm in diameter, breaking into 5-8 triangular segments.
Flower, fruit and leafLeaves have toothed margins, are smooth or have sparse stellate hairs with prominent veins on the undersurface, 2-9 cm long, 0.5-4 cm wide. Flowers yellow, solitary, 1-2 cm in diameter, with five petals joined at the base and with a shallow notch at the apex. Fruit a hard, brown capsule, 3-5 mm in diameter, breaking into 5-8 triangular segments.©S.D. Sawant
Flowering shoot.
TitleFlowering shoot
CaptionFlowering shoot.
Copyright©Chris Parker/Bristol, UK
Flowering shoot.
Flowering shootFlowering shoot.©Chris Parker/Bristol, UK
S. acuta is a vigorous competitor, suppressing other, more desirable plants.
TitleInfested area
CaptionS. acuta is a vigorous competitor, suppressing other, more desirable plants.
Copyright©S.D. Sawant
S. acuta is a vigorous competitor, suppressing other, more desirable plants.
Infested areaS. acuta is a vigorous competitor, suppressing other, more desirable plants.©S.D. Sawant

Identity

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Preferred Scientific Name

  • Sida acuta Burman f.

Preferred Common Name

  • sida

Other Scientific Names

  • Malvastrum carpinifolium (L.f.) A Gray
  • Malvinda carpinifolia (L.f.) Medik.
  • Sida acuta subsp. carpinifolia (L. f.) Borss. Waalk.
  • Sida acuta var. carpinifolia (L. f.) K. Schum.
  • Sida acuta var. intermedia S. Y. Hu
  • Sida acuta var. madagascariensis Hochr.
  • Sida berlandieri Turcz.
  • Sida bodinieri L.f.
  • Sida carpinifolia L. f.
  • Sida carpinifolia f. acuta (Burm. f.) Millsp.
  • Sida carpinifolia f. spiraeifolia (Link) Millsp.
  • Sida carpinifolia var. acuta (Burm. f.) Kurz
  • Sida chanetii Gand.
  • Sida frutescens Cav.
  • Sida garckeana Pol.
  • Sida jamaicensis Vell.
  • Sida lancea Gand.
  • Sida lanceolata Roxb.
  • Sida orientalis DC.
  • Sida planicaulis Cav.
  • Sida scoparia Lour.
  • Sida spiraeifolia Link
  • Sida spiraeifolia Willd.
  • Sida stauntoniana DC.
  • Sida ulmifolia Mill.
  • Sida vogelii Hook. f.

International Common Names

  • English: broom grass; broomweeds; cheeseweed; clock plant; common fanpetals; common wireweed; morning mallow; prickly sida; southern sida; spiny-head sida; wire weed
  • Spanish: babosilla; escoba blanca; escobita; escobita dulce; malva colorada; malva de Castilla; malva de platanillo
  • French: herbe à balais; herbe à panniers; herbe dure; herbe panier; sida à feuilles aiguës
  • Chinese: huang hua ren

Local Common Names

  • Australia: spinyhead sida
  • Bahamas: wire-weed
  • Brazil: guaxuma; relógio-de-vaqueiro; relógio-vassoura; vassonrinha curraleira; vassourinha-preta
  • Cambodia: kantrang bay sar
  • Colombia: escoba; escobilla
  • Cuba: malva bruja; malva de caballo
  • Dominican Republic: malva té
  • Ecuador/Galapagos Islands: escoba negra; escobilla negra
  • El Salvador: escoba; escobilla cabezuda; escobilla negra
  • Fiji: deni vuaka; paddy's lucerne
  • Germany: Samtmalve, Südliche
  • Haiti: balai cing heures; guimauve à petites fleurs; herbe connaît; petit lalo; ti-lalo
  • Honduras: huinar
  • Indonesia: galoenggang; sadagori
  • Lesser Antilles: balai; balai-onze-heures; balai-savane; balie; balye midi; balye onze; balye savann; soap bush; sweet broom
  • Malaysia: bunga telur belangkas; dukong anak; ketumbar hutan; lidah ular; pokok kelutut putih; sedeguri; seleguri; snake's tongue; spring-headed sida
  • Mexico: bobosilla; chichibe; escobilla blanca
  • Philippines: basbasot; escuba; surusighid; walis walisan
  • Samoa: mautofu
  • South Africa: taaiman
  • Sri Lanka: gas belila; kesar belila; malai tanki; palampasi; visha peti
  • Taiwan: syi ye jin wu shih hwa
  • Thailand: mai kward; mai kwat; yaa khat bai yaao; yung kwat; yung pat
  • Tonga: te'ehoosi
  • Trinidad and Tobago: ballier savanne
  • USA: southern sida
  • Venezuela: escoba amarilla; escoba dura
  • Vietnam: bai nhon

EPPO code

  • SIDAC (Sida acuta)

Summary of Invasiveness

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Originating in central America, the small perennial shrub, S. acuta has successfully invaded the tropics worldwide, largely as a contaminant in pasture seed. Its tolerance of a wide range of growing conditions has enabled S. acuta to become established in these diverse habitats. It infests various crops and habitats, but has been most problematic in pastures and rangelands, particularly in savannah-type biomes with pronounced wet and dry seasons. It can form dense monospecific stands in these regions, and has had a pronounced economic impact in northern Australia, Papua New Guinea and many Pacific Islands. Since the late 1980s, the foliage-feeding chrysomelid beetle Calligrapha pantherina has been introduced into many areas as a biological control agent specific to S. acuta and related Sida species. Introductions of C. pantherina have led to successes in control of S. acuta infestations, reducing seed production, and resulting in restoration of native vegetation in many cases.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Malvales
  •                         Family: Malvaceae
  •                             Genus: Sida
  •                                 Species: Sida acuta

Notes on Taxonomy and Nomenclature

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Numerous synonyms of S. acuta are cited by Borssum Waalkes (1966) and Fryxell (1985). The only one of these still to be used to some extent is S. carpinifolia, e.g. in Brazil (Lorenzi, 1982).

Description

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The following information is primarily from Holm et al. (1977), Waterhouse and Norris (1987) and Parsons and Cuthbertson (1992).

S. acuta is a small, erect, perennial shrub, branching profusely from the base. It usually ranges from 30-150 cm in height, but grows to 3 m in favourable conditions in northern Australia (Lonsdale et al., 1995). The stems are fibrous to almost woody, with a tough stringy bark. There is a deep, tough taproot. The leaves are alternate, lanceolate, acute, tapering towards both ends, and on a short, hairy petiole 3-6 mm long. The leaves have toothed margins, are smooth or have sparse stellate hairs and have prominent veins on the undersurface. The leaves are quite variable in size, from 2-9 cm long and 0.5-4 cm wide. The pair of stipules at the base of each leaf are not equal, with one frequently much narrower than the other. The flowers are yellow, solitary, 1-2 cm in diameter and on a short stalk 0.3-0.8 cm long. There are five petals, joined at the base and with a shallow notch at the apex. The fruit is a hard, brown capsule, 3-5 mm in diameter, breaking into 5-8 triangular segments. Each segment contains one seed and has a pair of sharp awns or 'beaks' 1-1.5 mm long which attach readily to animal fur or clothing. The seeds are small, reddish-brown to black, wedge-shaped, deeply indented on both sides, rounded on the back and about 1.5 mm long.

Distribution

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S. acuta is native to Central America, but has spread throughout the tropics and sub-tropics in the Pacific, Asia and Africa (Holm et al., 1977; Waterhouse and Norris, 1987; Parsons and Cuthbertson, 1992).

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

BhutanPresentUSDA-ARS, 2013
CambodiaWidespreadIntroducedHolm et al., 1977; Waterhouse, 1993
Chagos ArchipelagoPresentIntroduced Invasive PIER, 2013
ChinaPresentIntroduced Invasive Holm et al., 1977
-FujianPresentIntroducedFlora of China Editorial Committee, 2012
-GuangdongPresentIntroducedFlora of China Editorial Committee, 2012
-GuangxiPresentIntroducedFlora of China Editorial Committee, 2012
-HainanPresentIntroducedFlora of China Editorial Committee, 2012
-Hong KongPresentHolm et al., 1977
-YunnanPresentIntroducedFlora of China Editorial Committee, 2012
Christmas Island (Indian Ocean)WidespreadIntroduced Invasive Weeds of Australia, 2011Weed
Cocos IslandsWidespreadIntroduced Invasive Weeds of Australia, 2011Weed
IndiaWidespreadIntroduced Invasive Chaudhary, 1976
IndonesiaPresentIntroduced Invasive Holm et al., 1977; Waterhouse, 1993
-JavaPresentIntroduced Invasive ASEAN-Biodiversity, 2002
-Nusa TenggaraPresentIntroduced Invasive ASEAN-Biodiversity, 2002
IsraelPresentIntroducedUSDA-ARS, 2013
JapanPresentIntroduced Invasive Mito and Uesugi, 2004
JordanPresentIntroducedUSDA-ARS, 2013
LaosPresentIntroducedWaterhouse, 1993
MalaysiaWidespreadIntroduced Invasive Barnes and Chan, 1990; Waterhouse, 1993
MyanmarPresentIntroducedWaterhouse, 1993
NepalPresentIntroduced Invasive Sapkota, 2009
OmanPresentIntroducedUSDA-ARS, 2013
PhilippinesWidespreadIntroducedHolm et al., 1977; Waterhouse, 1993
SingaporePresentIntroduced Invasive Waterhouse, 1993
Sri LankaWidespreadIntroducedHolm et al., 1977
TaiwanWidespreadIntroduced Invasive Holm et al., 1977
ThailandWidespreadIntroducedWaterhouse, 1993; Noda et al., 1994
VietnamWidespreadIntroducedHolm et al., 1977; Waterhouse, 1993

Africa

BurundiPresentIntroducedUSDA-ARS, 2013
CameroonPresentIntroducedUSDA-ARS, 2013
CongoPresentIntroducedUSDA-ARS, 2013
Congo Democratic RepublicPresentIntroducedHolm et al., 1977
Côte d'IvoirePresentIntroducedHolm et al., 1977
EgyptPresentIntroducedUSDA-ARS, 2013
GabonPresentIntroducedUSDA-ARS, 2013
GhanaWidespreadIntroduced Invasive Holm et al., 1977; Niber, 1994
KenyaWidespreadIntroduced Invasive Holm et al., 1977
LiberiaPresentIntroducedUSDA-ARS, 2013
MadagascarPresentIntroducedUSDA-ARS, 2013
MalawiPresentIntroducedUSDA-ARS, 2013
MauritiusPresentIntroducedHolm et al., 1977
MayottePresentIntroduced Invasive PIER, 2013
MozambiquePresentIntroducedUSDA-ARS, 2013
NigeriaWidespreadIntroduced Invasive Holm et al., 1977
RwandaPresentIntroducedUSDA-ARS, 2013
SeychellesPresentIntroduced Invasive Hazell et al., 2008; PIER, 2009Agalega Island, Cousin Island, Frégate Island, Platte Island, Poivre Island
Sierra LeonePresentIntroducedUSDA-ARS, 2013
SomaliaPresentIntroducedUSDA-ARS, 2013
South AfricaPresentIntroduced Invasive USDA-ARS, 2013
Spain
-Canary IslandsPresentIntroduced Invasive DAISIE, 2013
SwazilandPresentIntroducedUSDA-ARS, 2013
TanzaniaPresentIntroducedUSDA-ARS, 2013
TogoPresentIntroducedUSDA-ARS, 2013
UgandaPresentIntroduced Invasive USDA-ARS, 2013
ZambiaPresentIntroducedUSDA-ARS, 2013

North America

MexicoPresentNative Not invasive Holm et al., 1977
USAPresentNativeHolm et al., 1977; USDA-ARS, 2009
-AlabamaPresentNativeUSDA-NRCS, 2013
-ArizonaPresentNativeUSDA-ARS, 2013
-FloridaPresentNativeUSDA-NRCS, 2013
-GeorgiaPresentNativeUSDA-ARS, 2013
-HawaiiWidespreadIntroduced Invasive Wagner et al., 1999Hawaii (Big Island), Kaui, Maui, Oahu, Molokai
-LouisianaPresentNativeUSDA-NRCS, 2013
-MississippiPresentNativeUSDA-NRCS, 2013
-New JerseyPresentNativeUSDA-NRCS, 2013
-PennsylvaniaPresentNativeUSDA-NRCS, 2013
-South CarolinaPresentNativeUSDA-ARS, 2013
-TexasPresentNativeUSDA-NRCS, 2013

Central America and Caribbean

AnguillaWidespreadNativeBroome et al., 2007
Antigua and BarbudaWidespreadNativeBroome et al., 2007
ArubaPresentNativeAcevedo-Rodríguez and Strong, 2012
BahamasPresentNativeAcevedo-Rodríguez and Strong, 2012
BarbadosWidespreadNativeBroome et al., 2007
BelizePresentNativeUSDA-ARS, 2013
British Virgin IslandsPresentNativeAcevedo-Rodríguez and Strong, 2012Guana, Tortola, Virgin Gorda
Costa RicaPresentNativeUSDA-ARS, 2013Weed
CubaPresentNative Invasive Holm et al., 1977; Oviedo Prieto et al., 2012Listed as one of the most noxious invasive species on this island
CuraçaoPresentNativeAcevedo-Rodríguez and Strong, 2012
DominicaWidespreadNativeBroome et al., 2007
Dominican RepublicPresentNative Not invasive Holm et al., 1977
El SalvadorPresentNative Not invasive Holm et al., 1977
GrenadaWidespreadNativeBroome et al., 2007
GuadeloupeWidespreadNativeBroome et al., 2007
GuatemalaPresentNativeUSDA-ARS, 2013
HaitiPresentNativeAcevedo-Rodríguez and Strong, 2012
HondurasPresentNative Not invasive Holm et al., 1979
JamaicaPresentNative Not invasive Callan, 1943; Holm et al., 1977
MartiniqueWidespreadNativeBroome et al., 2007
MontserratWidespreadNativeBroome et al., 2007
Netherlands AntillesWidespreadNativeBroome et al., 2007
NicaraguaPresentNativeUSDA-ARS, 2013
PanamaPresentNative Not invasive Holm et al., 1977
Puerto RicoPresentNative Invasive Acevedo-Rodriguez and Axelrod, 1999
Saint Kitts and NevisWidespreadNativeBroome et al., 2007
Saint LuciaWidespreadNativeBroome et al., 2007
Saint Vincent and the GrenadinesPresentNativeBroome et al., 2007
Trinidad and TobagoPresentNative Not invasive Holm et al., 1977
United States Virgin IslandsPresentNative Invasive Acevedo-Rodríguez and Strong, 2012St Croix, St Thomas, St John. Weed

South America

BrazilPresentNative Not invasive Holm et al., 1977
-BahiaPresentNativeForzza et al., 2012
-CearaPresentNativeForzza et al., 2012
-GoiasPresentNativeForzza et al., 2012
-MaranhaoPresentNativeForzza et al., 2012
-Minas GeraisPresentNativeForzza et al., 2012
-ParaPresentNativeForzza et al., 2012
-PernambucoPresentNativeForzza et al., 2012
-PiauiPresentNativeForzza et al., 2012
-TocantinsPresentNativeForzza et al., 2012
ColombiaPresentNative Not invasive Holm et al., 1977
EcuadorPresentNative Not invasive Porter, 1983
-Galapagos IslandsWidespreadIntroduced Invasive Charles Darwin Foundation, 2008; PIER, 2009Floreana Group, Isabela Group, San Cristobal Group, Santa Cruz Group, Santiago Group
French GuianaPresentNativeUSDA-ARS, 2013
GuyanaPresentNativeUSDA-ARS, 2013
PeruPresentNative Not invasive Holm et al., 1977
SurinamePresentNative Not invasive Holm et al., 1977
VenezuelaPresentNativeUSDA-ARS, 2013Weed

Europe

SpainPresentPresent based on regional distribution.

Oceania

American SamoaWidespreadIntroduced Invasive Waterhouse and Norris, 1987; Swarbrick, 1997; PIER, 2009Manu’a, Tutiila
AustraliaWidespreadIntroduced Invasive Waterhouse and Norris, 1987; Pitt, 1992; Fensham and Cowie, 1998; Smith, 2002; PIER, 2009Northern Territory, Queensland, Tiwi Islands
-Australian Northern TerritoryWidespreadIntroduced Invasive Weeds of Australia, 2011
-New South WalesPresentIntroduced Invasive Weeds of Australia, 2011
-QueenslandWidespreadIntroduced Invasive Weeds of Australia, 2011Weed
-Western AustraliaPresentIntroduced Invasive Weeds of Australia, 2011
Cook IslandsWidespreadIntroduced Invasive Waterhouse and Norris, 1987
FijiWidespreadIntroduced Invasive Holm et al., 1977; Waterhouse and Norris, 1987; PIER, 2009
French PolynesiaWidespreadIntroduced Invasive Waterhouse and Norris, 1987; PIER, 2009Gambier Islands, Marquesas, Society Islands, Tuamotu Archipelago, Tubuai (Austral) Islands
GuamWidespreadIntroduced Invasive Waterhouse and Norris, 1987
KiribatiPresentIntroduced Invasive Fosberg et al., 1979
Marshall IslandsPresentIntroduced Invasive Fosberg et al., 1979
Micronesia, Federated states ofWidespreadIntroduced Invasive Fosberg et al., 1979; PIER, 2009Chuuk Islands, Kosrae Island, Pohnpei Islands, Yap Islands
NauruPresentIntroduced Invasive Fosberg et al., 1979
New CaledoniaWidespreadIntroduced Invasive Waterhouse and Norris, 1987; PIER, 2009Île Art, Loyalty Islands, Ouvea Atoll, Île Tiga, New Caledonia Islands, Île Grande Terre
NiueWidespreadIntroduced Invasive Waterhouse and Norris, 1987; Whistler, 1988
Norfolk IslandPresentIntroduced Invasive Weeds of Australia, 2011Weed
Northern Mariana IslandsWidespreadIntroduced Invasive Fosberg et al., 1975; Fosberg et al., 1979
PalauPresentIntroduced Invasive PIER, 2009Angaur Island, Babeldaob Island
Papua New GuineaWidespreadIntroduced Invasive Waterhouse and Norris, 1987; Henty and Pritchard, 1988
SamoaWidespreadIntroduced Invasive Waterhouse and Norris, 1987; Whistler, 1988
Solomon IslandsWidespreadIntroduced Invasive Waterhouse and Norris, 1987
TongaWidespreadIntroduced Invasive Waterhouse and Norris, 1987; Whistler, 1988; PIER, 2009
VanuatuWidespreadIntroduced Invasive Waterhouse and Norris, 1987

History of Introduction and Spread

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Although native to central America, S. acuta has been spread through human agency throughout the tropics (Holm et al., 1977), including the major continental areas and a large portion of the oceanic islands. S. acuta is thought to have been introduced to northern Australia in the nineteenth century by Chinese prospectors who employed its fibres to make brooms (Flanagan et al., 2000). However, much of the history and introduction and spread of this fairly inconspicuous plant has not been documented. In many areas, such as the Galapagos Islands, the species has been naturalized long enough to become well integrated into the local ecosystems (Porter, 1983). In the West Indies, S. acuta is well integrated into the local floras to the point that is considered by many as native in the region.

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Australia China 1800s Yes Flanagan et al. (2000); Waterhouse and Norris (1987); Waterhouse and Norris (1987) Plant used in making brooms

Risk of Introduction

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With the broad distribution over which S. acuta is already established, it is likely to colonize many of the geographic areas in the subtropics and tropics where it has not already become naturalized.

Habitat

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S. acuta is found on most soil types, except seasonally flooded clays or soils derived from limestone (APB, 1993). It competes vigorously with other plant species, but does best in disturbed habitats in tropical or sub-tropical regions with a distinct wet and dry season. It has a deep taproot and can withstand drought, mowing and shallow tillage. It is a weed of degraded pastures, tree plantations, cereals, root crops, vegetables, planted forests, lawns, roadsides, and waste places (Pitt, 1992; Flanagan et al., 2000). In habitats where it occurs, it tends to flourish in riparian areas near watercourses. It has been reported at up to 1500 m altitude in Indonesia, at medium and higher elevations in Kenya and in the foothills of the Andes in Peru (Holm et al., 1977; Waterhouse and Norris, 1987; Parsons and Cuthbertson, 1992).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Secondary/tolerated habitat Harmful (pest or invasive)
Cultivated / agricultural land Secondary/tolerated habitat Natural
Managed grasslands (grazing systems) Principal habitat Harmful (pest or invasive)
Managed grasslands (grazing systems) Principal habitat Natural
Disturbed areas Present, no further details Natural
Rail / roadsides Present, no further details Natural
Urban / peri-urban areas Present, no further details Natural
Terrestrial ‑ Natural / Semi-naturalNatural grasslands Present, no further details Harmful (pest or invasive)
Natural grasslands Present, no further details Natural
Riverbanks Present, no further details Harmful (pest or invasive)
Riverbanks Present, no further details Natural
Scrub / shrublands Present, no further details Harmful (pest or invasive)
Scrub / shrublands Present, no further details Natural

Hosts/Species Affected

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S. acuta is a weed of plantation crops, cereals, root crops and vegetables throughout the Pacific and South-East Asia. It is a principal weed of maize in Mexico, sorghum in Australia and Thailand, tomatoes in the Philippines, onions in Brazil, and pastures in Australia, Fiji, Nigeria and Papua New Guinea. It is also a weed of tea in Taiwan and Sri Lanka, groundnuts in Ghana, cassava in Ghana and Nigeria, maize in Ghana, Nigeria and Thailand, coconuts in Trinidad, beans in Brazil, pastures under coconuts in Sri Lanka, pineapples in the Philippines, sugarcane and groundnuts in Australia, El Salvador and Trinidad, coffee in Colombia, rubber in Malaysia, upland rice in the Philippines and Nigeria, cotton in El Salvador and Thailand, and cowpeas and sweet potatoes in Nigeria (Holm et al., 1977; Chadhokar, 1978; Mott, 1980; Parsons and Cuthbertson, 1992; Ham and Eastick, 2004).

Biology and Ecology

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Reproductive Biology

Each flower produces 6 seeds, rarely 5 or 7 (Lonsdale et al., 1995). Seed production per plant averaged 101 and 236 at two locations in Australia with 95% of the seeds viable; seed production dropped to an average of 22 per plant with folivory due to the presence of the biological control insect Calligrapha pantherina (Lonsdale et al., 1995). Seeds are generally dormant when released requiring a post harvest ripening period at high temperatures for 1-3 months to break dormancy (Mott, 1980). Even when exposed on the soil surface for a full year in Australia, many seeds of S. acuta were viable, with 30% of seeds remaining viable after a dry season (Mott, 1980). Seeds of S. acuta have relatively high dormancy levels (Mott, 1980; McIvor and Howden, 2000). In a comparative study of 38 plant species in northern Australia, more S. acuta emerged under shaded conditions (8% out of 1000 seeds) than any other species (Gardener et al., 2001). In areas like northern Australia with distinct wet and dry seasons, most germination takes place early in the wet season, over a span of about 2 months (Mott, 1980).

The two awns on each seed segment attach readily to fur, hair or clothing. Seeds are also transported in mud on the feet of animals or vehicles. Seeds are also eaten by stock and pass undamaged through the gut. By these methods seeds become widely and rapidly dispersed. Coupled with a high natural germination rate and vigorous, competitive growth, S. acuta is a highly invasive and successful weed. 

Physiology and Phenology

Although innately a perennial, S. acuta can be annual in drier habitats, such as northern Australia (Flanagan et al., 2000). Plants senesce when the dry season begins in May or June, after germinating in November or December (Mott, 1980). Dry matter production is much greater under full sun conditions than shaded conditions, when S. acuta is vulnerable to competition (Chaudhary, 1976).

Seeds of S. acuta were found to contain phytoecdysteroids, which may provide protection for young seedlings against herbivores (Dinan et al., 2001). Phytochemical constituents have been shown to protect S. acuta from certain insects, such as larvae the noctuid moth, Earias vitella (Dongre and Rahalkar, 1993).

A savanna climate, featuring wet and dry periods, favours the phenology of S. acuta. In northern Australia, the seeds germinate during the first several months of the wet season that goes from November to April. The resulting plants produce flowers and set seed abundantly from March-May, with seeds able to survive well in dormancy through the dry season (Mott, 1980; Lonsdale et al., 1995).

Associations

S. acuta has been found associated with arbuscular mycorrhizae (Muthukumar et al., 2006).

Environmental Requirements

S. acuta is primarily a weed of the tropics and sub-tropics and is rarely found outside the zone between latitude 23°N and 23°S. It is a vigorous competitor, suppressing other, more desirable plants. The stems are unpalatable to stock (Nuwanyakpa et al., 1983), which selectively graze other plants, allowing S. acuta to dominate. Overgrazing accelerates this process. It survives mowing, slashing and cultivation. It tolerates a wide variety of soils and thrives in both dry and wet conditions.

Where there is no defined dry season, S. acuta will flower throughout the year. In monsoon climates, seeds germinate in the early part of the wet season, and most flowering takes place in the mid to late wet season. Each flower is open for one day only, opening in the morning and wilting by the late afternoon. Where there is a severe dry season, plants completely defoliate and then regrow from the rootstock with the initiation of the following wet season. Propagation is entirely by seeds which are produced in profusion and germinate readily following a post-harvest ripening period of 1-3 months at high temperatures (Mott, 1980). However, most seeds have a hard seedcoat which requires scarification, mechanically or by bacterial action, before the seeds will germinate (Parsons and Cuthberston, 1992). Consequently, about 30% of the seeds produced in Australia's Northern Territory during one wet season are still dormant at the beginning of the next.

Climate

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ClimateStatusDescriptionRemark
Am - Tropical monsoon climate Tolerated 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])
Cf - Warm temperate climate, wet all year Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Tolerated Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Acanthoscelides brevipes Herbivore Leaves
Asphondylia sidae Herbivore Stems
Brachycoryna pumila Herbivore Inflorescence/Leaves
Bucculatrix Herbivore Leaves
Calligrapha felina Herbivore Leaves
Calligrapha pantherina Herbivore Leaves Australia; Northern Territory; Queensland
Eutinobothrus sp. near pilosellus Herbivore Stems
Pyrgus adepta Herbivore Inflorescence/Leaves
Stegasta albocapitella Herbivore Leaves

Notes on Natural Enemies

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A biological control programme for Sida spp. in northern Australia began in 1984 with the establishment of an exploratory station in Mexico. A survey was made of the natural enemies of S. acuta in Mexico from April 1984 to November 1986 (Gillett et al., 1991), and in Australia's Northern Territory from 1984 to 1987 (Wilson and Flanagan, 1990).

The phytophagous insect fauna of S. acuta in the Northern Territory comprises 20 mainly rare, native or naturalized, polyphagous, external-feeding species.

Nine of the insects associated with S. acuta in Australia's Northern Territory are considered to be economic pests of agriculture (Wilson and Flanagan, 1990). These are the Rutherglen bug (Nysius vinitor), cottonseed bug (Oxycarenus luctuosus), hyaline grass bug (Liorhyssus hyalinus), cotton aphid (Aphis gossypii), hibiscus mealybug (Maconellicoccus hirsutus), cotton bollworm (Helicoverpa armigera), native budworm (Helicoverpa punctigera), cotton tipworm (Crocidosema plebejana), and a katydid (Caedicia sp.). In addition, the cotton stainer (Dysdercus andreae) has been recorded on S. acuta in Jamaica (Callan, 1943) and D. sidae and D. oceanicus in Fiji (Hinckley, 1963).

Additional natural enemies include Calonectria quinqueseptata (leaf spot of Hevea spp.) (Sulochana et al., 1982), Dysdercus ruficollis, Hypolimnas bolina, Meskea horor (Day et al., 1997) and Cuscuta spp. (parasitic dodders) in Sri Lanka (Jayasinge et al., 2004).

Means of Movement and Dispersal

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Seeds of S. acuta are easily spread by a variety of means including machinery, equipment, people, animals, seed or hay contamination and water (Pettit and Froend, 2001; Smith, 2002). Movement of S. acuta seed on tourist vehicles was documented for a national park in northern Australia (Lonsdale and Lane, 1994). Spread of S. acuta is commonly by seed, transported on vehicles or as contaminants in hay or seed (Smith, 2002). Seeds could also be transported directly by livestock either through clinging via sharp awns on the seeds or attaching to the hide of animals (Lonsdale et al., 1995).

Impact Summary

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CategoryImpact
Cultural/amenity Positive and negative
Economic/livelihood Positive and negative
Environment (generally) Positive and negative
Human health Positive

Economic Impact

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S. acuta is a vigorous competitor in degraded pastures, tree plantations, groundnuts, cereals, root crops, vegetables, planted forests, lawns, roadsides and waste places (Holm et al., 1977; Ham and Eastick, 2004), with the most serious infestations seen in pastures and rangeland (Holm et al., 1977; Schmutterer and Koch, 1979). Its lack of palatability help enable S. acuta to produce monospecific stands in pasture settings (Nuwanyakpa et al., 1983). S. acuta is considered amongst the ten most serious weeds in New Caledonia, Solomon Islands and Vanuatu (Waterhouse, 1985; Waterhouse and Norris, 1987), and is regarded by Holm et al. (1977) as among the 76 most serious weeds of the world. S. acuta was listed as one of 24 top invasive plant candidates for biological control among Pacific Island Countries and Territories, seven of which listed it as an important pest (Dovey et al., 2004). In Papua New Guinea, up to 80,000 ha in the Markham-Ramu valleys, were infested with thick stands of S. acuta up to 1.5 m in height following a drought year in 1997 (Kuniata and Korowi, 2004). Livestock owners were forced to cull numerous animals as a result. In the wake of the explosive increase in the S. acuta population, herbicide use was estimated to cost one company about $136,000 per year to control S. acuta on 37,000 ha of infested ranchland (Orapa, 2005). As a perennial weed S. acuta is most commonly recorded as a problem in perennial crops, where it has ample time for full development. Where weed succession advances to later seral stages, its perennial nature gives S. acuta and advantage (Afolayan, 1988). However, it establishes and flowers almost as rapidly as an annual and is able therefore to create problems in many crop situations.

Foliage of S. acuta has been observed to cause poisoning in cattle, with consumption of even a relatively small amount of foliage over an extended period of time leading to lisosomal storage disease (Furlan et al., 2008).

S. acuta also provides food, shelter and reproductive sites for insect pests of commercial crop plants. It is a major host of Anomis flava (cotton semi-looper), Maconellicoccus hirsutus (pink hibiscus mealybug), lantana mealy bug (Phenacoccus parvus), Okra mosaic virus (OMV), White-fly Transmitted Gemini (WTG) viruses and the recently characterized Sida yellow mosaic China virus (SiYMCNV) (Waterhouse, 1997; Umaharan et al., 1998; Xiong et al., 2005). It is also a wild host of Crocidosema plebejana (cotton tipworm), Phenacoccus manihoti (cassava mealybug) and the cotton insect, Dysdercus ruficollis. The pathogen Calonectria quinqueseptata (leaf spot of Hevea spp.) is also associated with S. acuta (Sulochana et al., 1982). The Sida yellow mosaic virus has been characterized on S. acuta and could potentially infect crops associated with the weed (Hernandez-Zepeda et al., 2007).

Environmental Impact

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Impact on Habitats

S. acuta may infest riparian areas, reducing habitat quality, as seen in Australia (Metcalfe, 2001). However, in some cases S. acuta may have a positive influence in reducing streambank erosion (Kumar et al., 1996). Simulated rainfall tests demonstrated the ability of transplanted seedlings to prevent loss of nitrogen in such settings (Kumar et al., 1997). 

Impact on Biodiversity

The formation of monospecific stands of this tough, spiny shrub in rangeland areas negatively impacts biodiversity (Lonsdale et al., 1995; Flanagan et al., 2000). S. acuta may also invade nearby natural areas; for example populations of S. acuta were identified infesting Australia’s Kakadu National Park, a world heritage site that is home to hundreds of species of native plants and animals (Cowie and Werner, 1993). Seeds of S. acuta were found being transported on the wheels of tourist vehicles within the same park (Londsdale and Lane, 1994). S. acuta is a larval host of the native butterfly, Hypolimnas bolina in Australia (Kemp, 1998). In its native range S. acuta provides forage for wildlife such as white-tailed deer, as well as cover for wildlife species like bobwhite quail, rabbits, and wild turkeys (Williams and Baxley, 2006).

Social Impact

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Where S. acuta has formed dense populations in pastures and rangelands, such as Papua New Guinea, northern Australia and many of the Pacific Islands, it has greatly impacted local economies, and demanded concerted control efforts. This has resulted as a strain on the already limited resources in these jurisdictions. The relatively low cost solution of introducing effective biological agents, such as Calligrapha pantherina has thus been much sought after in the south Pacific region (Flanagan et al., 2000; Dovey et al., 2004; Kuniata and Korowi, 2004; Julien et al., 2007).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Benefits from human association (i.e. it is a human commensal)
  • Fast growing
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
Impact outcomes
  • Damaged ecosystem services
  • Increases vulnerability to invasions
  • Modification of nutrient regime
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts agriculture
  • Negatively impacts animal health
  • Negatively impacts livelihoods
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - shading
  • Competition - smothering
  • Pest and disease transmission
  • Hybridization
  • Poisoning
  • 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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Cattle will graze S. acuta to a limited extent while the growth is still tender and it is utilized for livestock or poultry in some areas where it has been introduced (Egunjiobi, 1969; Serra et al., 1997), although it apparently exhibits some long-term toxicity to cattle (Furlan et al., 2008). Serra et al. (1997) observed that it had relatively high mineral content in pastures grazed by goats in the Philippines. Within its natural range it may provide valuable forage for wildlife such as white-tailed deer (Williams and Baxley, 2006).

It is regarded as a good source of fibre and a substitute for jute in Malaysia. It is used to make brooms in parts of South America and elsewhere, and is considered to have medicinal properties in parts of Asia and Africa (Waterhouse and Norris, 1987). Surveys among indigenous people identified S. acuta as an important source of traditional medicines in Africa, Asia and Central America (Karou et al., 2007). These uses include treating infections, fever, malaria, asthma, headache, diarrhea, skin diseases, dysentery, gonorrhea, rheumatism, eye cataracts and snakebite. It is seen as a valuable plant for treating nervous and urinary diseases and blood, bile and live disorders (Khare et al., 2002).

Its medical uses stem largely from relatively high levels of alkaloids and flavanoids, facilitating medicinal uses such as treatment of wounds or use as an antipyretic (Edeoga et al., 2005; Karou et al., 2007). Extracts of S. acuta exhibit antimicrobial activity on a variety of human infectious agents, including Staphylococcus aureus, Escherichia coli, Bacillus subtilis, Pseudomonas aeruginosa, and Mycobacterium phlei (Anani et al., 2000; Ekpo and Etim, 2009). Leaf extracts exhibit activity against bacteria but not against fungi, whereas root abstracts exhibit some activity against pathogenic fungi (Ekpo and Etim, 2009). Extracts from S. acuta exhibited strong antimalarial activity (Karou et al., 2003). Extracts from the root were found to possess strong hepatoprotective effects, thus validating traditional uses of S. acuta to treat liver disease (Sreedevi et al., 2009). Whole plant extracts were found to neutralize the effects of the venomous snake Bothrops atrox in Columbia (Otero et al., 2000).

S. acuta is able to accumulate heavy metals, and therefore may serve a phytoremediation role in contaminated sites (Gupta and Sinha, 2007). Extracts of S. acuta were also found offer protection of stored grains from the fungal pathogens Sitophilus oryzae and Prostephanus truncatus (Niber, 1994). Okunalo et al. (2007) used S. acuta as an indicator plant to detect contaminant levels in Kaduna, Nigeria.

Uses List

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Animal feed, fodder, forage

  • Fodder/animal feed

Environmental

  • Erosion control or dune stabilization
  • Wildlife habitat

Materials

  • Fibre
  • Pesticide

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Detection and Inspection

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Recommended resources for identification of S. acuta include PIER (2009), USDA-NRCS (2009), Viarouge et al. (1997), and Ivens (1968).

Similarities to Other Species/Conditions

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S. acuta can be readily confused with S. rhombifolia, and hybridization between the two species has been reported (Dekker, 1992). However, Viarouge et al. (1997) provide a valuable guide to identification of Sida species by vegetative characters, in which they point out that S. rhombifolia tends to have broader, more rhomboid leaves with dentation restricted to the upper half or two-thirds of the leaf margin, while the leaf margin in S. acuta is dentate throughout. The undersides of the leaves of S. rhombifolia are also more densely covered in stellate hairs. Other characters suggested by other authors include the stipules, which are uniformly narrow in S. acuta but of unequal width in S. rhombifolia; also the seeds of S. rhombifolia have only one awn; and the flowers are on stalks 1 to 3 cm long, much longer than those of S. acuta. On a local basis there may be confusion with some of the other Sida species which can occur as weeds; Viarouge et al. (1997) help to sort out nine species of western Africa. Ten species are described among weeds of Brazil (Lorenzi, 1982) and there is a useful table distinguishing the six that may occur in East Africa (Ivens, 1968). The Weeds of Australia (2011) factsheet for the species lists characters by which it can be distinguished from spiny sida (S. spinosa), common sida (S. rhombifolia), flannel weed (S. cordifolia), spiked sida (S. subspicata), spiked malvastrum (Malvastrum americanum) and prickly malvastrum (M. coromandelianum).

M. coromandelianum is more prostrate than S. acuta, has broader, irregularly serrated leaves with a suggestion of several uneven lobes, and has distinctive, circular, ribbed seed capsules. 

Prevention and Control

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Prevention

SPS measures

S. acuta is listed as a quarantine pest in a number of jurisdictions, particularly in the south Pacific region.

Public awareness

In countries where S. acuta has been of particular concern, strategies are being developed to better map and quantify the extent of the distribution of the plant, and thus raise awareness of its impact. As a weed of waste places and other marginal habitats, S. acuta often goes unnoticed until it invades pasture areas where it can cause economic damage.

Control

Cultural control and sanitary measures

Due to the tough, fibrous stems, deep taproot, unpalatability to livestock, continuous flowering and copious production of seeds with a degree of dormancy, S. acuta is very difficult to control. Single plants can be grubbed out prior to onset of flowering, ensuring the taproot is severed well below the crown to prevent regrowth (Parsons and Cuthbertson, 1992). Larger areas can be controlled by repeated cultivation until the supply of seeds in the soil is depleted. This is expensive and usually impractical. Satisfactory control is usually obtained by cultivating after the first rains of the monsoon have induced germination, followed by sowing of dense pastures (Waterhouse and Norris, 1987; Parsons and Cuthbertson, 1992). Strict control of grazing is required, as overgrazing encourages reinvasion of the weed. Competitive grasses may be effective in suppressing S. acuta, when combined with other control measures (Swarbrick, 1997), Certain pasture grasses may be grown to outcompete S. acuta, such as Urochloa mosambicensis (Cameron, 1996) or calopo (Calopogonium mucunoides) (Mott, 1980) in northern Australia.

Spread of S. acuta should be controlled by slashing or mowing before it flowers, and by keeping stock away from infested paddocks while it is fruiting. Seeds can be eaten by stock and pass undamaged through the gut, or may attach to the hide by means of the sharp awns on each seed. Seeds can be transported on vehicles or as contaminants in hay or seed (Smith, 2002).

Biological control 

A collaborative biological control programme for Sida spp. in northern Australia, involving the Northern Territory Department of Primary Industry and Fisheries and the Commonwealth Scientific and Industrial Research Organisation, began in 1984 with the establishment of an exploratory station in Mexico (Wilson and Flanagan, 1990; Hills, 1994). A survey was made of the natural enemies of S. acuta in Mexico from April 1984 to November 1986 (Gillett et al., 1991). Of the 31 phytophagous arthropods known to feed on S. acuta in its native range in Mexico, eight were considered to have potential as biological control agents.

Two species have been released in Australia. The chrysomelid beetle Calligrapha pantherina is causing extensive defoliation of S. acuta in some areas of northern Australia (Forno et al., 1992; Hills, 1993; Lonsdale et al., 1995; Day et al., 1997). Defoliation caused a reduction in seed production in northern Australia by an order of magnitude (8000 to 700 per sq m), but seed dormancy and the inability of beetles to locate isolated plants remain a challenge (Lonsdale et al., 1995). C. pantherina was released at 80 locations in northern Australia between 1989 and 1992, and had a significant impact on S. acuta within 10 years, especially near the coast (Flanagan et al., 2000). In addition to control of S. acuta populations, landowners have reported considerable return to native pasture.

C. pantherina is also capable of damaging S. rhombifolia, but is causing only limited damage to this plant in Australia due to climatic factors (Heard and Gardner, 1994). It is helpful to have S. rhombifolia as an alternate host (Forno et al., 1992). Sida spinosa is another congeneric introduced species in Australia that may act as an alternate host. Eutinobothrus pilosellus, not considered by Gillett et al. (1991) due to its uncertain taxonomy and specificity, is currently undergoing host specificity testing in Australia with a view to releasing it as a biological control agent of S. acuta.

Both C. pantherina and E. pilosellus are seen as good prospects for release in Pacific Island ecosystems invaded by S. acuta (Dovey et al., 2004; Julien et al., 2007). C. pantherina has provided good control since being released in Papua New Guinea in 2000 (Kuniata and Korowi, 2004), and has also been released in Fiji (2002) and on Vanuatu (2004) in the New Hebrides (Julien et al., 2007). Valuable experience has been gained over more than 10 years and in multiple locations (Hills, 1993; Flanagan et al., 2000; Julien et al., 2007) which has been helpful in deploying C. pantherina in new settings.

The other species considered by Gillett et al. (1991) to have potential were Acanthoscelides brevipes, Brachycoryna pumila, Calligrapha felina, Asphondylia sp. (possibly sidae), Stegasta albocapitella, Pyrgus adepta and a Bucculatrix species. The fungus Fusarium lateritium [Gibberella baccata], used for the control of S. spinosa (spiny sida) in the USA, has also been used against S. acuta with promising results (Boyette and Walker, 1983). The gall forming Lepidopteran Meskea horor was tested for host-specificity as a possible biological agent in Australia against Sida spp. However, M. horor was not specific enough as it could be reared on species representing all 4 tribes within the family Malvaceae (Day et al., 1997).

Physical/mechanical control

Tillage can be used to control S. acuta in pasture situations by slashing, chipping or cultivation (Swarbrick, 1997). Australian growers reported savings in mechanical control costs when the biological control agent Calligrapha pantherina was introduced (Flanagan et al., 2000). Sida species may also be controlled by electrical discharges (Brighenti and Brighenti, 2009).

Fertilizer treatments are sometimes recommended to enhance the efficacy of the biological control agent, C. pantherina when newly released; burning of grasslands may also impact the level of control (Kuniata and Korowi, 2004).

Chemical Control

Mature plants of S. acuta are difficult to control with herbicides. Young plants can be killed with amine 2,4-D or flowable atrazine (Reynolds, 1978; Pamplona and Moody, 1984). Mature plants are best treated by slashing early in the wet season, followed by spraying of the regrowth 2 weeks later with amine 2,4-D. Alternatively, if slashing is not possible, a spray of amine 2,4-D early in the wet season followed by another application later in the season may be effective (Parsons and Cuthbertson, 1992). Glyphosate, dicamba or picloram may be effective against mature plants (Waterhouse and Norris, 1987; Parsons and Cuthbertson, 1992), using ropewick or other type of directed application to avoid contact with the crop. Dicamba has also shown efficacy against S. acuta (Gonzalez-Ibanez, 1977). In northern Australia, metsulfuron methyl and fluroxypyr are used to control mature plants (Webb and Feez, 1987). Since the introduction of C. pantherina to northern Australia in the early 1990s, herbicide use has been reduced, with more than 40% of respondents attributing complete control to the agent by 1994 and 79% by 1999, resulting in substantial herbicide cost savings (Flanagan et al., 2000).

IPM

Fertilizer treatments are sometimes recommended to enhance the efficacy of the biological control agent, C. pantherina when newly released; burning of grasslands may also impact the level of control (Kuniata and Korowi, 2004).

Gaps in Knowledge/Research Needs

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There are many aspects of the biology, ecology and genetics of S. acuta that require further research. There is also an urgent need to better understand the dynamics of the relationship of S. acuta to the widely introduced biological control agent, Calligrapha pantherina under a variety of conditions and management parameters, as well as further research on other potential biological control agents. CLIMEX modelling techniques to predict further spread of S. acuta, including the context of climate change, would also be extremely useful.

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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.
USDA Germplasm Resources Information Networkhttp://www.ars-grin.gov/

Organizations

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Australia: ACIAR (Australian Centre for International Agricultural Research), GPO Box 1571, Canberra ACT 2601, Canberra, Australia, http://www.aciar.gov.au/

Australia: CSIRO (Commonwealth Scientific and Industrial Research Organisation), CSIRO Enquiries, Bag 10, Clayton South VIC 3169, http://www.csiro.au/

New Caledonia: Secretariat of the Pacific Community (SPC), BP D5, 98848, Noumea Cedex, 95 Promenade Roger, Laroque, Anse Vata, New-Caledonia, http://www.spc.org.nc/

Contributors

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10/07/13 Updated by:

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

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

18/12/2009 Updated by:

David Clements, Biology and Environmental Studies, Trinity Western University, 7600 Glover Road,  Langley, British Columbia, V2Y 1Y1, Canada

Distribution Maps

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