Tetragonia tetragonioides (New Zealand spinach)
Index
- Pictures
- Identity
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Description
- Plant Type
- Distribution
- Distribution Table
- History of Introduction and Spread
- Introductions
- Risk of Introduction
- Habitat
- Habitat List
- Biology and Ecology
- Climate
- Soil Tolerances
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Impact Summary
- Economic Impact
- Environmental Impact
- Social Impact
- Risk and Impact Factors
- Uses
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- References
- Contributors
- Distribution Maps
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Top of pagePreferred Scientific Name
- Tetragonia tetragonioides (Pall.) Kuntze
Preferred Common Name
- New Zealand spinach
Other Scientific Names
- Demidovia tatragonioides Pallas, Enum. Hort. Demidof 150, t. 1. 1781.
- Tatragonia cornuta Gaertner, Fruct. Sem. Pl. 2: 483. 1791.
- Tetragonia expansa Murray, Commentat. Soc. Regiae Sci. Gott. 6: 13. 1783
- Tetragonia expansa THUNB. ex MURR.
- Tetragonia halimifolia G. Foster, Fl. Ins. Austr. 39. 1786.
- Tetragonia inermis 1852. F. Mueller, Linnaea 25: 384.
- Tetragonia japonica Thunberg, Fl. Jap. 208. 1784.
- Tetragonia quadricornis Stokes, Bot. Mat. Med. 3: 127. 1812, nom. illegit.
- Tetragonia tetragonoides
International Common Names
- English: New Zealand spinach; perpetual spinach; sea spinach
- Spanish: espinaca de Nueva Zelandia; Espinacia de Nueva Zelandica; spinaca Nuevo Zelandia
- French: epinard; Epinard de la Nouvelle-Zelande; tetragon; tetragone; tetragone cornue
- Chinese: xin xi lan bo cai; yeung poh tsoi
- Portuguese: espinafre de Nova Zelandia
Local Common Names
- Chinese: summer spinach
- Russian: Novozelandskii shipinat; tetragonia shpinatnaia
- Australia: Botany Bay greens; Sydney greens; warrigal cabbage; warrigal cabbage; warrigal greens; warrigal spinach
- Brazil: espinafre da Nova Zelandia
- Croatia: Novozealandski shpinat
- Denmark: Newzealandsk spinat
- Germany: Neuseelaender Spinat; Neuseelandischer spinat
- Greece: spanaki Neas Zilandias
- Hungary: Ujzelandi paraj
- Italy: spinacio della Nouva Zelanda; Spinacio della Nouva-Zelande
- Japan: tsuruna; Tsuruna
- Netherlands: Nieuwzeelandse spinazie; Niewzeelandse Spinazie
- New Zealand: kokihi; paraihia; rengamutu; rengarenga; tutae-ika-moana
- Norway: Ny-Zealandsk spinat
- Philippines: baguio spinach
- Poland: szpinak Nowozelandzki
- Serbia: Novozelandski spanac(h)
- Slovenia: Novozelandska schpinacha
- South Africa: Nu Zeeland spinasie
- Sweden: Ny-seelaendisk spenat; Nyzeelandsk spenat
- Switzerland: neuseelander-spinat
EPPO code
- TEATE (Tetragonia tetragonioides)
Summary of Invasiveness
Top of pageT. tetragonioides is a leafy herb native to the Far East, parts of Australia, New Zealand and some Pacific Islands. It has been introduced to Africa, the Americas, Europe and parts of Asia. It is considered invasive in coastal habitats in Chile, Hawaii, Florida and California, and is one of several principal invaders in Reunion. The strongest case for its invasiveness is made in California, where it is controlled in natural areas. T. tetragonioides establishes in and competes with coastal, beach, dune, cliff and salt marsh vegetation. It is naturalized mainly in frost free coastal climates but it persists after cultivation in cold climates, such as in northern USA and Europe.
Taxonomic Tree
Top of page- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Dicotyledonae
- Order: Caryophyllales
- Family: Aizoaceae
- Genus: Tetragonia
- Species: Tetragonia tetragonioides
Notes on Taxonomy and Nomenclature
Top of pageA variety of latin binomials have been used for the New Zealand spinach Tetragonia tetragonioides (currently accepted). The most frequent synonym in the literature is Tetragonia expansa, and there may be as much literature referring to the latter as to the former.
Description
Top of pageFrom Vivrette (2004):
Stems: mat-forming, spreading to 30-140 cm. Leaves: 10-100 by 50-50 mm; petiole 10-30 mm, winged; blade pale green abaxially, dark green adaxially, midvein and lateral veins raised abaxially, ovate-rhombic to triangular, base truncate, papillate with larger papillae abaxially. Inflorescences: peduncle to 2 mm. Flowers: sometimes unisexual; calyx lobes spreading, yellow, ovate to half orbiculate, 2 mm; stamens clustered or scattered. Fruits: turbinate, 8-12 mm; horns 4-6. Seeds: smooth.
From Morris and Duretto (2009):
Annual or perennial herb, decumbent or ascending; stems to 1 m long. Leaves broadly hastate, rhomboid or lanceolate; petiole 10–45 mm long, papillose, decurrent; blade 10–80(–100?) mm long, 5–50 mm wide, dark green above, paler below with larger papillae. Flowers axillary, solitary or rarely 2 together, 8–10 mm diam.; pedicels to 5 mm long. Calyx lobes 4–5, 1 or 2 of them ovate-semiorbicular, larger than the rest, the smaller lobes ovate-triangular to lanceolate, obtuse or acute; adaxial surface yellowish, minutely papillose; abaxial surface green, papillose. Stamens 8–16, in groups alternating with the calyx lobes. Stigmas 3–8(–10?) in 2 groups, equal in number to the loculi. Fruit indehiscent, at first green, becoming dry and bony, to 13 mm long and 12 mm wide, globular or turbinate variously ridged at the summit; ridges produced into short equal or unequal horns. Seeds in 2 rows, c. 2.5 mm long, pyriform. Flowering and fruiting throughout year.
Distribution
Top of pageT. tetragonioides has been introduced to Africa, the Americas, Europe, Indonesia, Iran, Israel and the Philippines. It is widespread in Europe. Its seeds have been on sale in London since 1835 or earlier, and it was recorded in unlikely places such as Switzerland fairly early on after its initial introduction to Europe (Déséglise, 1881). Its popularity as an easy-to-grow spinach alternative means its introduced range is most likely larger than indicated in the distribution table.
There are disagreements over its native range. It seems that most agree that it is native from the Far East south to Australia (e.g. Harris, 1998; Hara et al., 2008; Morris and Duretto, 2009). Most accounts now regard it as non-native and naturalized in South America, including Chile (Reiche and Philippi, 1898; Zuloaga and Morrone, 1996; Ugarte et al., 2011; Missouri Botanical Garden, 2013; Zappi, 2014). However, older accounts still hold some influence in the literature, and its nativity in Chile and Argentina is still frequently claimed (Sturtevant, 1919; Roskruge, 2011). Occasionally the plant is claimed as native to Africa (e.g. Ugarte et al., 2011), but probably more because of the prevalence of the genus and the family in South Africa than due to a knowledge of the phylogeography of the species itself. In Australia, T. tetragonioides has established in New South Wales and Tasmania, the two states it is not native to (Randall, 2007).
Distribution Table
Top of pageThe distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
Last updated: 10 Feb 2022Continent/Country/Region | Distribution | Last Reported | Origin | First Reported | Invasive | Reference | Notes |
---|---|---|---|---|---|---|---|
Africa |
|||||||
Cabo Verde | Present | ||||||
Kenya | Present | Introduced | Has escaped from cultivation in some places | ||||
Madagascar | Present | 1955 | Introduced | Littoral Ford Dauphin ocean edge | |||
Mauritius | Present | 1993 | Introduced | Invasive | Became rare after invasive browsers were removed | ||
Morocco | Present | Introduced | Adventive | ||||
Mozambique | Present | Introduced | Has escaped from cultivation in some places | ||||
Réunion | Present, Widespread | Introduced | Invasive | In a list of the principal invasive species | |||
Rwanda | Present | Introduced | Has escaped from cultivation in some places | ||||
Saint Helena | Present | Introduced | Single herbarium specimen | ||||
Senegal | Present | Introduced | Has escaped from cultivation in some places | ||||
Somalia | Present | Introduced | Has escaped from cultivation in some places | ||||
South Africa | Present | Introduced | Eastern region coastal site | ||||
Uganda | Present | Introduced | Has escaped from cultivation in some places | ||||
Zambia | Present | Introduced | Has escaped from cultivation in some places | ||||
Zimbabwe | Present | Introduced | Has escaped from cultivation in some places | ||||
Asia |
|||||||
China | Present | Present based on regional distribution. | |||||
-Fujian | Present | Native | Sandy shores, also cultivated | ||||
-Guangdong | Present | Native | Sandy shores, also cultivated | ||||
-Jiangsu | Present | Native | Sandy shores, also cultivated | ||||
-Yunnan | Present | Native | Sandy shores, also cultivated | ||||
-Zhejiang | Present | Native | Sandy shores, also cultivated | ||||
Indonesia | Present, Only in captivity/cultivation | Introduced | |||||
Iran | Present | 2012 | Introduced | Cultivated in experiments | |||
Israel | Present, Only in captivity/cultivation | Introduced | Crop exported to US | ||||
Japan | Present | Present based on regional distribution. | |||||
-Hokkaido | Present, Only in captivity/cultivation | Native | South west of region | ||||
-Honshu | Present, Only in captivity/cultivation | Native | Common sometimes cultivated as vegetable | ||||
-Shikoku | Present, Only in captivity/cultivation | Native | |||||
Myanmar | Present | Native | |||||
Philippines | Present, Only in captivity/cultivation | Introduced | |||||
South Korea | Present, Only in captivity/cultivation | Native | Very common native plant on dunes in Korea | ||||
Taiwan | Present | Native | Sandy shores, also cultivated | ||||
Europe |
|||||||
Austria | Present | Introduced | First reported: <1885 | ||||
Belgium | Present | Introduced | 1888 | ||||
Czechia | Present | Introduced | 1918 | ||||
France | Present | Introduced | Established on dunes in Bordeaux on coast | ||||
-Corsica | Present | Introduced | 1994 | ||||
Hungary | Present, Localized | Introduced | |||||
Italy | Present | ||||||
Netherlands | Present | Introduced | |||||
Poland | Present | Introduced | 1931 | ||||
Portugal | Present | Introduced | |||||
-Azores | Present | Introduced | 1857 | ||||
-Madeira | Present | Introduced | 1864 | ||||
Spain | Present, Widespread | Introduced | |||||
-Balearic Islands | Present | Introduced | |||||
-Canary Islands | Present | Introduced | Not listed as invasive by others (Arevalo et al., 2005) | ||||
Switzerland | Present, Only in captivity/cultivation | Introduced | |||||
United Kingdom | Present, Localized | Introduced | Associated with bird seed in Britain | ||||
-Channel Islands | Present, Localized | Introduced | Not considered problematic by this author but widely scattered and in disturbed sites | ||||
North America |
|||||||
Costa Rica | Present | Introduced | Adventive in abandoned lot | ||||
Guadeloupe | Present | Introduced | |||||
Mexico | Present | Introduced | Cultivated and established on an upper beach | ||||
Nicaragua | Present | Introduced | Cultivated | ||||
Puerto Rico | Present, Only in captivity/cultivation | Introduced | Cultivated | ||||
United States | Present | Present based on regional distribution. | |||||
-California | Present, Localized | Introduced | 1891 | Invasive | Recorded wild at this date | ||
-Connecticut | Present | Introduced | |||||
-Delaware | Present, Localized | Introduced | Persistent after cultivation | ||||
-Florida | Present, Localized | Introduced | Invasive | Established on dunes and in salt marsh | |||
-Georgia | Present, Localized | Introduced | Persistent after cultivation | ||||
-Hawaii | Present, Localized | 2014 | Introduced | 1909 | Invasive | ||
-Maryland | Present | Introduced | |||||
-New York | Present | Introduced | |||||
-North Carolina | Present, Localized | Introduced | Persistent after cultivation | ||||
-Ohio | Present | Introduced | |||||
-Oklahoma | Present | Introduced | |||||
-Oregon | Present, Only in captivity/cultivation | Introduced | |||||
-South Carolina | Present | Introduced | |||||
-Washington | Present | Introduced | |||||
-Wisconsin | Present, Localized | Introduced | Adventive, found in strawberry patch | ||||
Oceania |
|||||||
Australia | Present, Widespread | Native | Invasive | Naturalised within Australia outside of its native range | |||
-New South Wales | Present, Localized | Introduced | Invasive | Low priority invasive species | |||
-Northern Territory | Present | Native | |||||
-Queensland | Present | Native | |||||
-South Australia | Present | Native | |||||
-Tasmania | Present | Native | |||||
-Western Australia | Present | Introduced | |||||
French Polynesia | Present | Native | |||||
New Caledonia | Present | Native | |||||
New Zealand | Present, Widespread | Native | |||||
Norfolk Island | Present | Native | Mentioned related to Philip Island invasive animal removal recovery | ||||
South America |
|||||||
Argentina | Present | Introduced | |||||
Brazil | Present | Present based on regional distribution. | |||||
-Parana | Present, Localized | Introduced | Naturalized | Naturalised | |||
-Rio de Janeiro | Present, Localized | Introduced | Naturalized | Naturalised | |||
-Santa Catarina | Present, Localized | Introduced | Naturalized | Naturalised | |||
-Sao Paulo | Present, Localized | Introduced | Naturalized | Naturalised | |||
Chile | Present, Localized | 2011 | Introduced | Invasive | Cultivated and wild | ||
-Easter Island | Present, Widespread | Native | |||||
Ecuador | Present, Only in captivity/cultivation | 2010 | Introduced | First recorded in Galapagos Islands in 2003 during alien plant inventories. Found on Isabela, Santa Cruz and San Cristobal | |||
Peru | Present | Arequipa, widely cultivated | |||||
Uruguay | Present | Mentioned as occurring Uruguay |
History of Introduction and Spread
Top of pageT. tetragonioides is regarded as native in from China and Korea to Japan, Australia and New Zealand (Allan et al., 1961; Ohwi, 1965; Kim, 2005; Morris and Duretto, 2009; eFloras, 2013). It is probably native in other sites in between these countries as it can spread naturally via ocean currents (Abe, 2006). However, this ability to spread by ocean currents is shared by others in the genus, which could potentially lead to an overestimation of its native range.
It was collected in South America in 1847 (Ugarte et al., 2011); by some accounts, T. tetragonioides may be native in South America (Morong and Britton, 1893; Sturtevant, 1919; Roskruge, 2011), but now most authors now class it as a cultivated, adventive or naturalized plant there (Macbride and Dahlgren, 1937; Zuloaga and Morrone, 1996; Ugarte et al., 2011; Zappi, 2014).
T. tetragonioides was reported by Captain Cook and eaten by his crew in 1770. Joseph Banks is credited with giving seeds to Kew Botanic Gardens, UK, in 1772, and by 1824 it was on sale in London. It then probably spread throughout Europe rather quickly (Sturtevant, 1919). Wild populations were established in France in the 1920s (Tempére, 1920).
T. tetragonioides was on sale in New York in seed catalogues in 1828 after initially being spread by members of a horticultural society (Sturtevant, 1919; Roskruge, 2011). It was recorded wild in California in 1891 and Hawaii in 1909. It was already known in South America by the mid to late 1800s (Morong and Britton, 1893; Reiche and Philippi, 1898; Sturtevant, 1919; Ugarte et al., 2011).
The timing of the spread is harder to pinpoint in Africa, but it is clearly widely planted there (Grubben, 2004). Its suitability as a vegetable in coastal habitats means that it has been introduced to many oceanic islands for human consumption (PIER, 2014). By all accounts the plant is probably more widely distributed than is documented here.
Introductions
Top of pageIntroduced to | Introduced from | Year | Reason | Introduced by | Established in wild through | References | Notes | |
---|---|---|---|---|---|---|---|---|
Natural reproduction | Continuous restocking | |||||||
California | <1891 | Horticulture (pathway cause) | Yes | Robbins (1940) | ||||
Chile | <1898 | |||||||
Galapagos Islands | <2003 | Horticulture (pathway cause) | No | Guézou et al. (2010) | Probably introduced no earlier than 1940s when significant settlement started only known from inhabited islands | |||
Hawaii | <1909 | Horticulture (pathway cause) | Yes | Wester (1992) | ||||
UK | New Zealand | 1772 | Horticulture (pathway cause) | Yes | Sturtevant (1919) | |||
USA | UK | 1828 | Horticulture (pathway cause) | Sturtevant (1919) |
Risk of Introduction
Top of pageT. tetragonioides is most likely to be introduced to new sites by gardeners wishing to grow it, and its seeds are available via internet and catalogue mail-order. The seeds are easily transported and remain viable for long periods in storage. Spread from one river flood plain or coastal site to another is also possible since the seeds are known to disperse via fresh and salt water currents (Abe, 2006; Nicol et al., 2009); though not strictly a halophyte, T. tetragonioides is reasonably salt tolerant (Roskruge, 2011).
Habitat
Top of pageT. tetragonioides is generally associated with coastal habitats in its native and introduced invasive range. It grows well on dunes, beaches, salt marshes, coastal cliffs and other coastal locations (Abe, 2006; Robbins, 1940; Roskruge, 2011). It is naturalized mainly in frost free coastal climates but it persists after cultivation in cold climates, such as in northern USA and Europe. It may also persist unassisted in abandoned gardens, or spread slowly from any place it is planted.
Habitat List
Top of pageCategory | Sub-Category | Habitat | Presence | Status |
---|---|---|---|---|
Terrestrial | ||||
Terrestrial | Managed | Cultivated / agricultural land | Present, no further details | Natural |
Terrestrial | Managed | Cultivated / agricultural land | Present, no further details | Productive/non-natural |
Terrestrial | Managed | Disturbed areas | Present, no further details | Natural |
Terrestrial | Managed | Disturbed areas | Present, no further details | Productive/non-natural |
Terrestrial | Natural / Semi-natural | Riverbanks | Principal habitat | Harmful (pest or invasive) |
Terrestrial | Natural / Semi-natural | Riverbanks | Principal habitat | Natural |
Terrestrial | Natural / Semi-natural | Riverbanks | Principal habitat | Productive/non-natural |
Terrestrial | Natural / Semi-natural | Rocky areas / lava flows | Principal habitat | Harmful (pest or invasive) |
Terrestrial | Natural / Semi-natural | Rocky areas / lava flows | Principal habitat | Natural |
Littoral | Coastal areas | Principal habitat | Harmful (pest or invasive) | |
Littoral | Coastal areas | Principal habitat | Natural | |
Littoral | Coastal areas | Principal habitat | Productive/non-natural | |
Littoral | Coastal dunes | Principal habitat | Harmful (pest or invasive) | |
Littoral | Coastal dunes | Principal habitat | Natural | |
Littoral | Coastal dunes | Principal habitat | Productive/non-natural | |
Littoral | Salt marshes | Principal habitat | Harmful (pest or invasive) | |
Littoral | Salt marshes | Principal habitat | Natural | |
Littoral | Salt marshes | Principal habitat | Productive/non-natural | |
Freshwater | Irrigation channels | Present, no further details | Natural | |
Freshwater | Irrigation channels | Present, no further details | Productive/non-natural | |
Freshwater | Rivers / streams | Present, no further details | Natural | |
Freshwater | Rivers / streams | Present, no further details | Productive/non-natural | |
Brackish | Estuaries | Principal habitat | Natural | |
Brackish | Estuaries | Principal habitat | Productive/non-natural |
Biology and Ecology
Top of pageGenetics
Chromosome number 2n = 32
Reproductive biology
T. tetragonioides is predominantly self-pollinated but cross-pollination may occur (Grubben, 2004). The fruits fall to the ground on ripening so it can reseed itself. Germination can be irregular but generally takes less than 20 days, and may occur naturally in spring in either temperate or cold climates (Roskruge, 2011). Under cultivation plants reached harvestable size 50-55 days after planting (Roskruge, 2011).
Physiology and phenology
Germination takes up to 20 days and first flowers 59-63 days after planting. Late maturing plants can produce an abundance of seeds (Roskruge, 2011).
Associations
T. tetragonioides is associated with coastal plants such as beach morning glory, Ipomoea imperati.
Environmental requirements
T. tetragonioides prefers mesic conditions but there is evidence that it is drought tolerant (Hara et al., 2008). It grows well on dunes, beaches, salt marshes and other coastal locations (Abe, 2006; Robbins, 1940; Roskruge, 2011). Plants are reasonably salt-tolerant, though seedlings may not withstand as much salt as mature plants. T. tetragonioides is also tolerant of a variety of soil types, preferring pH 5.8-7.5 (Roskruge, 2011). It is reported to prefer frost-free sites (Grubben, 2004; Roskruge, 2011), although it may encounter frost in parts of its native range, and can persist after cultivation in much colder sites (such as Wisconsin, Connecticut, Oregon and Washington, USA, and Suffolk, UK), suggesting that as long as it can flower and fruit in a single growing season, the seeds will not be killed by freezing temperatures.
Climate
Top of pageClimate | Status | Description | Remark |
---|---|---|---|
Af - Tropical rainforest climate | Tolerated | > 60mm precipitation per month | |
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 | Tolerated | < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25]) | |
Aw - Tropical wet and dry savanna climate | Tolerated | < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25]) | |
B - Dry (arid and semi-arid) | Preferred | < 860mm precipitation annually | |
Cf - Warm temperate climate, wet all year | Preferred | Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year | |
Cs - Warm temperate climate with dry summer | Preferred | Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers | |
Cw - Warm temperate climate with dry winter | Preferred | Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters) | |
Df - Continental climate, wet all year | Preferred | Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year) |
Soil Tolerances
Top of pageSoil reaction
- acid
- neutral
Soil texture
- light
- medium
Special soil tolerances
- infertile
- saline
- shallow
Natural enemies
Top of pageNatural enemy | Type | Life stages | Specificity | References | Biological control in | Biological control on |
---|---|---|---|---|---|---|
Alternaria alternata | Pathogen | Plants|Whole plant | not specific | |||
Aphis gossypii | Herbivore | Plants|Leaves | not specific | |||
Botryotinia fuckeliana | Pathogen | Plants|Whole plant | not specific | |||
Brevipalpus californicus | not specific | |||||
Helminthosporium | Pathogen | Plants|Whole plant | not specific | |||
Heterodera schachtii | not specific | |||||
Meloidogyne hapla | Parasite | Plants|Roots | not specific | |||
Meloidogyne incognita | Parasite | Plants|Roots | not specific | |||
Meloidogyne javanica | Parasite | Plants|Roots | not specific | |||
Myzus persicae | Herbivore | Plants|Leaves | not specific | |||
Tetranychus urticae | Herbivore | Plants|Leaves | not specific | |||
Verticillium albo-atrum | Pathogen | Plants|Whole plant | not specific |
Notes on Natural Enemies
Top of pageNone of the pests in the Natural Enemies table are described as serious pests of T. tetragonioides (Roskruge, 2011). Some of the natural enemies can carry pathogens of potatoes; specifically, Helminthosporium sp. is commonly known as silver scurf in potatoes, and Verticillium alboatrum is a wilt disease found on potatoes (APHIS, 2004; Roskruge, 2011).
Means of Movement and Dispersal
Top of pageNatural dispersal (non-biotic)
The floating horned seeds of T. tetragonioides can be transported by water, including rivers and oceans (Abe, 2006; Nicol et al., 2009). Plants have established on a recently formed volcanic island in the Pacific Ocean, near Japan, apparently from seed transported naturally on ocean currents (Abe, 2006).
Vector transmission (biotic)
T. tetragonioides has not been associated with animal dispersal in the literature, but transport of plants for bird nesting materials could be possible, as could ingestion by tortoises.
Accidental introduction
Plants have been dispersed in bird seed in Britain (Hanson and Mason, 1985).
Intentional introduction
This plant has been deliberately introduced around the world as an easy-to-grow spinach alternative (Sturtevant, 1919; Cambie and Ferguson, 2003; Roskruge, 2011).
Pathway Causes
Top of pageCause | Notes | Long Distance | Local | References |
---|---|---|---|---|
Botanical gardens and zoos | e.g. Kew Garden is likely source for plants in Europe | Yes | Yes | Sturtevant (1919) |
Flooding and other natural disasters | Floodplain plant in native range | Yes | Yes | Nicol et al. (2009) |
Garden waste disposal | Common garden plant worldwide | Yes | Grubben and Denton (2004); Roskruge (2011); Sturtevant (1919) | |
Horticulture | Common garden plant worldwide | Yes | Yes | Grubben and Denton (2004); Roskruge (2011) |
Internet sales | Common garden plant worldwide, and long available in gardening catalogues | Yes | Grubben and Denton (2004); Roskruge (2011); Sturtevant (1919) | |
Medicinal use | Possible since it is known as an ulcer reducer | Yes | Cambie and Ferguson (2003); Okuyama and Yamazaki (1983) | |
Nursery trade | Common garden plant worldwide, and long available in gardening catalogues | Yes | Yes | Grubben and Denton (2004); Roskruge (2011); Sturtevant (1919) |
Pet trade | Bird seed | Yes | Yes | Hanson and Mason (1985) |
Seed trade | Common garden plant worldwide, and long available in gardening catalogues | Yes | Yes | Grubben and Denton (2004); Roskruge (2011); Sturtevant (1919) |
Pathway Vectors
Top of pageVector | Notes | Long Distance | Local | References |
---|---|---|---|---|
Floating vegetation and debris | Seeds known to float in water - principal natural dispersal mechanism | Yes | Abe (2006) | |
Luggage | Unknown frequency but it is a desirable garden plant | Yes | Yes | |
Available for sale in plant catalogues in the 1800s | Yes | Yes | Roskruge (2011) | |
Soil, sand and gravel | Possible since it grows on dunes | Yes | Yes | Kim (2005) |
Water | This is the natural means of dispersal | Yes | Yes | Abe (2006) |
Plant Trade
Top of pagePlant parts liable to carry the pest in trade/transport | Pest stages | Borne internally | Borne externally | Visibility of pest or symptoms |
---|---|---|---|---|
Flowers/Inflorescences/Cones/Calyx | Yes | Yes | Pest or symptoms not visible to the naked eye but usually visible under light microscope | |
Growing medium accompanying plants | Yes | Yes | Pest or symptoms not visible to the naked eye but usually visible under light microscope | |
Leaves | Yes | Yes | Pest or symptoms not visible to the naked eye but usually visible under light microscope | |
Seedlings/Micropropagated plants | Yes | Yes | Pest or symptoms not visible to the naked eye but usually visible under light microscope | |
Stems (above ground)/Shoots/Trunks/Branches | Yes | Yes | Pest or symptoms not visible to the naked eye but usually visible under light microscope |
Plant parts not known to carry the pest in trade/transport |
---|
Bark |
Bulbs/Tubers/Corms/Rhizomes |
Fruits (inc. pods) |
True seeds (inc. grain) |
Wood |
Impact Summary
Top of pageCategory | Impact |
---|---|
Cultural/amenity | Positive |
Economic/livelihood | Positive |
Environment (generally) | Positive and negative |
Human health | Positive |
Economic Impact
Top of pageThere are no reports of T. tetragonioides impacting crops. Costs of control in terms of labour and herbicide costs are not recorded. It is grown as a crop in Israel, and as a common garden plant around the world (APHIS, 2004; Grubben, 2004; Roskruge, 2011).
Environmental Impact
Top of pageImpact on habitats
T. tetragonioides is known to establish significant populations in coastal habitats outside of its native range, including dunes, salt marshes and coastal cliffs in Hawaii, Florida, California and Reunion (Robbins, 1940; Starr and Starr, 2006; Zomlefer et al., 2007; Soubeyran, 2008).
Impact on biodiversity
The spread of T. tetragonioides outside of its native range in coastal habitats is not associated with many accounts of its negative impacts on individual species. The spreading habit (almost vine-like) and abundant fleshy leaves of T. tetragonioides give it significant competitive ability. It competes with common native plants like Ipomoea imperati on white sand beaches in Hawaii (Rick Warshauer USGS-Biological Resources Division, Kilauea, Hawaii). It is one of several dominant species in Korean dunes (Kim, 2005).
Social Impact
Top of pageT. tetragonioides is a common garden plant around the world and has potential medicinal applications (Okuyama and Yamazaki, 1983; Cambie and Ferguson, 2003; APHIS, 2004; Grubben, 2004; Roskruge, 2011).
Risk and Impact Factors
Top of page- Invasive in its native range
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Is a habitat generalist
- Pioneering in disturbed areas
- Fast growing
- Has high reproductive potential
- Has propagules that can remain viable for more than one year
- Reproduces asexually
- Ecosystem change/ habitat alteration
- Modification of successional patterns
- Monoculture formation
- Threat to/ loss of native species
- Competition - monopolizing resources
- Competition - shading
- Competition - smothering
- Competition - strangling
- Pest and disease transmission
- Interaction with other invasive species
- Rapid growth
- Highly likely to be transported internationally accidentally
- Highly likely to be transported internationally deliberately
- Highly likely to be transported internationally illegally
- Difficult to identify/detect as a commodity contaminant
Uses
Top of pageEconomic value
T. tetragonoides is rarely cultivated as a crop for sale in large quantities, though it appears to have been imported into the United States from Israel for sale (APHIS, 2004), and may occasionally be sold in markets in Africa (Grubben, 2004). It is cultivated worldwide by gardeners as an easy to grow leafy green.
Social benefit
The fresh shoots of T. tetragonoides can be eaten raw, but leaves are generally cooked. 100 grams of T. tetragonoides is 94 g water, protein 1.5 g, fat 0.2 g, carbohydrate 2.5 g, Ca 58 mg, P 28 mg, Fe 0.8 mg, vitamin A 4400 IU, thiamin 0.04 mg, riboflavin 0.13 mg, niacin 0.5 mg, folate 15 µg and ascorbic acid 30 mg; it provides 59 kJ or 14 calories (Grubben, 2004). However, much of the calcium is present as oxalates and not available to the human body. Eating it raw has been discouraged because of the high saponin content (Grubben, 2004).
Uses List
Top of pageEnvironmental
- Wildlife habitat
General
- Sociocultural value
Human food and beverage
- Vegetable
Medicinal, pharmaceutical
- Traditional/folklore
Detection and Inspection
Top of pageA number of floras describe T. tetragonioides (Allan et al., 1961; Ohwi, 1965; Morris and Duretto, 2009; eFloras, 2013). It has been described in a dichotomous key for agricultural seeds available in German and translated into English (Brouwen and Stahlin, 1980).The distinctive horned 4-chambered seeds could be relatively easy to spot for someone familiar with the species.
Similarities to Other Species/Conditions
Top of pageThe climbing New Zealand spinach, Tetragoniatrigyna Banks and Sol. ex Hook.f., has smaller leaves and stems than T. tetragonioides (Roskruge, 2011).
Prevention and Control
Top of pageDue to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
Rapid response
Small populations have been controlled manually on Midway and Farrolon Islands, in the Northwest Hawaiian Islands (Forest Starr, pers. comm. Starr Environmental, Maui Hawaii, 2013)
Public awareness
T. tetragonioides is included in some websites documenting invasive species, such as PIER (2014).
Physical/mechanical control
Plants can be pulled up, although the effectiveness of this has not been reported.
Chemical control
Control with glyphosate may be effective based on other plants in Aizoaceae (DiTomaso, 2013).
Control by utilization
Not likely to be effective considering that growers of T. tetragonioides are most likely to harvest leaves and shoots for food rather than pulling whole plants.
Ecosystem restoration
Removal of T. tetragonioides coupled with planting of desirable plants and shrubs may prevent it establishing in the seedbank.
Gaps in Knowledge/Research Needs
Top of pageThe natural and human mediated spread of T. tetragonioides is not well documented: it was claimed as native in South America by some authors but it appears that botanists in South America now regard it as non-native. Genetic sequencing of wild populations and cultivated plants throughout its range could give insights into the spread of the plant. The claim that the plants in Europe were sourced via seeds that were brought back to England by Sir Joseph Banks could be investigated.
References
Top of pageAcevedo-Rodríguez P, Strong MT, 2012. Catalogue of Seed Plants of the West Indies. Washington, D.C., USA: Smithsonian Institution Scholarly Press, 1192 pp. [Smithsonian Contributions to Botany 98.]
Allan HH, 1961. Flora of New Zealand, I. Indigenous Tracheophyta. Wellington, New Zealand: Government Printer.
Alonso Paz EA, Bassagoda MJ, 2003. Survey of the flora and plant communities of Cerro Verde, Rocha, Uruguay. (Relevamiento de la flora y comunidades vegetales del Cerro Verde, Rocha, Uruguay.) Comunicaciones Botánicas, 127:1-20.
APHIS, 2004. Importation of New Zealand Spinach (Tetragonia tetragonioides) Palas. from Israel into the United States (A Qualitative, Pathway-initiated Risk Assessment). United States Department of Agriculture.
Barker S, 1970. Quondong Station, South Australia: a field context for applied rangeland research. Transactions of the Royal Society of South Australia, 94:179-190.
Campos JA, Herrera M, 2010. Analysis of the allochthonous flora of Bizkaia (Basque Country, Spain. (Análisis de la flora alóctona de Bizkaia (País Vasco, España).) Lazaroa, 30:7-33.
Cronk Q, 1983. Tetragonia tetragonoides (Pall.) Kuntze (Herbarium specimen). Royal Botanic Garden Edinburgh, Edinburgh. Edinburgh, UK: Royal Botanic Garden Edinburgh.
Déséglise A, 1881. Florula genevensis advena.
DiTomaso JM, Kyser GB, Oneto SR, Wilson RG, Orloff SB, Anderson LW, Wright SD, Roncoroni JA, Miller TL, Prather TS, Ransom C, Beck KG, Duncan C, Wilson KA, Mann JJ, 2013. Weed Control in Natural Areas in the Western United States. Davis, California, USA: Weed Research and Information Center, University of California, 544 pp.
eFloras, 2013. 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
GBIF, 2013. Global Biodiversity Information Facility. Global Biodiversity Information Facility (GBIF). http://data.gbif.org/species/
Halvorson WL, 1992. Alien Plants at Channel Islands National Park. Alien Plant Invasions in Native Ecosystems of Hawaii: Management and Research. Honolulu, Hawaii, USA: University of Hawaii Cooperative National Park Resources Studies Unit, 64-96.
Hanson CG, Mason JL, 1985. Bird seed aliens in Britain. Watsonia, 15:237-252.
Hara M, Tokunaga K, Kuboi T, 2008. Isolation of a drought-responsive alkaline a-galactosidase gene from New Zealand spinach. Plant Biotechnology, 25:497-501.
Harris G, 1998. Invasive New Zealand weeds: our native plant invaders. CalEPPC News, 6:8-9.
Herbst DR, Wagner WL, 1992. Alien plants on the northwestern Hawaiian Islands. Alien plant invasions in native ecosystems of Hawaii: management and research. Honolulu, Hawaii, USA: University of Hawaii Cooperative National Park Resources Studies Unit, 189-224.
Imada CT, 2012. Hawaiian Native and Naturalized Vascular Plants Checklist. Bishop Musem Technical Report. Honolulu, Hawaii, USA: Bishop Museum.
Kartesz JT, 2013. The biota of North America program (BONAP). North American Plant Atlas.
Kress WJ, Lace JH, Farr E, Daw Yin YK, 2003. A checklist of the trees, shrubs, herbs, and climbers of Myanmar. Washington, DC, USA: Department of Systematic Biology, Botany, National Museum of Natural History.
Macbride JF, 1937. Flora of Peru [ed. by Dahlgren, B. E.]. Chicago, USA: Field Museum of Natural History.
Meyer J-Y, 2009. Strategic Action Plan to Fight Against Invasive Introduced Plantes in Rapa Nui (Easter Island). (Plan de Accion Estratégico para Luchar Contra las Plantes Introducidas Invasoras en Rapa Nui (Isla De Pascua).) Ministry of Education, Higher Education and Research, Government of French Polynesia.
Missouri Botanical Garden, 2013. Tropicos database. St Louis, USA: Missouri Botanical Garden. http://www.tropicos.org/
Morong T, Britton NL, 1893. II - An Enumeration of the Plants Collected by Dr. Thomas Morong in Paraguay, 1888-1890. Annals of the New York Academy of Sciences, 7:45-280.
Morris DI, 2009. 101 Aizoacea. Flora of Tasmania [ed. by Duretto, M. F.]. Hobart, Australia: Tasmanian Herbarium, Tasmanian Museum and Art Gallery, 10. http://demo1.tmag.tas.gov.au/
Nicol J, Weedon J, Marsland K, 2009. Understorey vegetation monitoring of the Chowilla river red gum watering sites. South Australian Research and Development Institute (Aquatic Sciences). Adelaide, Australia: South Australian Research and Development Institute (Aquatic Sciences), 76 pp.
Ohwi J, 1965. Flora of Japan. Washington, DC, USA: Smithsonian Institute.
Okuyama E, Yamazaki M, 1983. The principles of Tetragonia tetragonoides having anti-ulcerogenic activity. II. Isolation and structure of cerebrosides. Chemical and Pharmaceutical Bulletin Japan, 31:2209-2219.
PIER, 2014. Pacific Islands Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html
Reiche C, Philippi F, 1898. Flora of Chile (Flora de Chile). Santiago de Chile, Chile: Imprenta Cervantes.
Robbins W, 1940. Alien plants growing without cultivation in California, 637:128 pp.
Soubeyran Y, 2008. Exotic species invading French overseas communities. Inventory of fixtures and recommendations. (Les espèces exotiques envahissantes dans les collectivités françaises d'outre-mer. Etat des lieux et recommandations.) Exotic species invading French overseas communities. French committee of the IUCN.
Starr F, Starr K, 2006. Oahu Offshore Islets Botanical Survey. Hawaii State Department of Land and Natural Resources and Offshore Islet Restoration Committee.
Sturtevant EL, 1919. State of New York-Department of Agriculture 27th Annual Report. New York, USA: JB Lyon Company, State Printers.
Tempére MG, 1920. Notes on some new and interesting plants of the Arcachonnaise region. (Notes sur quelques plantes nouvelles ou intéressantes de la région Arcachonnaise.) Actes de la Société Linnéenne De Bordeaux:27-28.
Ugarte E, Lira F, Fuentes N, Klotz S, 2011. Vascular alien flora, Chile. Check List, 7(3):365-382 pp.
USDA-ARS, 2013. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx
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
Vivrette NJ, 2004. Tetragonia (L.). In: Flora of North America: Magnoliophyta: Caryophyllidae, Part 1. Oxford University Press, 77 pp.
Weakley AS, 2012. Flora of the Southern and Mid-Atlantic States. Chapel Hill, North Carolina, USA: University of North Carolina, 1072 pp. URL: http://www.herbarium.unc.edu/flora.htm [Accessed 04 September 2013]
Wester L, 1992. Origin and distribution of adventive alien flowering plants in Hawaii. Honolulu, HI, USA: University of Hawaii, pp. In: Alien plant invasions in native ecosystems of Hawaii: management and research [ed. by Stone, C. P. \Smith, C. W. \Tunison, J. T.]. Honolulu, HI, USA: University of Hawaii, 99-154.
Wisflora, 2014. Wisflora: Checklist of the Vascular Plants of Wisconsin. Madison, Winsconsin, USA: Wisconsin State Herbarium, University of Wisconsin. http://www.botany.wisc.edu/wisflora/
Wishner C, 1998. Flora of the Santa Monica Mountains: synonymized checklist and index. Crossosoma, 23:3.
Wunderlin RP, Hansen BF, 2014. Atlas of Florida Vascular Plants. http://florida.plantatlas.usf.edu/
Zappi D, 2014. Aizoaceae. (Aizoaceae in Lista de Espécies da Flora do Brasil.) List of Species in the Flora of Brazil. Rio de Janeiro, Brazil: Botanical Garden of Rio de Janeiro.
Zomlefer WB, Giannasi DE, Judd WS, 2007. A floristic survey of National Park Service areas of Timucuan Ecological and Historic Preserve (including Fort Caroline National Memorial), Duval County, Florida. Journal of the Botanical Research Institute of Texas:1157-1178.
Zuloaga FO, Morrone O, 1996. I. Pteridophyta, Gymnosperamae and Angiospermae (Monocotyledoneae). (Catálogo de las Plantas Vasculares de la República Argentina I. Pteridophyta, Gymnosperamae y Angiospermae (Monocotyledoneae).) Catalogue of Vascular Plants of the Argentine Republic.
Distribution References
Allan HH, 1961. Flora of New Zealand, I. Indigenous Tracheophyta., Wellington, New Zealand: Government Printer.
Alonso Paz EA, Bassagoda MJ, 2003. Survey of the flora and plant communities of Cerro Verde, Rocha, Uruguay. (Relevamiento de la flora y comunidades vegetales del Cerro Verde, Rocha, Uruguay). In: Comunicaciones Botánicas, 127 1-20.
APHIS, 2004. Importation of New Zealand Spinach (Tetragonia tetragonioides) Palas. from Israel into the United States (A Qualitative, Pathway-initiated Risk Assessment)., United States Department of Agriculture.
Barker S, 1970. Quondong Station, South Australia: a field context for applied rangeland research. In: Transactions of the Royal Society of South Australia, 94 179-190.
CABI Data Mining, Undated. CAB Abstracts Data Mining.,
CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
Campos JA, Herrera M, 2010. Analysis of the allochthonous flora of Bizkaia (Basque Country, Spain). (Análisis de la flora alóctona de Bizkaia (País Vasco, España)). In: Lazaroa, 30 7-33.
Cronk Q, 1983. (Tetragonia tetragonoides (Pall.) Kuntze (Herbarium specimen)). In: Royal Botanic Garden Edinburgh, Edinburgh, UK: Royal Botanic Garden Edinburgh.
Déséglise A, 1881. Florula genevensis advena.,
eFloras, 2013. eFloras., St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria .
GBIF, 2013. Global Biodiversity Information Facility. http://www.gbif.org/species
Halvorson WL, 1992. Alien Plants at Channel Islands National Park. In: Alien Plant Invasions in Native Ecosystems of Hawaii: Management and Research, Honolulu, Hawaii, USA: University of Hawaii Cooperative National Park Resources Studies Unit. 64-96.
Hanson C G, Mason J L, 1985. Bird seed aliens in Britain. Watsonia. 237-252.
Harris G, 1998. Invasive New Zealand weeds: our native plant invaders. In: CalEPPC News, 6 8-9.
Herbst DR, Wagner WL, 1992. Alien plants on the northwestern Hawaiian Islands. In: Alien plant invasions in native ecosystems of Hawaii: management and research, Honolulu, Hawaii, USA: University of Hawaii Cooperative National Park Resources Studies Unit. 189-224.
Imada CT, 2012. Hawaiian Native and Naturalized Vascular Plants Checklist. In: Bishop Musem Technical Report, Honolulu, Hawaii, USA: Bishop Museum.
Kartesz JT, 2013. The biota of North America program (BONAP). In: North American Plant Atlas,
Kress WJ, Lace JH, Farr E, Daw Yin YK, 2003. A checklist of the trees, shrubs, herbs, and climbers of Myanmar., Washington, DC, USA: Department of Systematic Biology, Botany, National Museum of Natural History.
Macbride JF, 1937. Flora of Peru., [ed. by Dahlgren BE]. Chicago, USA: Field Museum of Natural History.
Meyer J-Y, 2009. Strategic Action Plan to Fight Against Invasive Introduced Plantes in Rapa Nui (Easter Island). (Plan de Accion Estratégico para Luchar Contra las Plantes Introducidas Invasoras en Rapa Nui (Isla De Pascua))., Ministry of Education, Higher Education and Research, Government of French Polynesia.
Missouri Botanical Garden, 2013. Tropicos database., St Louis, USA: Missouri Botanical Garden. http://www.tropicos.org/
Morong T, Britton NL, 1893. II - An Enumeration of the Plants Collected by Dr. Thomas Morong in Paraguay, 1888-1890. In: Annals of the New York Academy of Sciences, 7 45-280.
Morris DI, 2009. 101 Aizoacea. In: Flora of Tasmania, Hobart, Australia: Tasmanian Herbarium, Tasmanian Museum and Art Gallery. 10. http://demo1.tmag.tas.gov.au/
Nicol J, Weedon J, Marsland K, 2009. Understorey vegetation monitoring of the Chowilla river red gum watering sites. In: South Australian Research and Development Institute (Aquatic Sciences), Adelaide, Australia: South Australian Research and Development Institute South Australian Research and Development Institute (Aquatic Sciences). 76 pp.
Ohwi J, 1965. Flora of Japan., Washington, DC, USA: Smithsonian Institute.
PIER, 2014. Pacific Islands Ecosystems at Risk., Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html
Reiche C, Philippi F, 1898. Flora of Chile. (Flora de Chile)., Santiago de Chile, Chile: Imprenta Cervantes.
Robbins W, 1940. Alien plants growing without cultivation in California., 637 128 pp.
Soubeyran Y, 2008. Exotic species invading French overseas communities. Inventory of fixtures and recommendations. (Les espèces exotiques envahissantes dans les collectivités françaises d'outre-mer. Etat des lieux et recommandations). In: Exotic species invading French overseas communities, French committee of the IUCN.
Starr F, Starr K, 2006. Oahu Offshore Islets Botanical Survey., Hawaii State Department of Land and Natural Resources and Offshore Islet Restoration Committee.
Tempére MG, 1920. Notes on some new and interesting plants of the Arcachonnaise region. (Notes sur quelques plantes nouvelles ou intéressantes de la région Arcachonnaise). In: Actes de la Société Linnéenne De Bordeaux, 27-28.
Ugarte E, Lira F, Fuentes N, Klotz S, 2011. Vascular alien flora, Chile. In: Check List, 7 (3) 365-382.
USDA-ARS, 2013. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx
Weakley AS, 2012. Flora of the Southern and Mid-Atlantic States., Chapel Hill, North Carolina, USA: University of North Carolina. 1072 pp. http://www.herbarium.unc.edu/flora.htm
Wester L, 1992. Origin and distribution of adventive alien flowering plants in Hawaii. In: Alien plant invasions in native ecosystems of Hawaii: management and research, [ed. by Stone CP, Smith CW, Tunison JT]. Honolulu, HI, USA: University of Hawaii. 99-154.
Wisflora, 2014. Wisflora: Checklist of the Vascular Plants of Wisconsin., Madison, Winsconsin, USA: Wisconsin State Herbarium, University of Wisconsin. http://www.botany.wisc.edu/wisflora/
Wishner C, 1998. Flora of the Santa Monica Mountains: synonymized checklist and index. In: Crossosoma, 23 3.
Wunderlin RP, Hansen BF, 2014. Atlas of Florida Vascular Plants., http://florida.plantatlas.usf.edu/
Zappi D, 2014. Aizoaceae. List of Species in the Flora of Brazil. (Aizoaceae in Lista de Espécies da Flora do Brasil)., Rio de Janeiro, Brazil: Botanical Garden of Rio de Janeiro.
Zomlefer WB, Giannasi DE, Judd WS, 2007. A floristic survey of National Park Service areas of Timucuan Ecological and Historic Preserve (including Fort Caroline National Memorial), Duval County, Florida. In: Journal of the Botanical Research Institute of Texas, 1157-1178.
Zuloaga FO, Morrone O, 1996. I. Pteridophyta, Gymnosperamae and Angiospermae (Monocotyledoneae). (Catálogo de las Plantas Vasculares de la República Argentina I. Pteridophyta, Gymnosperamae y Angiospermae (Monocotyledoneae)). In: Catalogue of Vascular Plants of the,
Contributors
Top of page17/03/14 Original text by:
Christopher Buddenhagen, Florida State University, USA
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