- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Air Temperature
- Soil Tolerances
- Natural enemies
- Notes on Natural Enemies
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Threatened Species
- Risk and Impact Factors
- Similarities to Other Species/Conditions
- Prevention and Control
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Spartina densiflora Brongn.
Other Scientific Names
- Chauvinia chilensis Steud.
- Spartina juncea var. laxiflora St.-Yves
- Spartina juncea var. montevidense (Arechav.) St.-Yves
- Spartina montevidensis Arechav.
- Spartina patagonica Speg.
International Common Names
- English: Austral cordgrass; Chilean cordgrass; denseflower cordgrass; denseflowered cord grass; Humboldt Bay cordgrass; Humboldt cordgrass
Local Common Names
- Argentina: espartillo; esparto
- Colombia: llinto
- Portugal: capim-praturá
- Uruguay: espartillo; esparto
- SPTDE (Spartina densiflora)
Summary of InvasivenessTop of page
S. densiflora, known by the common name denseflower cordgrass, is native to the coast of southern South America and has been introduced to the west coast of North America and parts of the Mediterranean coast where it is a noxious weed in some areas. S. densiflora is an extremely aggressive species that outcompetes native plants and alters both the physical structure and biological composition of tidal marshes and mudflats. The rapid growth rate and lack of dormancy period make S. densiflora a threat to local biodiversity where it is invasive. The California Invasive Plant Council classifies its potential impact on native ecosystems as ‘High – Alert’, and it is listed as Class ‘A’ Noxious weed and a Quarantine weed in Oregon, and a Class ‘A’ Noxious weed in Washington and comes under the Wetland and aquatic weed quarantine (USDA-NRCS, 2016).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Monocotyledonae
- Order: Cyperales
- Family: Poaceae
- Genus: Spartina
- Species: Spartina densiflora
Notes on Taxonomy and NomenclatureTop of page
Spartina, commonly known as cordgrass or cord-grass, is a genus of plants in the grass family. The genus consists of 15 species, including Spartina densiflora Brongn. (USDA-ARS, 2016). S. densiflora is presently one of the three most widely distributed species of the genus Spartina, along with S. alterniflora and S. anglica (Bortolus, 2006).
The first known samples of S. densiflora were collected between 1822 and 1825 in Concepcion, Chile. The species were described by Adolphe Brongniart and published in Voyage Autour du Monde 2 in 1829 (Brongniart, 1829).
In the decades after the publication of S. densiflora in 1829, more halophyte species were described in Chile (Chauvinia chilensis; Steudel, 1855), Uruguay (S. montevidensis; Arechavaleta, 1894) and Argentina (S. patagonica; Spegazzini, 1897), which are now recognized as phenotypic variations of S. densiflora (Parodi, 1919, 1967; Mobberley, 1956). But some of these names are still used incorrectly in the literature, S. montevidensis being the most common (Bortolus, 2006). Significant genetic variation is indicated by the large number of forms, varieties, subvarieties and subspecies described by Missouri Botanical Garden (2016).
The generic name Spartina is derived from the Greek word ‘spartina’ (a rope made of straw, spartium), referring to the fibrous leaves. The Latin epithet densiflora means ‘dense flower’.
DescriptionTop of page
The following is from Barkworth (2016):
S. densiflora is a perennial grass that grows in erect, tufted clumps of slender stems that can reach 1.5 m tall. The plant generally lacks rhizomes but when present, they are short, to 10 mm thick. Culms: 27-150 cm, forming large clumps, indurate, usually with short extravaginal shoots appressed to the culms. Sheaths: glabrous, lower sheaths smooth, indurate and shining, upper sheaths dull and somewhat striate; Ligules: 1-2 mm; Blades: The blade-like narrow leaves are gray-green and rolled inward, especially when new. Leaves are 12-43 cm long, 3-8 mm wide, abaxial surfaces glabrous, adaxial surfaces and margins scabrous. Inflorescence: a panicle with dense, compact colorless flowers,10-30 cm long, 4-8 mm wide, sinuous in outline, often twisted, with 2-15 branches; Branches: 1-11 cm long, longer branches narrower than the shorter branches, all branches tightly appressed, moderately imbricate, axes not prolonged beyond the distal spikelets, with 10-30 spikelets. Spikelets: 8-14 mm, tipped with bristles; Glumes: glabrous or sparsely hispidulous, keels hispidulous, margins sparsely hispidulous; Lower glumes 4-7 mm, usually obtuse; Upper glumes 8-14 mm, 1-veined, usually acuminate; Lemmas: minutely hispidulous, keels glabrous proximally, hispidulous distally, apices acuminate to obtuse; Paleas: acuminate, keels glabrous basally, hispidulous distally; Anthers: 3-5 mm.
Plant TypeTop of page Grass / sedge
DistributionTop of page
S. densiflora is native to the coastline of southern South America from southern Brazil (Parana, Rio Grande do Sul, Santa Catarina and Sao Paulo), Uruguay and Argentina (Buenos Aires, Chaco, Chubut, Cordoba, Federal District, La Pampa, Rio Negro, San Luis, Santa Cruz and Santa Fe) on the east coast and Chile on the west coast. The species is introduced and naturalized in North America (Oregon, Washington, Texas and California), Morocco and Spain (USDA-ARS, 2016).
According to the USDA-NRCS (2016)S. densiflora is also present in British Columbia, Canada. And unlike USDA-ARS (2016), it does not report on presence of S. densiflora in Texas, USA. DAISIE (2016) reports that S. densiflora is also established in Portugal.
Distribution TableTop of page
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/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Morocco||Present||Introduced||1978||Invasive||ISSG, 2016; USDA-ARS, 2016||Merja Zerga Lagoon|
|Canada||Present||Present based on regional distribution.|
|-British Columbia||Present||Introduced||USDA-NRCS, 2016|
|USA||Present, few occurrences||EPPO, 2014|
|-California||Present||Introduced||late 1800s||Invasive||ISSG, 2016; USDA-ARS, 2016; USDA-NRCS, 2016||Humboldt Bay, San Francisco Bay (Estuary)|
|-Oregon||Present||Introduced||USDA-ARS, 2016; USDA-NRCS, 2016|
|-Texas||Present||Introduced||Barkworth, 2016; USDA-ARS, 2016|
|-Washington||Present||Introduced||2001||Invasive||ISSG, 2016; USDA-ARS, 2016; USDA-NRCS, 2016||Grays Harbor, Race Lagoon|
|Argentina||Present||Native||USDA-ARS, 2016||Buenos Aires, Chaco, Chubut, Cordoba, Federal District, La Pampa, Rio Negro, San Luis, Santa Cruz and Santa Fe|
|Brazil||Present||Present based on regional distribution.|
|-Rio Grande do Sul||Present||Native||USDA-ARS, 2016|
|-Santa Catarina||Present||Native||USDA-ARS, 2016|
|-Sao Paulo||Present||Native||USDA-ARS, 2016|
|Venezuela||Present||Missouri Botanical Garden, 2016|
|Spain||Present||Introduced||Invasive||DAISIE, 2016; ISSG, 2016; USDA-ARS, 2016||Odiel River, Tinto River; Established|
History of Introduction and SpreadTop of page
S. densiflora was first introduced to USA, to Humboldt Bay, California possibly during the 1800s by the lumber trade, and is currently spreading dramatically at several other invaded sites (Clifford, 2002). In 1976, S. densiflora was transplanted from Humboldt Bay to the San Francisco Estuary as part of a marsh restoration project adjacent to Corte Madera Creek in Marin County. S. densiflora has now spread beyond the original marsh plantings to the entire two mile length of Corte Madera Creek and is spreading beyond the creek into San Francisco Bay (California State Coastal Conservancy, 2016), where it has reappeared in areas where it was believed to be eradicated (Smith et al., 2001; Ayres et al., 2004). There are reports that suggest that the progress of S. densiflora invasions in the northeast Pacific has accelerated during the last few years in comparison with previous decades (e.g. Grossinger et al., 1998; Pickart, 2001; Smith et al., 2001). The latest invasion was recorded in 2001 at Grays Harbor (Pfauth et al., 2003). Considering its wide distribution within South America, southwest Europe and along the west coast of North Africa, the invasion of Grays Harbor suggests that this species is able to invade most estuarine environments along the west coast of the USA and Canada (Bortolus, 2006).
Only recently was S. densiflora reported in Western Europe and North Africa. The first report locating S. densiflora outside the Americas was from Spain (Tutin et al., 1980; Nieva, 1996). The introduction of S. densiflora to the European continent is considered accidental, and probably resulted from the lumber trade between South America and Spain (Nieva et al., 2001).
In recent decades, S. densiflora has aggressively spread over the Gulf of Cadiz, from Cape San Vicente, Portugal, to Gibraltar, Spain. According to Nieva et al. (2002)S. densiflora has probably already been introduced into Galicia, north-west Spain, as well.
Although S. densiflora is found at 51°33′S in Argentina, it has been suggested that this species may have physiological constraints limiting its winter production and therefore the extent of its distribution and abundance at higher latitudes in Europe (Bortolus, 2006).
S. densiflora was transported from Spain to Morocco in 1978. The introduction of the species could have been the result of a mislabelling of species for a botanical garden, or via solid ballast. S. densiflora is now established in Morocco and is reported as an invasive species (Bortolus, 2006).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|California||South America||late 1800s||Timber trade (pathway cause)||Yes||California State Coastal Conservancy (2016)||Introduced to Humboldt Bay, California, possibly during the nineteenth century by lumber trade|
|Morocco||Spain||1978||Botanical gardens and zoos (pathway cause)
Timber trade (pathway cause)
|Yes||Bortolus (2006)||Introduction could have been the result of a mislabel of species for a botanical garden, or via solid ballast|
|Spain||South America||1800s-1978||Timber trade (pathway cause)||Yes||Nieva et al. (2001)||Introduction to the European continent is considered accidental. It probably resulted from the lumber trade between South America and Spain|
Risk of IntroductionTop of page
S. densiflora has been previously spread via the lumber trade, and occurence of S. densiflora populations near oyster growing areas in Washington (Tomales Bay and Puget Sound), USA, suggests human aided spread via transport of oyster spat. Seeds can also travel on tidal currents (California Invasive Plants Council, 2016).
S. densiflora is listed in the California Invasive Plant Inventory. The California Invasive Plant Council classifies the potential impact on native ecosystems as ‘High – Alert’. S. densiflora is rated as ‘High’ based on an evaluation which considers ecological impacts, invasion potential and ecological distribution (California Invasive Plants Council, 2016; USDA-NRCS, 2016).
S. densiflora is listed as a Class ‘A’ Noxious weed and Quarantine weed in the US state of Oregon. Also in the state of Washington, S. densiflora is listed as a Class ‘A’ Noxious weed and comes under the Wetland and aquatic weed quarantine (USDA-NRCS, 2016).
HabitatTop of page
S. densiflora is capable of invading a broad spectrum of habitats: from brackish to hypersaline (Nieva, 1996; Kittelson and Boyd, 1997), from intertidal to strictly terrestrial (Mobberley, 1956; Clifford, 2002; Nieva et al., 2002), and from subtropical to austral. Soils that can support S. densiflora vary from well drained and oxygenated (Vicari et al., 2002), to muddy and anoxic (Cabrera and Zardini, 1978; Nicora and Rugolo de Agrasar, 1987). S. densiflora inhabits protected estuaries and bays as well as open coastlines where it successfully populates cobble beaches, hard (volcanic stones) and soft (sand, clay and limestone) rocky shores. In marshes it usually colonizes the middle and high zones (Bortolus, 2005, 2006).
In freshwater and brackish environments it shows the fastest colonization and expansion rates, probably because of the salinity stress condition in marine coastal areas (Castillo et al., 2005).
Habitat ListTop of page
|Terrestrial ‑ Natural / Semi-natural||Riverbanks||Present, no further details|
|Wetlands||Present, no further details|
|Rocky areas / lava flows||Present, no further details|
|Coastal areas||Present, no further details|
|Mud flats||Present, no further details|
|Intertidal zone||Present, no further details|
|Salt marshes||Present, no further details|
Biology and EcologyTop of page
The chromosome number is 2n = 60 (Barkworth, 2016).
S. densiflora is one of several species which produces hybrids with Spartina alterniflora (Parodi, 1919; Cabrera, 1970; Bortolus, 2001), which makes it a dangerous threat to native plant communities (Bortolus, 2006).
S. densiflora can reproduce both sexually and asexually, by vegetative tiller production and seed germination, but the asexual role is very small in comparison (Nieva et al., 2001). The flowering period of S. densiflora in its native area is from November to May. In Spain, the flowering period is from June to December and in the USA, from April to July (Bortolus, 2006). Within its native range, S. densiflora does not exhibit any clear dormancy period during the year, which gives it a competitive advantage over other species (Bortolus, 2006).
Populations of S. densiflora demonstrate a pattern of sequential development of identical growth units derived from tillers. S. densiflora forms live shoots to support an annual die-back phase (Castillo and Figueroa, 2009). In the USA, spring to early autumn is the time for rapid growth and development of S. densiflora. When it blooms, from April to July, it experiences the die-back phase with the loss of flowering culms (Noxious Weed Control Board, 2010).
Physiology and Phenology
S. densiflora is a perennial (EOL, 2016), and shows great phenotypic plasticity. The tiller density, biomass production, flowering period and phenotype of S. densiflora are highly variable between regions. It may vary between a tall form of 1.5 m and a short form of few centimetres tall. The density of inflorescences, spikes, spikelets, and their shape and size also vary importantly among plants (ISSG, 2016).
S. densiflora has a C4 metabolism and is able to develop different growing strategies to enable it to cope better with local environmental variables and micro-habitat conditions (Kittelson and Boyd, 1997; Nieva et al., 2005). S. densiflora shows a faster ramet turnover in exposed low marsh areas than in high marsh areas. This results in increased mechanical resistance to wave action, which favours its success in this zone (Nieva et al., 2005; Bortolus, 2006).
Population Size and Structure
S. densiflora forms large, often dense colonies consisting of tight clumps of herbage that grow quickly (EOL, 2016). In old S. densiflora grasslands, rhizomes of different ramets living together are usually packed tightly in a complex, multidirectional phalanx-growing (clumping ramets) pattern. Phalanx-growing salt-marsh plants are often considered more successful competitors than guerrilla-growing (spreading) plants, because although they grow slowly, they colonize available above- and below-ground space with closely packed modules. Guerrilla-growing plants are capable of rapid lateral expansion but with loosely packed modules. However, S. densiflora seems to be able to combine the best of both growing strategies by overlapping guerrilla clones growing in different directions, generating multi-clone phalanx-growing modules (Bortolus, 2006).
In the high marsh of Buenos Aires, Argentina, S. densiflora grows together with Salicornia ambigua (= Sarcocornia perennis), Distichlis spicata, Limonium sp., Cressa sp. and Juncus acutus (Vervoost, 1967; Cabrera, 1970; Iribarne, 2001). In this region S. densiflora is also found sharing the intertidal zone with Spartina longispica and Spartina alterniflora, which occupy the high, middle and low intertidal zones respectively (Bortolus, 2006).
S. densiflora grows best in freshwater and brackish marshes but it tolerates strongly saline habitats as well. The species has high Arsenic (As) tolerance, associated with the capacity to accumulate Arsenic in its roots and largely avoid its transport to the leaves (Mateos-Naranjo et al., 2012).
Soils that can support S. densiflora vary from well drained and oxygenated (Vicari et al., 2002), to muddy and anoxic (Cabrera and Zardini, 1978; Nicora and Rugolo de Agrasar, 1987). According to Calflora (2016)S. densiflora grows on soils with a pH of 5.2-8.6.
Several sources report different elevations for S. densiflora. According to Calflora (2016)S. densiflora occurs on elevations between 0-80 m. The EOL (2016) reports on an elevation in between 1-130 m. Lastly, Baird and Thieret (2016) state that S. densiflora only occurs below 10 m.
ClimateTop of page
|BS - Steppe climate||Preferred||> 430mm and < 860mm annual precipitation|
|BW - Desert climate||Preferred||< 430mm annual precipitation|
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Preferred||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
Air TemperatureTop of page
|Parameter||Lower limit||Upper limit|
|Mean annual temperature (ºC)||4||27|
RainfallTop of page
|Parameter||Lower limit||Upper limit||Description|
|Dry season duration||4||7||number of consecutive months with <40 mm rainfall|
|Mean annual rainfall||482||1092||mm; lower/upper limits|
Soil TolerancesTop of page
Special soil tolerances
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
Notes on Natural EnemiesTop of page
There has been a case recorded of an aggressive spike infestation by Claviceps fungi in Argentina and Chile (Ringuelet, 1935; Fischer et al., 2005). This fungus might have a negative impact on the sexual reproductive success of S. densiflora. But until now there have been no studies evaluating the effects of this fungus on the production and reproduction of S. densiflora (Bortolus, 2006).
Pathway CausesTop of page
|Botanical gardens and zoos||Its introduction could have been the result of the mislabelling of a species for a botanical garden||Yes||Bortolus, 2006|
|Fisheries||Possible spread via human aided transport of Oyster spat in Washington, USA||Yes||California Invasive Plant Council, 2016|
|Habitat restoration and improvement||Transplanted from Humboldt Bay to the San Francisco Estuary as part of a marsh restoration project||Yes||California State Coastal Conservancy, 2016|
|Timber trade||Probably introduced from South America to the USA and the Mediterranean by lumber trade||Yes||California State Coastal Conservancy, 2016; Nieva et al., 2001|
Pathway VectorsTop of page
|Bulk freight or cargo||Probably introduced from South America to the USA and the Mediterranean by lumber trade||Yes||California State Coastal Conservancy, 2016; Nieva et al., 2001|
|Germplasm||Introduction could have been the result of the mislabelling of a species for a botanical garden||Yes||Bortolus, 2006|
|Water||Dispersal via tidal currents is feasible||Yes||California Invasive Plant Council, 2016|
Impact SummaryTop of page
|Environment (generally)||Positive and negative|
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Rallus longirostris obsoletus (California clapper rail)||USA ESA listing as endangered species USA ESA listing as endangered species||California||Competition - monopolizing resources||California State Coastal Conservancy, 2016|
|Reithrodontomys raviventris (salt-marsh harvest mouse)||EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered species||California||Competition - monopolizing resources||California State Coastal Conservancy, 2016|
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Highly adaptable to different environments
- Is a habitat generalist
- Fast growing
- Has high reproductive potential
- Reproduces asexually
- Ecosystem change/ habitat alteration
- Modification of hydrology
- Monoculture formation
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Competition - monopolizing resources
- Rapid growth
UsesTop of page
There are several reports on the ability of S. densiflora to accumulate toxic heavy metals and organic pollutants from the environment, which makes it possibly useful for phytoremediation.
Andrades-Moreno et al. (2014) report that S. densiflora is growing on sediments with high concentrations of heavy metals in the salt marshes of the joint estuary of Tinto and Odiel rivers, in the south-west of Spain. According to Andrades-Moreno et al. (2014), these marshes are one of the most polluted areas by heavy metals in the world.
S. densiflora demonstrated a remarkable tolerance to phenanthrene stress (phenanthrene is a polycyclic aromatic hydrocarbon). S. densiflora increased phenanthrene degradation in soils. It exhibited higher rates of disappearance of phenanthrene than other species (Redondo-Gómez et al., 2011).
S. densiflora also presents a high tolerance to arsenic (As)-induced stress. The As-tolerance of the species is associated with the capacity to accumulate As in its roots and largely avoid its transport to the leaves. This fact indicates that this species could be useful for arsenic phytostabilization purposes (Mateos-Naranjo et al., 2012).
S. densiflora offers a variety of resources for birds. They use it as nest-building material, like the long-winged harrier (Circus buffoni). They find protection from predators, like the Argentinean common rail (Pardirralus sanguinolentus) and the wren-spinetail (Spartonoica maluroides), or a place to find prey (C. buffoni), and at least one of them (Rhea americana; Herrera, 2000; Isacch et al., 2001) eats S. densiflora. The edges of the Spartina marsh are important for many birds as shelter against strong cold winds (e.g. for ducks, swans and shorebirds; Bortolus et al., 1998; Martinez and Ferrero, 2001).
Several mammals find in S. densiflora marshes a last refuge in regions dominated by urban areas. The Spartina marshes, patchily distributed along the coast, serve as corridors for many mammals providing hunting, feeding and breeding areas (Bortolus, 2006).
Similarities to Other Species/ConditionsTop of page
Similar species according to ISSG (2016) are Scirpus maritimus, Spartina alterniflora, Spartina anglica, Spartina foliosa, Spartina maritima, Spartina patens and Triglochin maritima.
In California, USA, S. densiflora has often been mistaken for S. foliosa, from which it differs in its indurate culms, narrow, inrolled leaves and tendency to grow among Salicornia in the upper intertidal zone or in open mud (Barkworth, 2016).
Prevention and ControlTop of page
For small populations of S. densiflora manual or mechanical extraction, burning or covering are management techniques that can work effectively. Mechanical removal of S. densiflora has proven to be more effective than hand removal methods. The covering method involves draping fabric that seals out sunlight and kills off the S. densiflora underneath. Studies have shown that mowing can be an effective method for control on a large scale (Kittelson and Milton, 1997; ISSG, 2016).
Flooding has been described as one of the most important environmental factors determining the distribution of salt marsh plants. Abbas et al. (2012) have studied the impact of different water levels on the germination and establishment of S. densiflora, and the germination rate tended to decrease gradually with depth. The results show that artificial inundation of invaded marshes to a water depth of 8 cm would prevent the establishment of S. densiflora from the seed bank (Abbas et al., 2012).
The aquatic herbicides imazapyr or glyphosate with a surfactant are applied to S. densiflora to eradicate and control populations. These two herbicides are currently the only aquatic chemicals approved for use in estuarine environments in the state of California by the US Environmental Protection Agency and the California Department of Pesticide Regulation (CDPR) (ISSG, 2016).
ReferencesTop of page
Abbas AM; Rubio-Casal AE; Cires Ade; Figueroa ME; Lambert AM; Castillo JM, 2012. Effects of flooding on germination and establishment of the invasive cordgrass Spartina densiflora. Weed Research (Oxford), 52(3):269-276.
Andrades-Moreno L; Castillo Idel; Parra R; Doukkali B; Redondo-Gómez S; Pérez-Palacios P; Caviedes MA; Pajuelo E; Rodríguez-Llorente ID, 2014. Prospecting metal-resistant plant-growth promoting rhizobacteria for rhizoremediation of metal contaminated estuaries using Spartina densiflora. Environmental Science and Pollution Research, 21(5):3713-3721.
Arechavaleta J, 1894. Uruguayan grasses (Las gramineas uruguayas). Anales del Museo Nacional de Montevideo, 1:376-382.
Ayres DR; Smith DL; Zaremba K; Klohr S; Strong DR, 2004. Spread of exotic cordgrasses and hybrids (Spartina spp.) in the tidal marshes of San Francisco Bay, California, USA. Biological Invasions, 6:221-231.
Baird JR; Thieret JW, 2016. Spartina densiflora. Jepson eFlora [ed. by Jepson Flora Project]. http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?tid=45043
Barkworth ME, 2016. Manual of Grasses for North America. Utah, USA: Utah State University. http://herbarium.usu.edu/webmanual
Bortolus A, 2001. Ecological interactions between the digger crab Chasmagnathus granulata and espartillares of Spartina densiflora, mesofauna associated and characteristics of the substrate in Mar Chiquita coastal lagoon (Buenos Aires, Argentina). PhD Thesis (Interacciones ecologicas entre el cangrejo cavador Chasmagnathus granulata y los espartillares de Spartina densiflora, mesofauna asociada y caracteristicas del sustrato en la laguna costera Mar Chiquita (Buenos Aires, Argentina). PhD Thesis). Buenos Aires, Argentina: Universidad Nacional de Mar del Plata.
Bortolus A, 2005. Finding a lost world in the mythic Patagonia: setting physiographic and ecological baseline information of the austral salt marshes of South America. In: Argentine Conference Botanica. Rosario, Argentina.
Bortolus A; Iribarne O; Martinez MM, 1998. Relationship between waterfowl and the seagrass Ruppia maritima in a Southwestern Atlantic coastal lagoon. Estuaries, 21:710-717.
Brongniart MA, 1829. Phanerogamie. Voyage autour du monde sur la corvette La Coquille, pendant les anees, 2(14):1822-1825.
Cabrera AL, 1970. Part II. Gramineas (Parte II. Gramineas). Flora de la Provincia de Buenos Aires. Buenos Aires, Argentina: INTA.
Cabrera AL; Zardini EM, 1978. Manual of flora around Buenos Aires (Manual de la flora de los alrededores de Buenos Aires), second edition. Buenos Aires, Argentina: Editorial Buenos Aires.
Calflora, 2016. California flora database. Berkeley, California, USA. http://www.calflora.org/
California Invasive Plant Council, 2016. Spartina densiflora (dense-flowered cordgrass). http://www.cal-ipc.org/ip/management/plant_profiles/Spartina_densiflora.php
California State Coastal Conservancy, 2016. San Francisco Estuary Invasive Spartina Project. Oakland, California, USA: California State Coastal Conservancy. http://www.spartina.org
Castillo JM; Figueroa E, 2009. Effects of abiotic factors on the life span of the invasive cordgrass Spartina densiflora and the native Spartina maritima at low salt marshes: Changes in life span of cordgrasses. Aquatic Ecology, 43(1):51-60.
Castillo JM; Rubio-Casal AE; Redondo S; Álvarez-López AA; Luque T; Luque C; Nieva FJ; Castellanos EM; Figueroa ME, 2005. Short-term responses to salinity of an invasive cordgrass. Biological Invasions, 7(1):29-35.
Clifford M, 2002. Dense-flowered cordgrass (Spartina densiflora) in Humboldt Bay. Summary and literature review. A report for the California State Coastal Conservancy, Oakland. Oakland, California: California State Coastal Conservancy.
Daehler CC; Strong DR, 1996. Status, prediction and prevention of introduced cordgrass Spartina spp. invasions in Pacific estuaries, USA. In: Biological Conservation [ed. by Carey JR, Moyle P, Rejmánek M, Vermeij GJ], 51-58.
DAISIE, 2016. Delivering Alien Invasive Species Inventories for Europe. European Invasive Alien Species Gateway. www.europe-aliens.org/default.do
Encyclopedia of Life, 2016. Encyclopedia of Life. http://www.eol.org
EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm
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09/06/2016 Original text by:
Ymkje van de Witte, Consultant, UK
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