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Mesembryanthemum crystallinum
(crystalline iceplant)

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Datasheet

Mesembryanthemum crystallinum (crystalline iceplant)

Summary

  • Last modified
  • 06 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Mesembryanthemum crystallinum
  • Preferred Common Name
  • crystalline iceplant
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • M. crystallinum, commonly known as crystalline iceplant, is a prostrate halophyte native to north Africa, south-western Africa, western Asia and southern Europe. It has been both intentionally (as an ornamental...

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Pictures

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PictureTitleCaptionCopyright
Mesembryanthemum crystallinum (crystalline iceplant); flowering habit. La Geria region of Lanzarote, Canary Islands.
TitleFlowering habit
CaptionMesembryanthemum crystallinum (crystalline iceplant); flowering habit. La Geria region of Lanzarote, Canary Islands.
Copyright©Yummifruitbat/via wikipedia - CC BY-SA 2.5
Mesembryanthemum crystallinum (crystalline iceplant); flowering habit. La Geria region of Lanzarote, Canary Islands.
Flowering habitMesembryanthemum crystallinum (crystalline iceplant); flowering habit. La Geria region of Lanzarote, Canary Islands.©Yummifruitbat/via wikipedia - CC BY-SA 2.5
Mesembryanthemum crystallinum (crystalline iceplant); habit, immature plant. Growing at LZ-702 in Las Casitas, Yaiza, Lanzarote, Canary Islands. March, 2011.
TitleHabit
CaptionMesembryanthemum crystallinum (crystalline iceplant); habit, immature plant. Growing at LZ-702 in Las Casitas, Yaiza, Lanzarote, Canary Islands. March, 2011.
Copyright©Frank Vincentz/via wikipedia - CC BY-SA 3.0
Mesembryanthemum crystallinum (crystalline iceplant); habit, immature plant. Growing at LZ-702 in Las Casitas, Yaiza, Lanzarote, Canary Islands. March, 2011.
HabitMesembryanthemum crystallinum (crystalline iceplant); habit, immature plant. Growing at LZ-702 in Las Casitas, Yaiza, Lanzarote, Canary Islands. March, 2011.©Frank Vincentz/via wikipedia - CC BY-SA 3.0
Mesembryanthemum crystallinum (crystalline iceplant); close-up of the distinctive bladder cells.
TitleBladder cells
CaptionMesembryanthemum crystallinum (crystalline iceplant); close-up of the distinctive bladder cells.
Copyright©Schnobby/via wikipedia - CC BY-SA 3.0
Mesembryanthemum crystallinum (crystalline iceplant); close-up of the distinctive bladder cells.
Bladder cellsMesembryanthemum crystallinum (crystalline iceplant); close-up of the distinctive bladder cells.©Schnobby/via wikipedia - CC BY-SA 3.0

Identity

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Preferred Scientific Name

  • Mesembryanthemum crystallinum L.

Preferred Common Name

  • crystalline iceplant

Other Scientific Names

  • Cryophytum crystallinum (L.) Rothm.
  • Gasoul crystallinum (L.) N.E. Br.

International Common Names

  • English: common iceplant; iceplant
  • Spanish: barrilla de canarias; erba diacciola; escarchada; escarchosa
  • French: ficoïde cristalline; ficoide glaciale

Local Common Names

  • Germany: Eiskraut
  • Portugal: barrilha
  • Sweden: isört

Summary of Invasiveness

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M. crystallinum, commonly known as crystalline iceplant, is a prostrate halophyte native to north Africa, south-western Africa, western Asia and southern Europe. It has been both intentionally (as an ornamental) and accidentally (in sand used in ships ballasts) introduced into a number of countries in north and south America and Australia. This species is extremely stress-tolerant and during drought can switch from C3 photosynthesis to Crassulacean Acid Metabolism (CAM). Throughout its life M. crystallinum accumulates salt which is leached from plants upon death. The increase in the concentration of salt prevents the establishment of native plant species enabling seeds of M. crystallinum to germinate and grow and dominate in the salt marsh habitat. Monotypic stands of this species have been reported in Australia and California, USA. In California it is also reported as being a major threat to Chorizanthe pungens var. pungens [C. pungens], a native species included on the California Native Plant Society (CNPS) Inventory of Rare and Endangered plants.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Caryophyllales
  •                         Family: Aizoaceae
  •                             Genus: Mesembryanthemum
  •                                 Species: Mesembryanthemum crystallinum

Notes on Taxonomy and Nomenclature

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M. crystallinum belongs to the Aizoaceae family. The genus Mesembryanthemum contains around 25 different species that are typically found in the northern hemisphere, however they are also present in the south of Africa (CSIRO, 2004).

M. crystallinum was originally named Mesembrianthemum from mesembria, meaning mid-day, as the flowers were believed to open only in sun. The discovery of night-blooming species changed the spelling to indicate a flower with it’s fruits in the middle (mesos, middle and bryon, fruit). Crystallinum refers to the ice-like bubbles on the leaves with the common name arising from the claim that the leaves are cold to the touch on hot days (Natural History of Orange County, California 2015). The word Cryophytum is Greek for ice and plant (Natural History of Orange County, California 2015).

Description

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The following description is taken from the Flora of North America Editorial Committee (2015). Plants annual to biennial. Stems trailing, dichotomously branched, to 1 m. Leaves sessile or petiolate; petiole, ± clasping; blade ovate to spatulate, flat, 2-20 cm, margins undulate. Inflorescences terminal or axillary, cymes; proximal bracts opposite, leaflike; distal bracts alternate, reduced; flowering profusely. Flowers 7-10 mm diam.; hypanthium aging red, round; calyx lobes 5, unequal; petals 20-40, connate into tube, white, aging pink; stamens 30. Capsules coarsely papillate. Seeds 200, rough with minute tubercles. 

Plant Type

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Biennial
Succulent

Distribution

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M. crystallinum is native to north Africa, south-western Africa, Western Asia and southern Europe. However there are differing views as to whether it is a native or non-native of southern Europe (Kloot, 1983; Weber, 2004). It has been introduced and naturalized in a number of countries including three states of the USA, Mexico, Australia and southern South America (Queensland Government, 2011).

It has been reported however that the distribution of M. crystallinum in Egypt is decreasing and under threat due to human activities (Abd El-Gawad and Shehata, 2014).

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

IsraelPresentNativeUSDA-ARS, 2015

Africa

AlgeriaPresentNativeUSDA-ARS, 2015
EgyptPresentNativeEl Shayeb et al., 2002; Abd and El-Gawad Shehata, 2014; USDA-ARS, 2015
LibyaPresentNativeUSDA-ARS, 2015
MoroccoPresentNativeUSDA-ARS, 2015
NamibiaPresentNativeAgarie et al., 2009; Weeplian and Ho, 2014; USDA-ARS, 2015
Saint HelenaPresentNativeUSDA-ARS, 2015
South AfricaPresentNativeUSDA-ARS, 2015Eastern Cape, Northern Cape, Western Cape
Spain
-Canary IslandsPresentNativeUSDA-ARS, 2015
TunisiaPresentNativeUSDA-ARS, 2015

North America

MexicoPresentIntroducedUSDA-ARS, 2015Baja California, naturalized
USAPresentPresent based on regional distribution.
-ArizonaPresentIntroducedUSDA-ARS, 2015
-CaliforniaPresentIntroduced Invasive Vitousek, 1992; USDA-ARS, 2015
-PennsylvaniaPresentIntroducedUSDA-NRCS, 2015Naturalized

South America

ArgentinaPresentIntroducedUSDA-ARS, 2015Chubut, Rio Negro, naturalized
ChileAdams et al., 1998; Agarie et al., 2009; Madrigal-González et al., 2013; USDA-ARS, 2015
PeruPresentIntroducedUSDA-ARS, 2015Naturalized

Europe

CroatiaPresentNativeUSDA-ARS, 2015
FrancePresentNativeUSDA-ARS, 2015
-CorsicaPresentNativeUSDA-ARS, 2015
ItalyPresentNativeUSDA-ARS, 2015
PortugalPresentPresent based on regional distribution.
-AzoresPresentNativeUSDA-ARS, 2015
-MadeiraPresentNativeUSDA-ARS, 2015
SpainPresentNativeUSDA-ARS, 2015

Oceania

AustraliaPresentPresent based on regional distribution.
-New South WalesPresentIntroducedCouncil of Heads of Australasian Herbaria, 2015
-South AustraliaPresentIntroducedCouncil of Heads of Australasian Herbaria, 2015
-TasmaniaPresentIntroducedCouncil of Heads of Australasian Herbaria, 2015
-VictoriaPresentIntroducedCouncil of Heads of Australasian Herbaria, 2015
-Western AustraliaPresentIntroducedAdams et al., 1998; Council of Heads of Australasian Herbaria, 2015

History of Introduction and Spread

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M. crystallinum was introduced into the USA in the early 1500’s from sand ballast deposited from ships, in particular in Pennsylvania (Flora of North America Editorial Committee, 2015; Water Shed Health, 2015). In 1881, in the USA, M. crystallinum was advertised in seeds catalogues and was intentionally introduced into Australia as an ornamental plant (Kloot, 1987). It was also introduced for erosion control but was soon found to be ineffective (FloraBase, 2015). 

Risk of Introduction

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M. crystallinum produces a large number of seed which can be translocated into new areas by winds and by rabbits, rats and mice (AZ-WIPWG, 2005; Cal-IPC, 2015). It has a wide environmental tolerance and can grow in temperate climates and cold and warm deserts but is shade intolerant (AZ-WIPG, 2005).

Habitat

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M. crystallinum favours disturbed sites such as roadsides, cliffs, over-grazed areas and areas of coastal erosion, 0-100 m in altitude (FloraBase, 2015; Flora of North America Editorial Committee, 2015). It is however also found on undisturbed sites. It can grow well in saline environments and nutrient poor soils. 

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
 
Terrestrial – ManagedCultivated / agricultural land Secondary/tolerated habitat Harmful (pest or invasive)
Managed grasslands (grazing systems) Secondary/tolerated habitat Harmful (pest or invasive)
Disturbed areas Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Natural
Littoral
Coastal areas Principal habitat Harmful (pest or invasive)
Coastal areas Principal habitat Natural

Biology and Ecology

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Genetics

M. crystallinum has been reported to have a diploid number of 2n = 18 (Flora of North America Editorial Committee, 2015).

Reproductive Biology

There are five phases of growth for M. crystallinum; germinating seedlings, juvenile, adult, flowering fruiting and seed formation which respond differently to environmental stresses (Adams et al., 1998; Abd El-Gawad and Shehata, 2014). It flowers from early spring to summer, a process triggered by environmental stresses such as high salinity, drought or low temperatures. Flowers are hermaphrodite and are pollinated by insects. Flowers open during the day and close at and night (CSIRO, 2004). Under laboratory conditions M. crystallinum can produce around 15,000 seeds per plant (AZ-WIPWG, 2005). These may remain viable in the soil seed bank for up to two years (Plant Right, 2015). Seeds germinate during cold months and dormancy can be broken by any form of disturbance e.g. grazing, roads, drought and fire (Vivrette, 1999).

Physiology and Phenology

M. crystallinum is extremely stress-tolerant. During the dry season and periods of water stress, the mode of photosynthesis in adult plants of M. crystallinum switches from C3 to Crassulacean Acid Metabolism (CAM). This enables the plant to open its stomata, mainly at night, allowing carbon dioxide to be fixed as malic acid which the plant then converts to carbohydrates by standard photosynthesis during the day (Schmalzer and Hinkle, 1987). This process enhances the net gain of carbon and therefore increases seed production (CSIRO, 2004). Once seeds germinate, seedlings undergo a period of rapid vegetative growth which stops during the summer season. Plants possess specialized trichomes called epidermal bladder cells which have a number of proposed functions. These include as a storage of water, accumulation of salts, protection from UV rays and a role in plant defense (Barkla et al., 2012).

Longevity

Depending on environmental conditions, M. crystallinum can be an annual, biennial or perennial (CSIRO, 2004). However, its lifecycle is usually completed within a number of months.

Population Size and Structure

In some parts of Australia where it has been introduced, M. crystallinum occurs as scattered patches (CSIRO, 2004). It can however form more dense mats which cover the ground when present in favourable environments. For example, monotypic stands have been recorded in parts of Australia and California (AZ-WIPG, 2005) and in Egypt, its native range (Abd El-Gawad and Shehata, 2014).

Environmental Requirements

M. crystallinum is found on a wide range of soils including those with high salinity, with varying moisture contents and pH though it’s found not to grow in areas of shade (Plants for a Future, 2015). It has been suggested that optimal growth of M. crystallinum is obtained with a moderate salinity, between 0.1-0.2 M of NaCl. Higher concentrations of up to 0.5 M are also tolerated but this level often triggers reproductive development (CSIRO, 2004). M. crystallinum is not tolerant of frost and plants are killed.

Climate

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ClimateStatusDescriptionRemark
BS - Steppe climate Preferred > 430mm and < 860mm annual precipitation
BW - Desert climate Preferred < 430mm annual precipitation
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers

Soil Tolerances

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Soil drainage

  • free

Soil texture

  • heavy
  • light
  • medium

Special soil tolerances

  • saline

Means of Movement and Dispersal

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Natural Dispersal

The large number of seeds produced by M. crystallinum may readily be dispersed by winds (AZ-WIPWG, 2005).

Vector Transmission

It has been suggested that seeds of M. crystallinum may be dispersed by rabbits and mice (Cal-IPC, 2015).

Accidental Introduction

Seeds of M. crystallinum are small and can easily be attached to clothing, shoes and tyres and can be accidentally introduced into new areas (AZ-WIPWG, 2005). It is also thought to have been accidentally introduced into new areas as a contaminant in sand used as a ballast for ships (Cal-IPC, 2015)

Intentional Introduction

M. crystallinum was intentionally introduced into a number of countries for ornamental purposes, landscaping and erosion control (Cal-IPC, 2015). Seeds of M. crystallinum are also available for sale online.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Internet sales Yes Yes
Ornamental purposes Yes Cal-IPC, California Invasive Plant Council

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessionsSeeds may attach to clothing Yes The Arizona Wildland Invasive Plant Working Group, AZ-WIPWG
Host and vector organismsSeeds may dispersed by rabbits and mice Yes Cal-IPC, California Invasive Plant Council
Land vehicles Yes The Arizona Wildland Invasive Plant Working Group, AZ-WIPWG
Ship ballast water and sedimentAs a contaminant in sand used in ship ballasts Yes Cal-IPC, California Invasive Plant Council
Wind Yes The Arizona Wildland Invasive Plant Working Group, AZ-WIPWG

Impact Summary

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CategoryImpact
Environment (generally) Negative

Environmental Impact

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As a halophyte, M. crystallinum causes the accumulation of salt in the top soil (Abd El-Gawad and Shehata, 2014). When mature plants die the stored salt is leached into the soil thereby dramatically increasing the salinity. This makes it difficult for native species to grow, even after clearance and favours the germination of M. crystallinum seedlings (Vivrette and Muller, 1977; Schmalzer and Hinkle, 1987; Zavaleta et al., 2001; D’Antonio and Meyerson, 2002; Libik et al., 2004). This decreases biodiversity and monotypic stands of M. crystallinum have been reported in Australia and California, USA (AZ-WIPWG, 2005). M. crystallinum can also rapidly absorb moisture from the soil and leads to the build-up high levels of nitrate (Flora Base, 2015). Along with two other invasive species, M. crystallinum has been reported as a major threats to Chorizanthe pungens var. pungens [C. pungens], a native species included on the California Native Plant Species (CNPS) Inventory of Rare and Endangered plants.

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Chorizanthe pungens var. pungens (Monterey spineflower)NatureServe NatureServe; USA ESA listing as threatened species USA ESA listing as threatened speciesUSA

Social Impact

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Social Impact

Due to the accumulation of salt in M. crystallinum contains potentially toxic levels of oxalate. This can cause stock poisoning however plants are rarely eaten (FloraBase, 2015).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Pioneering in disturbed areas
  • Has propagules that can remain viable for more than one year
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
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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Social Benefit

The leaves of M. crystallinum can be cooked or eaten raw as a spinach substitute, as well as being pickled. The seed can be used a famine food when there is no other suitable food source (Sturtevant, 1888; Plants for the Future, 2015).M. crystallinum also has a number of medicinal properties where it is used to treat dysentery, liver and kidney diseases and to relieve pain and itching.

Environmental Services

Due to its ability to accumulate salt, M. crystallinum has been used for the desalination of soil and has been suggested that it could be used in bioremediation (Abd El-Gawad and Shehata, 2014).

Uses List

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Environmental

  • Land reclamation
  • Ornamental
  • Soil improvement

Human food and beverage

  • Vegetable

Medicinal, pharmaceutical

  • Traditional/folklore

Similarities to Other Species/Conditions

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M. crystallinum is morphologically similar to a number of species within the genus. These include M. nodiflorum and M. aitonis. However M. nodiuforum plants tends to be smaller, more upright with linear leaves overall and M. aitonis has fewer pimples and during flowering petals the same length as the sepals (CSIRO, 2015; Herbiguide, 2015).

Prevention and Control

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Control

Physical/Mechanical Control

Small isolated plants can be removed by hand during the spring and early summer (FloraBase, 2015). Dense patches of M. crystallinum may be cut close to the ground (Weber, 2004).

Chemical Control

Some herbicide can be used to control dense patches but application is recommened before seed set (Weber, 2004). In Australia however, herbicide resistance has been reported (FloraBase, 2015).

References

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Abd El-Gawad AM; Shehata HS, 2014. Ecology and development of Mesembryanthemum crystallinum L. in the deltaic Mediterranean coast of Egypt. Egyptian Journal of basic and applied science, 1(1):29-37.

Adams P; Nelson DE; Yamada S; Chmara W; Jensen RG; Bohnert HJ; Griffiths H, 1998. Tansley review no. 97. Growth and development of Mesembryanthemum crystallinum (Aizoaceae). New Phytologist, 138(2):171-190.

Agarie S; Kawaguchi A; Kodera A; Sunagawa H; Kojima H; Nose A; Nakahara T, 2009. Potential of the common ice plant, Mesembryanthemum crystallinum as a new high-functional food as evaluated by polyol accumulation. Plant Production Science, 12(1):37-46.

Barkla BJ; Vera-Estrella R; Pantoja O, 2012. Protein profiling of epidermal bladder cells from the halophyte Mesembryanthemum crystallinum. Proteomics, 12(18):2862-2865. http://onlinelibrary.wiley.com/doi/10.1002/pmic.201200152/full

Cal-IPC (California Invasive Plant Council), 2015. California Invasive Plants Council. www.cal-ipc.org. Berkeley, California, USA: California Invasive Plants Council.

Council of Heads of Australasian Herbaria, 2015. Australia's virtual herbarium. Australia: Council of Heads of Australasian Herbaria. http://avh.ala.org.au/#tab_simpleSearch

CSIRO, 2004. Mesembryanthemum crystallinum L., Australia: CSIRO. http://www.cpbr.gov.au/cpbr/WfHC/Mesembryanthemum/index.html

D'Antonio C; Meyerson LA, 2002. Exotic plant species as problems and solutions in ecological restoration: a synthesis. Restoration Ecology, 10(4):703-713.

El Shayeb FM; El Tantawy H; El Kholi A, 2002. Phytosociological studies on the wild Mesembryanthemum species in Egypt. Quantitive analysis of the representative community. Journal of Biological Sciences, 2(4):275-279.

Elorza MS; Bernado FG; Olivan AS; Iglesias LPG, 2010. Invasiveness of alien vascular plants in six arid zones of Europe, Africa and America. Lazaroa, 31:109-126.

Fish and Wildlife Service US, 2009. Monterey spineflower (Chorizanthe pungens var. pungens) 5-year review: summary and evaluation., USA: US Fish and Wildlife Service, 21 0pp. http://ecos.fws.gov/docs/five_year_review/doc2393.pdf

Flora of North America Editorial Committee, 2015. Flora of North America North of Mexico. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=1

FloraBase, 2015. The Western Australian Flora. Western Australia, Australia: Department of Environment and Conservation. http://florabase.dec.wa.gov.au/

Herbiguide, 2015. Herbiguide., Australia. http://www.herbiguide.com.au/

Herppich WB; Huyskens-Keil S; Schreiner M, 2008. Effects of saline irrigation on growth, physiology and quality of Mesembryanthemum crystallinum L., a rare vegetable crop. Journal of Applied Botany and Food Quality, 82(1):47-54.

Kloot PM, 1983. The role of common iceplant (Mesembryanthemum crystallinum) in the deterioration of medic pastures. Australian Journal of Ecology, 8(3):301-306.

Kloot PM, 1987. The naturalised flora of South Australia 4. Its manner of introduction. Journal of Adelaide Botanical Garden, 10(2):223-240.

Libik M; Pater B; Elliot S; Slesak I; Miszalski Z, 2004. Malate accumulation in different organs of Mesembryanthemum crystallinum L. following age-dependent or salinity-triggered CAM metabolism. Zeitschrift für Naturforschung. Section C, Biosciences, 59(3/4):223-228. http://znaturforsch.com/c.htm

Madrigal-González J; Cea AP; Sánchez-Fernández LA; Martínez-Tillería KP; Calderón JE; Gutiérrez JR, 2013. Facilitation of the non-native annual plant Mesembryanthemum crystallinum (Aizoaceae) by the endemic cactus Eulychnia acida (Cactaceae) in the Atacama Desert. Biological Invasions, 15(7):1439-1447. http://rd.springer.com/article/10.1007/s10530-012-0382-y

Missouri Botanical Garden, 2015. Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

Natural History of Orange County California, 2015. Mesembryanthemum crystallinum L. http://nathistoc.bio.uci.edu/Plants%20of%20Upper%20Newport%20Bay%20(Robert%20De%20Ruff)/Alliaceae/Aizoaceae/Mesembryanthemum%20crystallinum.htm

Plant Right, 2015. Mesembryanthemum crystallinum. http://www.plantright.org/species/mesembryanthemum-crystallinum

Plants and Fungi of South Western NSW, 2015. Mesembryanthemum crystallinum fact sheet. http://keys.lucidcentral.org/keys/v3/scotia/key/Plants%20and%20Fungi%20of%20south%20western%20NSW/Media/Html/Mesembryanthemum_crystallinum.htm

Plants for a Future, 2015. Plants for a future database. London, UK. http://www.pfaf.org

Queensland Government, 2015. Weeds of Australia, Biosecurity Queensland edition. Queensland, Australia. http://keyserver.lucidcentral.org/weeds/

Schmaizer PA; Hinkle CR, 1987. Species biology and potential for controlling four exotic plants (Ammophila arenaria, Carpobrotus edulis, Cortadera jubata and Gasoul crystallinum) on Vandenberg Air Force Base, California. NASA Technical Memorandum (NASA-TM-10980).

Sturtevant EL, 1888. History of garden vegetables (Continued). The American Naturalist, 22(261):802-808.

The Arizona Wildland Invasive Plant Working Group (AZ-WIPWG), 2005. Plant assessment form - Mesembryanthemum crystallinum. http://sbsc.usgs.gov/research/projects/swepic/SWVMA/PLANTPDF/Mesembryanthemum_crystallinum_AZ_PAF

USDA-ARS, 2015. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

USDA-NRCS, 2015. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Vitousek PM, 1992. Effects of alien plants on native ecosystems. http://manoa.hawaii.edu/hpicesu/book/1992_chap/03.pdf

Vivrette NJ, 1999. 5th California Islands Symposium (1999)., USA 141-142.

Vivrette NJ; Muller CH, 1977. Mechanism of invasion and dominance of coastal grassland by Mesembryanthemum crystallinum. Ecological Monographs, 47(3):301-318.

Water Shed Health, 2015. Mesembryanthemum crystallinum and M. nodiflorum., USA. http://www.watershedhealth.org/weedwatch/docs/matrix/Mesembryanthemum_spp_042907

Weber E, 2004. Invasive plant species of the world: a reference guide to environmental weeds. Wallingford, UK: CABI Publishing, 560 pp.

Weeplian T; Ho Y, 2014. Effects of photoperiod and photosynthetic photon flux on seed germination and growth of Mesembryanthemum crystallinium L seedling. In: International Conference on Plant Factory, DOI: 10.13140/2.1.3024.8000. Koyoto, Japan.

Zavaleta ES; Hobbs RJ; Mooney HA, 2001. Viewing invasive species removal in a whole-ecosystem context. Trends in Ecology & Evolution, 16(8):454-459.

Contributors

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23/09/15 Original text by:

Emma Thompson, CABI, UK

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