Ulva (Enteromorpha) flexuosa subsp. flexuosa and flexuosa subsp. paradoxa (Wolfen ex Roth) J. Agardh, 1883

Common Name: Green alga, grass kelp

Synonyms and Other Names:

Hollow green weed, Conferva paradoxa, Enteromorpha clathrata var. hopkirkii, Enteromorpha intestinalis var. tubulosa, Enteromorpha erecta, Enteromorpha flexuosa subsp. paradoxa, Enteromorpha hopkirkii, Enteromorpha paradoxa, Enteromorpha paradoxa var. typica, Enteromorpha plumose, Enteromorpha prolifera var. tubulosa , Enteromorpha tubulosa, Scytosiphon erectus, Ulva paradoxa 



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Identification: Ulva flexuosa has a smooth, tubular thallus that is linear and uniform in width, with numerous branches in the lower third. It can have one to several fronds. Ulva flexuosa ranges from translucent dark to light green, with chloroplasts taking up part to most of the outer region of its cells. It has a very small rhizoidal region (Woolcott and King 1999). In general, its subspecies are differentiated by their degree of branching: U. flexuosa subsp. flexuosa tends to be unbranched, while Ulva flexuosa subsp. paradoxa has branched filaments and is typically more slender (Sahoo et al. 2003). Additional characteristics are summarized below.

Ulva flexuosa subsp. flexuosa is yellow-green with tubular fronds, cylindrical stalks, and a blade expanding above the stalk which ends in a rounded tip. It is attached to the substrate by a small, round basal disc (Sahoo et al. 2003). Ulva flexuosa subsp. flexuosa has sparse branching, one to two pyrenoids (for carbon dioxide fixation), and quadrangular or rectangular cells arranged in approximately regular rows (Mares et al. 2011).

Ulva flexuosa subsp. paradoxa has an intensely branched stalk with rectangular cells arranged in long rows. These rows are somewhat unordered in the central parts of broader thalli, thinning to a single row of cells at the ends of branchlets (Mares et al. 2011).


Size: < 30 cm long and 0.5-1.0 cm in diameter (Sahoo et al. 2003)


Native Range: Unknown. This species is widespread. It occurs around the world in inland and/or coastal waters of Latin America and the Caribbean, Asia, Europe, Australia, the U.S.A., Mexico, and various islands in the Pacific Ocean (Kowalski 1975, Trono 1975, Webber 1975, Riouall 1976, Sivalingam 1977, Morton 1978, Vaughan 1978, Bird and McIntosh 1979, Lobel and Ogden 1981, Kies and Dworsky 1982, Rodriguez de Rios and Lobo 1984, Grant and Prasad 1985, Guner et al. 1985, Ho 1987, Hadi et al. 1989, Khotimchenko 1993, Beach et al. 1995, Martinez-Murillo and Aladro-Lubel 1996, Magnusson 1997, Schories et al. 1997, Fernandez et al. 1998, Woolcott and King 1999, Hodgson and McDermid 2000, Tabudravu et al. 2002, Matik-Skoko et al. 2004, Lourenco et al. 2006, Sahoo et al. 2006).

Ulva flexuosa is considered a cosmopolitan species that has a worldwide distribution (Lougheed and Stevenson 2004).


Map Key
This map only depicts Great Lakes introductions.

 
Great Lakes Nonindigenous Occurrences: Ulva flexuosa was recorded for the first time in 2003 in Muskegon Lake, Michigan, where the Muskegon River empties into Lake Michigan. Within the same drainage, it has also been recorded from White Lake and Mona Lake (Lougheed 2005, Lougheed and Stevenson 2004).


Table 1. Great Lakes region nonindigenous occurrences, the earliest and latest observations in each state/province, and the tally and names of HUCs with observations†. Names and dates are hyperlinked to their relevant specimen records. The list of references for all nonindigenous occurrences of Ulva (Enteromorpha) flexuosa subsp. flexuosa and flexuosa subsp. paradoxa are found here.

State/ProvinceFirst ObservedLast ObservedTotal HUCs with observations†HUCs with observations†
MI200320214Black-Macatawa; Muskegon; Pere Marquette-White; Thornapple

Table last updated 4/26/2024

† Populations may not be currently present.


Ecology: Ulva flexuosa is primarily a marine species but can tolerate freshwater conditions and is considered euryhaline. It is fast growing and prefers littoral zones with some wave action. In the Lake Michigan watershed, it occurs in mesotrophic to eutrophic aquatic habitats. Its establishment could have been spurred by increased salt concentrations from industrial sources and de-icing salts since the 1850s, as well as a lack of native grazing pressure.

In Muskegon Lake, the two subspecies co-occur in littoral zone macrophyte beds, although U. flexuosa subsp. paradoxa has been recorded to comprise 61% of the macroalgae present by biovolume, while U. flexuosa subsp. flexuosa only comprised 31%. In White Lake, Ulva sp. has been known to comprise 5% of the macroalgae present, and in Mona Lake, 20% (Lougheed and Stevenson 2004).

Ulva flexuosa is known from diverse habitats around the world and has been recorded in salinities ranging from 0.5–34‰. It has been recorded in tidal pools, on mudflats, in sea dikes, in polyhaline man-made waters, in oligohaline ponds, in drainage ditches, on rocks, and in warm calm waters below low tide (Kapraun 1970, Koeman and van den Hoek 1984, Hadi et al. 1989, Sahoo et al. 2003).

To reproduce, dioecious gametophytes produce male and female gametes that have two flagellae, while sporophytes produce zoospores with four flagellae (Koeman and van den Hoek 1984). Zoospores can be stored in the dark for over 50 days and still germinate with a 35% survival rate, suggesting that transport in ballast water is possible (Beach et al. 1995, Kolwalkar et al. 2007). Swarmers may be released with increasing exposure at night and greater desiccation, whereas vegetative growth may typically occur with shorter exposure times (Mairh et al. 1985).


Means of Introduction: Ulva flexuosa was very likely transported to the Great Lakes basin in ballast water or on ship hulls (Lougheed and Stevenson 2004).


Status: Established where recorded.


Great Lakes Impacts:
Summary of species impacts derived from literature review. Click on an icon to find out more...

EnvironmentalSocioeconomicBeneficial



The Great Lakes range of U. flexuosa is currently limited to Muskegon, Mona and White Lake. However, as it is able to form blooms in Muskegon Lake—which has a lower conductivity than Lake Erie—there is potential for it to thrive in other locations in the Great Lakes (Lougheed and Stevenson 2004).

Ulva flexuosa has a moderate environmental impact in the Great Lakes.
Realized:
The first recorded bloom of U. flexuosa in the Great Lakes occurred in Muskegon Lake, where it dominated the macrophyte community. Ulva flexuosa supports a relatively low density and diversity of epiphytes compared to native macrophytes (Lougheed and Stevenson 2004). Moreover, U. flexuosa has formed dense, free floating mats in Muskegon Lake during bloom conditions.

Potential:
Blooms of U. flexuosa in Europe have been associated with the extirpation of species of red algae and epiphytes (Schories et al. 1997). If U. flexuosa was to replace Oedogonium and Cladophora, which support many diatoms, food webs depending on such epiphytes could be negatively affected (Lougheed and Stevenson 2004).

When U. flexuosa blooms in nutrient rich marine coastal areas, a decline in biodiversity of native algae has been observed, as well as cascading food web effects that negatively impact macroinvertebrates, shorebirds, and fish (Lougheed and Stevenson 2004). Mats of U. flexuosa have also impacted abiotic conditions, as the dense algal cover alters redox conditions and chemical interactions between the sediment and water column (Lougheed and Stevenson 2004). It is hypothesized that shading of macrophytes could become problematic in Great Lakes if blooms were sustained for long periods of time (Lougheed and Stevenson 2004).

Ulva flexuosa has a moderate socio-economic impact in the Great Lakes.
Realized:
Ulva flexuosa is known to foul a variety of permanent man-made structures and boats. This can lead to expensive repairs of fouled boat motors. There are even reports of it growing on boats treated with antifouling paint (Kolwalker et al. 2007, Lougheed and Stevenson 2004).

Mats of U. flexuosa washed up on the shores of Muskegon Lake during the bloom of 2003 discouraged beach use by tourists (Lougheed and Stevenson 2004). In addition to the cost to boat users, these blooms also decreased the value of lakeshore property and discouraged tourism (Lougheed and Stevenson 2004).

Ulva flexuosa has a moderate beneficial impact in the Great Lakes.
Potential:
Ulva flexuosa is used as a bioindicator for metal contamination, including lead and iron (Ho 1987, Tabudravu et al. 2002). Ulva flexuosa was also found to be an economically efficient species for bioabsorption in industrial settings. It could be used as an eco-friendly alternative for wastewater treatment in dye manufacturing, tannery, textile, and cosmetic industries (Sivasamy et al. 2012).

 


Management:  

Regulations (pertaining to the Great Lakes region).
There are no known regulations for this species.

Note: Check federal, state/provincial, and local regulations for the most up-to-date information.

Control
Biological
There are no known biological control methods for this species .

Physical
There are no known physical control methods for this species.

Chemical

The presence of U. flexuosa is associated with high nutrient availability and high levels of salinity (Lougheed and Stevenson 2004, Shories et al. 1997). The reduction of pollution and nutrient run-off could decrease the viable habitat for U. flexuosa.

Note: Check state/provincial and local regulations for the most up-to-date information regarding permits for control methods. Follow all label instructions.


References (click for full reference list)


Other Resources:
Author: Sturtevant, R.A., M.M. McCarthy, and A. Fusaro.


Contributing Agencies:
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Revision Date: 9/12/2019


Citation for this information:
Sturtevant, R.A., M.M. McCarthy, and A. Fusaro., 2024, Ulva (Enteromorpha) flexuosa subsp. flexuosa and flexuosa subsp. paradoxa (Wolfen ex Roth) J. Agardh, 1883: U.S. Geological Survey, Nonindigenous Aquatic Species Database, Gainesville, FL, and NOAA Great Lakes Aquatic Nonindigenous Species Information System, Ann Arbor, MI, https://nas.er.usgs.gov/queries/greatlakes/FactSheet.aspx?Species_ID=2726, Revision Date: 9/12/2019, Access Date: 4/26/2024

This information is preliminary or provisional and is subject to revision. It is being provided to meet the need for timely best science. The information has not received final approval by the U.S. Geological Survey (USGS) and is provided on the condition that neither the USGS nor the U.S. Government shall be held liable for any damages resulting from the authorized or unauthorized use of the information.