Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Portulaca quadrifida
(chickenweed)

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Datasheet

Portulaca quadrifida (chickenweed)

Summary

  • Last modified
  • 19 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Portulaca quadrifida
  • Preferred Common Name
  • chickenweed
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • P. quadrifida is a succulent, annual, mat-forming species. The native range of this species is not clearly defined and many describe it as pantropical. P. quadrifida has been introduced into a number o...

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Pictures

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PictureTitleCaptionCopyright
Portulaca quadrifida (chickenweed); habit, close view, showing stems, leaves and flowers. Bergianska Trädgården, Stockholm, Sweden. July, 2012.
TitleHabit
CaptionPortulaca quadrifida (chickenweed); habit, close view, showing stems, leaves and flowers. Bergianska Trädgården, Stockholm, Sweden. July, 2012.
Copyright©Christer T. Johansson-2012/via wikipedia - CC BY-SA 3.0
Portulaca quadrifida (chickenweed); habit, close view, showing stems, leaves and flowers. Bergianska Trädgården, Stockholm, Sweden. July, 2012.
HabitPortulaca quadrifida (chickenweed); habit, close view, showing stems, leaves and flowers. Bergianska Trädgården, Stockholm, Sweden. July, 2012.©Christer T. Johansson-2012/via wikipedia - CC BY-SA 3.0
Portulaca quadrifida (chickenweed); habit. Bergianska Trädgården, Stockholm, Sweden. July, 2012.
TitleHabit
CaptionPortulaca quadrifida (chickenweed); habit. Bergianska Trädgården, Stockholm, Sweden. July, 2012.
Copyright©Christer T. Johansson-2012/via wikipedia - CC BY-SA 3.0
Portulaca quadrifida (chickenweed); habit. Bergianska Trädgården, Stockholm, Sweden. July, 2012.
HabitPortulaca quadrifida (chickenweed); habit. Bergianska Trädgården, Stockholm, Sweden. July, 2012.©Christer T. Johansson-2012/via wikipedia - CC BY-SA 3.0

Identity

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Preferred Scientific Name

  • Portulaca quadrifida L.

Preferred Common Name

  • chickenweed

Other Scientific Names

  • Illecebrum verticillatum L.
  • Portulaca formosana (Hayatta) Hayatta
  • Portulaca meridiana L.f.
  • Portulaca microphylla A.Rich.
  • Portulaca walteriana Poelln.

International Common Names

  • English: single-flowered purslane; small-leaved purslane; ten o’clock plant
  • French: pourpier
  • Chinese: si ban ma chi xian

Local Common Names

  • : kinyorwe
  • Cuba: espinaca; verdolaga
  • Dominican Republic: pinganilla; regina; ringa
  • India: chaunlayi; chotaluniya; godupavili kura; pacin; passalai keerai; pavin; sinna paruppu keerai
  • Lesser Antilles: pourpier double
  • Myanmar: mya-byit-gale; nya-byit
  • Pakistan: lunak
  • Philippines: sayikan
  • Portugal: beldroega
  • Puerto Rico: yerba de pollo
  • Thailand: phak bia nu

Summary of Invasiveness

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P. quadrifida is a succulent, annual, mat-forming species. The native range of this species is not clearly defined and many describe it as pantropical. P. quadrifida has been introduced into a number of new countries around the world. There are no records of P. quadrifida behaving as an invasive weed outside its native range, where it acts as an agricultural weed. This species is not particularly competitive but, as a halophyte, it is resistant to drought and areas of low soil fertility and therefore readily becomes dominant under such circumstances. There is also a large amount of variation within this species enabling it to adapt to new environments. As a plant with food and herbal properties, there are possibilities for deliberate introduction of this species into new areas where it could be problematic.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Caryophyllales
  •                         Family: Portulacaceae
  •                             Genus: Portulaca
  •                                 Species: Portulaca quadrifida

Notes on Taxonomy and Nomenclature

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The genus Portulaca contains around 150 species (PROTA, 2014). Portulaca is in the family Portulacaceae, which has traditionally been treated to contain several genera, but very recently circumscribed to contain only the type genus (Nyffeler and Eggli, 2010). P. quadrifida has been known by a number of other synonyms but the original Linnean name persists and there is no confusion with any closely related species. It is however regarded as a variable species and occurs in a number of different ploidy forms (2n= 18, 36, 48).

Description

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The following description of P. quadrifida is taken from Flora Zambesiaca (2014).

Prostrate annual herb with a somewhat swollen tap-root; stems often reddish and sometimes rooting at the nodes, sometimes reaching 25 cm long and c. 1 cm in diam. Leaves opposite; lamina fleshy, up to 10 × 4 mm, but often much less and frequently somewhat shrunk in dried specimens, elliptic, lanceolate, elliptic-oblong or rarely cordate-ovate, apex acute or obtuse, both surfaces more or less flattened; petiole c. 1 mm. long; stipular hairs whitish, numerous, 3–5 mm long. Flowers 1–4 at the ends of the branches, surrounded by 4 leaves often somewhat larger than the cauline ones and by numerous hairs c. 4 mm long. Sepals 2–4 × 1·5–3·5 mm ± triangular, obtuse, united at the base. Petals 4 (5), yellow or orange or very rarely pink or purplish, almost free, c. 5 × 3 mm, elliptic to ovate. Stamens 8–12. Ovary conical-ovoid; style c. 5 mm long, thickened at the apex and with 4 spreading stigmas. Capsule conical-ovoid, dehiscing horizontally c. 1/3 of the way up. Seeds many, c. 1 mm in diameter, greyish, reniform, verrucose with blunt tubercles.

The stamen number and flower size of P. quadrifida however may be variable (PROTA, 2014).

Plant Type

Top of page Annual
Broadleaved
Herbaceous
Seed propagated
Succulent
Vegetatively propagated

Distribution

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The distribution of P. quadrifida is not clearly defined. USDA-ARS (2014) describe the distribution of P. quadrifida as ‘pantropical, except Australia’ but the Flora of China Editorial Committee (2014) considers it as ‘probably native to Africa, now pantropical’. Wagner and Lorence (2015) in the Flora of West Indies also treat this species as native to much of the Caribbean.

For the purposes of this datasheet, P. quadrifida is treated as native to Africa and Western Asia and introduced elsewhere. See distribution table for countries in which this species is recorded.

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

ChinaPresentIntroducedFlora of China Editorial Committee, 2015; GBIF, 2015
-GuangdongPresentIntroducedFlora of China Editorial Committee, 2015
-HainanPresentIntroducedFlora of China Editorial Committee, 2015
-YunnanPresentIntroducedFlora of China Editorial Committee, 2015
IndiaPresentNativeGBIF, 2015
-GujaratPresentNativeSrivastava et al., 2013
-Indian PunjabPresentNativeSaimbhi et al., 1992
-KarnatakaPresentNativeBalikai, 1999
-Tamil NaduPresentNativeSubbulakshmi et al., 2009
-Uttar PradeshPresentNativeKachare et al., 2005
IndonesiaPresentNativeGBIF, 2015First specimen in 1923
KuwaitPresentNativeGBIF, 2015
MaldivesPresentGBIF, 2015
NepalPresentNativeSiwakoti, 2000
OmanPresentNativeGBIF, 2015
PakistanPresentNativeGBIF, 2015
Saudi ArabiaPresentNativeGBIF, 2015
SingaporePresentGBIF, 2015
Sri LankaPresentNativeGBIF, 2015
TaiwanPresentFlora of China Editorial Committee, 2015First specimen from 1920
ThailandPresentGBIF, 2015
United Arab EmiratesPresentNativeKarim, 1993
YemenPresentNativeGBIF, 2015

Africa

AngolaPresentNativeGBIF, 2015
BeninPresentNativeGBIF, 2015
BotswanaPresentNativeGBIF, 2015
Burkina FasoPresentNativeGBIF, 2015
CameroonPresentNativeGBIF, 2015
Central African RepublicPresentNativeGBIF, 2015
ComorosPresentNativeGBIF, 2015
Congo Democratic RepublicPresentNativePROTA, 2014Cultivated
Côte d'IvoirePresentNativeGBIF, 2015
EgyptPresentNativeFlora of Pakistan, 2015
EthiopiaPresentNativeGBIF, 2015
GhanaPresentNativeGBIF, 2015
Guinea-BissauPresentNativeGBIF, 2015
KenyaPresentNativeGBIF, 2015
LiberiaPresentNativeGBIF, 2015
MadagascarPresentNativeGBIF, 2015
MaliPresentNativeGBIF, 2015
MayottePresentNativeGBIF, 2015
MozambiquePresentNativeGBIF, 2015
NamibiaPresentNativeGBIF, 2015
NigeriaPresentNativeGBIF, 2015
RwandaPresentNativePROTA, 2014Cutlivated
SeychellesPresentNativeGBIF, 2015
SomaliaPresentNativeGBIF, 2015
South AfricaPresentNativeGBIF, 2015
SudanPresentNativeGBIF, 2015
SwazilandPresentNativeGBIF, 2015
TanzaniaPresentNativeGBIF, 2015
TogoPresentNativeGBIF, 2015
UgandaPresentNativeGBIF, 2015

Central America and Caribbean

Antigua and BarbudaPresentWagner and Lorence, 2015
BarbadosPresentWagner and Lorence, 2015
British Virgin IslandsPresentGBIF, 2015
CaribbeanPresentWagner and Lorence, 2015
CubaPresentWagner and Lorence, 2015
Dominican RepublicPresentWagner and Lorence, 2015
Greater AntillesPresentWagner and Lorence, 2015
GrenadaPresentWagner and Lorence, 2015
GuadeloupePresentGBIF, 2015
HaitiPresentWagner and Lorence, 2015
JamaicaPresentGBIF, 2015; Wagner and Lorence, 2015
Lesser AntillesPresentWagner and Lorence, 2015
MontserratPresentWagner and Lorence, 2015
Netherlands AntillesPresentGBIF, 2015
Puerto RicoPresentGBIF, 2015; Wagner and Lorence, 2015
Saint Kitts and NevisPresentWagner and Lorence, 2015
Saint LuciaPresentWagner and Lorence, 2015
United States Virgin IslandsPresentGBIF, 2015First specimen in 1895

South America

GuyanaPresentGBIF, 2015

Oceania

FijiPresentGeesink, 1969
KiribatiPresentGeesink, 1969
Marshall IslandsPresentGBIF, 2015
SamoaPresentGeesink, 1969; GBIF, 2015
Solomon IslandsPresentGBIF, 2015

History of Introduction and Spread

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There is little information with regards to the introduction and spread of P. quadrifida however, GBIF (2014) records specimens from the Caribbean (US Virgin Islands) from 1895, Indonesia from 1923 and Taiwan from 1920.

Risk of Introduction

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The risk of P. quadrifida being accidentally introduced into new areas is low as it is a slow-growing species with minimal risk of contamination of crop seed. As a plant of interest medicinally, however, there is some risk of deliberate introduction.

Habitat

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P. quadrifida is found in the wild on bare patches of ground and among rocks, on sandy or stony soils, from sea-level up to 2000 m altitude. It is often involuntarily introduced by the agency of man and readily occupies newly disturbed areas, compost and rubbish heaps and fields. It is tolerant of a wide range of soils but prefers sand or sandy loams (PROTA, 2014). Singh (2005) notes that P. quadrifida may occur on alkaline soils but is not so common on saline soils. African Plant Database (2014) describe P. quadrifida as a plant of sandy river banks, open stony dry, or sub-desert grassland and shrubland, Acacia savanna; trampled areas, roadside and edges of cultivated fields. 

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Principal habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Natural
Rail / roadsides Principal habitat Natural
Urban / peri-urban areas Principal habitat Natural
Terrestrial ‑ Natural / Semi-naturalRiverbanks Secondary/tolerated habitat Natural
Wetlands Secondary/tolerated habitat Natural
Rocky areas / lava flows Secondary/tolerated habitat Natural
Scrub / shrublands Secondary/tolerated habitat Natural
Deserts Secondary/tolerated habitat Natural
Arid regions Principal habitat Natural
Littoral
Coastal areas Secondary/tolerated habitat Natural

Hosts/Species Affected

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In its native range of India, P. quadrifida is a significant weed in maize (Zea mays) (Subbulakshmi, 2009) and onions (Allium cepa) (Kachare et al., 2005). It is also a major weed of green gram (Vigna radiata) in the ‘summer’ season in India but less in black gram (V. mungo) in the monsoon season (Singh et al., 1991). It also occurs in pigeon pea (Cajanus cajan), cabbage (Brassica oleracea var. capitata), cauliflower (Brassica oleracea var. botrytis), cucurbits (Cucurbitaceae family) and teosinte (Zea species) in India and in coffee (Coffea species) in Kenya and sorghum (Sorghum bicolor) in Sudan.

Biology and Ecology

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Genetics

The Chromosome number for P. quadrifida is variously reported as 2n= 18, 36 and 48 (Sharma and Bhattachyya, 1956; Missouri Botanic Garden, 2014; PROTA, 2014), with 2n= 48 being the most frequently quoted.

The wide distribution and large variation, seen in P. quadrifida, points to great genetic variability which enables the species to adapt to new environments (PROTA, 2014).

Reproductive Biology

P. quadrifida reproduces by producing seeds which are largely dormant due to their hard seed coats. Scarification with sulphuric acid, dry storage for more than three months or high temperature pretreatment (58°C) for about one month can overcome dormancy. Germination is controlled by external factors such as light, the water content of the soil and temperature. Alternation between 12 and 35°C gives higher rates of germination than a constant intermediate temperature. The temperature requirement appears to be the main regulator of germination and seeds can germinate over wide ranges of moisture content and photoperiods (Shukla, 1972). Fresh seeds need light for germination, but this requirement disappears in older seeds.

It is also possible for P. quadrifida to regenerate vegetatively from small stem fragments (PROTA, 2014).

Physiology and Phenology

P. quadrifida has C4 metabolism (PROTA, 2014) and is therefore well adapted to benefit from high temperature conditions. The C4 metabolism helps to optimize photosynthesis in conditions of high heat and bright sunlight while enduring periods of limited water availability (Koch and Kennedy, 1982). As a prostrate herb, it is not otherwise especially competitive, but with extreme resistance to drought, it can become dominant in hot, arid conditions. The author collected a herbarium specimen of this species in Ethiopia and found that it was still alive and growing from between dry newspaper 12 months later (Chris Parker, Consultant, personal observation, 2015).

Generative development seems not to be influenced by photoperiod. Flowering in Pakistan is from August to March but flowering and fruiting can occur at any time of year when conditions are hot. This may be year-round in some areas. Under conditions of sub-lethal desiccation, the weed may propagate through shoot fragments. As only the most extreme desiccation is lethal, this is a very common phenomenon.

Longevity

P. quadrifida is an annual species and the established plants are not believed to perennate. The longevity of seeds in the soil is presumed to be at least a few years but no estimates have been seen.

Environmental Requirements

Although P. quadrifida is frequently referred to as a halophyte, therefore adapted to saline soils (Kokad and Ahmad, 2010), Singh (2005) suggests it is more tolerant of alkalinity than of salinity. Ecocrop (2014) indicates a low or moderate tolerance, not exceeding an electrical conductivity of 4 dS/m (equivalent to 40mM NaCl). Although Ecocrop (2014) indicates that P. quadrifida benefits from moderate or high soil fertility, it may thrive in extremely infertile conditions.

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])
Cs - Warm temperate climate with dry summer Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 22 36

Rainfall Regime

Top of page Bimodal
Summer
Uniform
Winter

Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • acid
  • alkaline
  • neutral

Soil texture

  • heavy
  • light

Special soil tolerances

  • infertile
  • saline
  • shallow
  • sodic

Notes on Natural Enemies

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There is no information available on the natural enemies of P. quadrifida. It is however likely that it is affected by many of those listed for P. oleracea.

Means of Movement and Dispersal

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Natural Dispersal

Seeds may float on the surface of water and are not normally blocked by sieves because of their small size. As a consequence, the species is dispersed through irrigation systems. Seeds of P. quadrifida are also easily spread by wind (PROTA, 2014).

Vector Transmission

Seeds of P. quadrifida may be ingested by birds and dispersed locally to new locations through their droppings (PROTA, 2014). 

Accidental Introduction

It is possible for P. quadrifida to be accidentally introduced to new areas through the movement of soil. This is possible as this species is able to tolerate desiccation and can grow vegetatively and regenerate from small fragments of stem. 

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Food Yes
Garden waste disposal Yes
Live food or feed trade Yes
Medicinal use Yes

Impact Summary

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CategoryImpact
Economic/livelihood Negative

Economic Impact

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P. quadrifida is regarded as an agricultural weed in its native range, where it competes with many crop species. The Flora of China Editorial Committee (2014) comments that it is sometimes a troublesome weed of cultivation, as the plants break up very easily and small fragments can act as propagules. This would have a negative economic impact however estimates of crop loss caused by the weed have not been calculated.

Environmental Impact

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There is little information available with regards to the environmental impacts of P. quadrifida. Although not very competitive, due to its extreme resistance to drought, it can become dominant in hot, arid conditions forming dense mats and therefore decreasing native biodiversity. As a halophyte, the accumulation of salt in the soil will also inhibit the growth of other species.

The association of this species with extremely infertile soil conditions leads farmers (in Ethiopia) to consider it the cause of infertility. It is referred to as ‘syphilis of the soil’ in the local language. There is, however, no evidence for any such effect.

Risk and Impact Factors

Top of page Invasiveness
  • Has a broad native range
  • Abundant in its native range
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Negatively impacts agriculture
Impact mechanisms
  • Competition - monopolizing resources
Likelihood of entry/control
  • Difficult/costly to control

Uses

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P. quadrifida is widely used as a salad or vegetable and may be cultivated e.g. Democratic Republic of Congo and in Rwanda. It may also be fed to pigs and poultry (PROTA, 2014). P. quadrifida typically has a high oxalate content which is potentially toxic but this is reduced by traditional cooking methods and rarely poses a risk. In Rajasthan, India, the leaves, after boiling, are used in preparing bread by mixing with millet flour and in Tamil Nadu the leaves and tender shoots are cooked and eaten as greens (Flowers of India, 2014).

P. quadrifida is also used as a traditional medicine in Africa and in China e.g. as a veterinary lactation stimulant or as an anti-abortifacient. It is also used for parasitic infection, for kidneys, pulmonary troubles, stomach troubles, venereal disease and as a diuretic, pain-killer or vermifuge (JSTOR Global Plants, 2014). Other uses mentioned by PROTA (2014) include as a diuretic, to treat rheumatism and gynaecological diseases, as a sedative, analgesic and cardiotonic, to treat fever, disorders of the urinary tract, worm diseases, as a tonic and choleretic, to treat dysentery and to apply externally to ulcers, eczema and dermatitis.

A study by Mulla and Paramjyothi Swamy (2012) confirmed anti-cancer activity against human colon cancer HT-29 cells. Naidu and Babu (2009) confirmed anti-fungal activity against Aspergillus niger and Candida albicans.

Uses List

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Animal feed, fodder, forage

  • Fodder/animal feed

Human food and beverage

  • Vegetable

Medicinal, pharmaceutical

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

Similarities to Other Species/Conditions

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The mat-forming habit and prostrate stems which can root from the nodes easily distinguish P. quadrifida from other members of the genus (Gilbert and Phillips, 2000).

In Africa P. pilosa has a similar habit to P. quadrifida and Geesink (1972) reported that a number of specimens from New Caledonia and the Pacific recorded as P. pilosa were often actually P. quadrifida. They can be distinguished from each other as P. pilosa has alternate leaves, five petals and more numerous stamens whereas P. quadrifida has four petals and the leaves are opposite (PROTA, 2014). Additionally P. hereroensis has often been confused with P. quadrifida but this species is an annual which does not root adventitiously (Gilbert and Phillips, 2000).

Prevention and Control

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Physical/Mechanical Control

Normal hoeing is relatively ineffective against P. quadrifida as the plant can survive extremely desiccating conditions and can re-grow from small fragments of stem. It is recommended either to burn or bury all removed material to prevent re-establishment.

Solarization for 30 and 45 days provided 100% control in North India (Nasr-Esfahani, 1993).

Movement Control

Due to its ability to survive desiccation and re-growth from small fragments of stem, it is especially important to prevent movement of weeded material via cultivation equipment.

Chemical Control

There are reports of P. quadrifida being controlled by herbicides and it is reported as being susceptible to MCPA and 2,4-D (Pfeiffer and Burleigh, 1958). In India, selective control was achieved with metoxuron, metribuzin, methabenzthiazuron, terbutryn, methabenzthiazuron + fluchloralin and fluchloralin + metribuzin (Randhawa et al., 1981). Additionally, directed applications of glyphosate + 2, 4-D have also been used in cropped fields (Tnau Agritech Portal, 2014).

References

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African Plant Database, 2015. African Plants Database (version 3.4.0). Geneva and Pretoria, Switzerland and South Africa: Conservatoire et Jardin botaniques de la Ville de Genève and South African National Biodiversity Institute. http://www.ville-ge.ch/musinfo/bd/cjb/africa/index.php?langue=an

Balikai RA, 1999. New record of plants of grapevine mealybug. Insect Environment, 5(2):81.

Ecocrop, 2014. Data sheet - Portulaca quadrifida. Rome, Italy: FAO. http://ecocrop.fao.org/ecocrop/srv/en/dataSheet?id=8882

Flora of China Editorial Committee, 2015. Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

Flora of Pakistan, 2015. Flora of Pakistan/Pakistan Plant Database (PPD). Tropicos website. USA: St. Louis, Missouri and Cambridge, Massachusetts. http://www.tropicos.org/Project/Pakistan

Flora Zambesiaca, 2015. Flora Zambesiaca online (eFloras). Richmond, Surrey, UK: Kew Databases. http://apps.kew.org/efloras/search.do

Flowers of India, 2014. Flowers of India. http://www.flowersofindia.net/

GBIF, 2015. Global Biodiversity Information Facility. http://www.gbif.org/species

Geesink R, 1969. An account of the genus Portulaca in Indo-Australia and the Pacific (Portulacaceae). Blumea, 17:274-301.

Geesink R, 1972. Portulacaceae. Flora Malesiana - Series 1, Spermatophyta, 7(1):121-133.

Gilbert MG; Phillips SM, 2000. A review of the opposite-leaved species of Portulaca in Africa and Arabia. Kew Bulletin, 55(4):769-802.

JSTOR Global Plants, 2014. JSTOR Global Plants Database. Ann Arbor, MI and New York, NY, USA: JSTOR. http://plants.jstor.org/

Kachare M; Pandey S; Kumar S, 2005. Integrated weed management in Kharif onion (Allium cepa L.) CV N-53. Farm Science Journal, 14(2):89-90.

Karim FM, 1993. New records of the Flora of the United Arab Emirates (Part 3). Arab Gulf Journal of Scientific Research, 11(3):391-401.

Koch KE; Kennedy RA, 1982. Crassulacean acid metabolism in the succulent C4 dicot, Portulaca oleracea L. under natural environmental conditions. Plant Physiology, 69(4):757-761

Kokad S; Ahmad S, 2010. Characterizing salt-tolerant plants using ecosystem and economic utilization potentials for Pakistan. Managing Natural Resources for Sustaining Future Agriculture; Research Briefings, 2(10):20 pp.

Missouri Botanical Garden, 2014. Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

Mulla SK; Swamy P, 2012. Anticancer activity of ethanol and polyphenol extracts of Portulaca quadrifida Linn. on human colon cancer cell lines. International Journal of Pharma and Bio Sciences, 3(3):P-488-P-498. http://www.ijpbs.net/vol-3/issue-3/pharma/53.pdf

Naidu LM; Babu NK, 2009. Evaluation of in vitro antimicrobial activity of Portulaca quadrifida L. International Journal of Plant Sciences (Muzaffarnagar), 4(2):494-497.

Nasr-Esfahani M, 1993. Soil-solarization for the control of weeds in Pantanagar. Integrated weed management for sustainable agriculture. Proceedings of an Indian Society of Weed Science International Symposium, Hisar, India, 18-20 November 1993 Hisar, Haryana, India; Indian Society of Weed Science, Vol. III:233-237

Nyffeler R; Eggli U, 2010. Disintegrating Portulacaceae: a new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon, 59(1):227-240.

Pfeiffer RK; Burleigh R, 1958. Papers. 1st Nat. Weed Contr. Conf., Gt Britain 1953. 10 pp.

PROTA, 2014. PROTA4U web database. Grubben GJH, Denton OA, eds. Wageningen, Netherlands: Plant Resources of Tropical Africa. http://www.prota4u.org/search.asp

Randhawa KS; Sandhu KS; Saimbhi MS, 1981. Studies on chemical weed control in (Pisum sativum L.) under different soil conditions. Abstracts of papers,Annual Conference of Indian Society of Weed Science, 1981:30.

Saimbhi MS; Sandhu KS; Daljit Singh; Kooner KS; Dhiman JS; Dhillon NPS, 1992. Use of herbicide for weed and disease control in cabbages. Advances in Horticultural Science, 6(3):125-128; 11 ref.

Sharma AK; Bhattacharyya NK, 1956. Cytogenetics of some members of Portulacaceae and related families. Caryologia, 8:257-274.

Shukla SP, 1972. Nature of dormancy and germination regulation in seeds of Portulaca quadrifida L. Japanese Journal of Ecology, 22(1):12-18.

Singh G; Ram IC; Singh D, 1991. Crop-weed competition studies in greenhouse and blackgram. Tropical Pest Management, 37(2):144-148.

Singh NT, 2005. Irrigation and soil salinity in the Indian subcontinent: past and present [ed. by Singh, N. T.]. Bethlehem, USA: Lehigh University Press, 404 pp.

Siwakoti M, 2000. Additions to the Flora of Nepal - V. Journal of Economic and Taxonomic Botany, 24(3):655-656.

Srivastava A; Joshi AG; Raole VM, 2013. Leaf epidermal micromorphology of Portulaca L. species found in Vadodara, Gujarat, India. Journal of Botany, 2013:Article ID 368238. http://www.hindawi.com/journals/jb/2013/368238/

Subbulakshmi S; Subbian P; Saravanan N; Prabakaran NK, 2009. Weed shift in a maize (Zea mays L.) - sunflower (Helianthus annuus L.) cropping system. Acta Agronomica Hungarica, 57(2):111-117. http://www.akademiai.com/content/119692/

Tnau Agritech Portal, 2014. Weed management: problem and parasitic weed management. Tamil Nadu, India. http://www.agritech.tnau.ac.in/agriculture/agri_weedmgt_problemandparasiticmgt.html

USDA-ARS, 2014. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

Wagner WL; Lorence DH, 2015. Flora of the West Indies. Washington DC, USA: Smithsonian Institution. http://botany.edu/antilles/WestIndies/index

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.

Contributors

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16/07/2014 Original text by:

Chris Parker, Consultant, Bristol, UK.

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