Algae (sing. alga) are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms. The largest and most complex marine forms are called seaweeds. They are photosynthetic, like plants, and "simple" because they lack the many distinct organs found in land plants. Though the prokaryotic cyanobacteria (commonly referred to as blue-green algae) were traditionally included as "algae" in older textbooks, many modern sources regard this as outdated and restrict the term algae to eukaryotic organisms. All true algae therefore have a nucleus enclosed within a membrane and chloroplasts bound in one or more membranes. Algae constitute a paraphyletic and polyphyletic group, as they do not all descend from a common algal ancestor, although their chloroplasts seem to have a single origin.
Algae lack the various structures that characterize land plants, such as phyllids and rhizoids in nonvascular plants, or leaves, roots, and other organs that are found in tracheophytes. They are distinguished from protozoa in that they are photosynthetic. Many are photoautotrophic, although some groups contain members that are mixotrophic, deriving energy both from photosynthesis and uptake of organic carbon either by osmotrophy, myzotrophy, or phagotrophy. Some unicellular species rely entirely on external energy sources and have limited or no photosynthetic apparatus.
All algae have photosynthetic machinery ultimately derived from the cyanobacteria, and so produce oxygen as a by-product of photosynthesis, unlike other photosynthetic bacteria such as purple and green sulfur bacteria.
Algae are most prominent in bodies of water but are also common in terrestrial environments. However, terrestrial algae are usually rather inconspicuous and far more common in moist, tropical
regions than dry ones, because algae lack vascular tissues
and other adaptations to live on land. Algae are also found in other situations, such as on snow
and on exposed rocks in symbiosis
with a fungus as lichen
The various sorts of algae play significant roles in aquatic ecology. Microscopic forms that live suspended in the water column (phytoplankton) provide the food base for most marine food chains. In very high densities (so-called algal blooms) these algae may discolor the water and outcompete, poison, or asphyxiate other life forms. Seaweeds grow mostly in shallow marine waters, however some have been recorded to a depth of 300 m.Some are used as human food or harvested for useful substances such as agar, carrageenan, or fertilizer.
Study of algae
The study of marine and freshwater algae is called phycology or algology.
The US Algal Collection is represented by almost 300,000 accessioned and inventoried herbarium specimens.
have been traditionally included among the algae, referred to as the Cyanophytes or blue-green algae, recent works on algae usually exclude them due to large differences such as the lack of membrane-bound organelles, the presence of a single circular chromosome
, the presence of peptidoglycan
in the cell walls, and ribosomes different in size and content from eukaryotes . Rather than in chloroplasts, they conduct photosynthesis on specialized infolded cytoplasmic membranes called thylakoid membranes
. Therefore, they differ significantly from the algae despite occupying similar ecological niches.
By modern definitions algae are eukaryotes and conduct photosynthesis within membrane-bound organelles called chloroplasts. Chloroplasts contain circular DNA and are similar in structure to cyanobacteria, presumably representing reduced cyanobacterial endosymbionts. The exact nature of the chloroplasts is different among the different lines of algae, reflecting different endosymbiotic events. The table below lists the three major groups of algae and their lineage relationship is shown in the figure on the left. Note many of these groups contain some members that are no longer photosynthetic. Some retain plastids, but not chloroplasts, while others have lost them entirely.
| Supergroup affiliation
|| Summary |
||These algae have primary chloroplasts, i.e. the chloroplasts are surrounded by two membranes and probably developed through a single endosymbiotic event. The chloroplasts of red algae have chlorophylls a and d (often), and phycobilins, while those of the green alga have chloroplasts with chlorophyll a and b. Higher plants are pigmented similarly to green algae and probably developed from them, and thus Chlorophyta is a sister taxon to the plants; sometimes they are grouped as Viridiplantae. |
| Excavata and Rhizaria
These groups have green chloroplasts containing chlorophylls a and b . Their chloroplasts are surrounded by four and three membranes, respectively, and were probably retained from an ingested green alga.|
Chlorarachniophytes, which belong to the phylum Cercozoa, contain a small nucleomorph, which is a relict of the alga's nucleus.
Euglenids, which belong to the phylum Euglenozoa, live primarily in freshwater and have chloroplasts with only three membranes. It has been suggested that the endosymbiotic green algae were acquired through myzocytosis rather than phagocytosis.
|Chromista and Alveolata
These groups have chloroplasts containing chlorophylls a and c, and phycobilins. The latter chlorophyll type is not known from any prokaryotes or primary chloroplasts, but genetic similarities with the red algae suggest a relationship there.|
In the first three of these groups (Chromista), the chloroplast has four membranes, retaining a nucleomorph in cryptomonads, and they likely share a common pigmented ancestor, although other evidence casts doubt on whether the Heterokonts, Haptophyta, and Cryptomonads are in fact more closely related to each other than other groups.
The typical dinoflagellate chloroplast has three membranes, but there is considerable diversity in chloroplasts among the group, and it appears there were a number of endosymbiotic events here. The Apicomplexa, a group of closely related parasites, also have plastids called apicoplasts. Apicoplasts are not photosynthetic but appear to have a common origin with dinoflagellates chloroplasts.
It was W.H.Harvey (1811 — 1866) who first divided the algae into four divisions based on their pigmentation. This is the first use of a biochemical criterion in plant systematics. Harvey's four divisions were: red algae (Rhodophyta), brown algae (Heteromontophyta), green algae (Chlorophyta) and Diatomaceae (Dixon, 1973 p.232).
Forms of algae
Most of the simpler algae are unicellular flagellates
, but colonial and non-motile forms have developed independently among several of the groups. Some of the more common organizational levels, more than one of which may occur in the life cycle
of a species, are
- Colonial: small, regular groups of motile cells
- Capsoid: individual non-motile cells embedded in mucilage
- Coccoid: individual non-motile cells with cell walls
- Palmelloid: non-motile cells embedded in mucilage
- Filamentous: a string of non-motile cells connected together, sometimes branching
- Parenchymatous: cells forming a thallus with partial differentiation of tissues
In three lines even higher levels of organization have been reached, with full tissue differentiation. These are the brown algae, —some of which may reach 50 m in length (kelps)—the red algae, and the green algae. The most complex forms are found among the green algae (see Charales and Charophyta), in a lineage that eventually led to the higher land plants. The point where these non-algal plants begin and algae stop is usually taken to be the presence of reproductive organs with protective cell layers, a characteristic not found in the other alga groups.
The first plants on earth evolved from shallow freshwater algae much like Chara some 400 million years ago. These probably had an isomorphic alternation of generations and were probably heterotrichous. Fossils of isolated land plant spores suggest land plants may have been around as long as 475 million years ago.
Algae and symbioses
Some species of algae form symbiotic relationships
with other organisms. In these symbioses
, the algae supply photosynthates (organic substances) to the host organism providing protection to the algal cells. The host organism derives some or all of its energy requirements from the algae. Examples include
- lichens: a fungus is the host, usually with a green alga or a cyanobacterium as its symbiont. Both fungal and algal species found in lichens are capable of living independently, although habitat requirements may be greatly different from those of the lichen pair.
- corals: algae known as zooxanthellae are symbionts with corals. Notable amongst these is the dinoflagellate Symbiodinium, found in many hard corals. The loss of Symbiodinium, or other zooxanthellae, from the host is known as coral bleaching.
- sponges: green algae live close to the surface of some sponges, for example, breadcrumb sponge (Halichondria panicea). The alga is thus protected from predators; the sponge is provided with oxygen and sugars which can account for 50 to 80% of sponge growth in some species.
, the three main algal Phyla
, have life-cycles which show tremendous variation with considerable complexity. In general there is an asexual phase where the seaweed's cells are diploid
, a sexual phase where the cells are haploid
followed by fusion of the male and female gametes
. Asexual reproduction is advantageous in that it permits efficient population increases, but less variation is possible. Sexual reproduction allows more variation but is more costly because of the waste of gametes that fail to mate, among other things. Often there is no strict alternation between the sporophyte and gametophyte phases and also because there is often an asexual phase, which could include the fragmentation of the thallus.
Numbers and distribution
In the British Isles the UK Biodiversity Steering Group Report estimated there to be 20,000 algal species in the UK, freshwater and marine, about 650 of these are seaweeds. Another checklist of freshwater algae reported only about 5000 species. It seems therefore that the 20,000 is an overestimate or an error (John, 2002 p.1).
The Smithsonian collection of algae has over 300,000 specimens.
Worldwide it is thought that there are over 5,000 species of red algae, 1,500 — 2,000 of brown algae and 8,000 of green algae. In Australia it is estimated that there are over 1,300 species of red algae, 350 species of brown algae and approximately 2,000 species of green algae totalling 3,650 species of algae in Australia.
Around 400 species appear to be an average figure for the coastline of South African west coast.
669 marine species have been described from California (U.S.A.).
642 entities are listed in the check-list of Britain and Ireland (Hardy and Guiry, 2006).
No publication has been found which attempts to discuss the general distribution of algae in the seas worldwide. However, notes and comments have been made by some authors.
The floristic discontinuities may appear to determined by geographical features such as Antarctica
, long distances of ocean or general land masses. However, the distances between Norway, the Faroes and Iceland do not show great changes in distribution.
There has been dispersal in some species by ships, water currents and the like; further, some algae can quickly become entangled and make drifting mats. Two red species have been introduced from the Pacific to Europe and the Mediterranean: Bonnemaisonia hamifera Hariot and Asparagopsis armata Harvey, A. armata is a native of Australia.Colpomenia peregrina is a native of the Pacific but has also invaded Europe.
Britain and Ireland
- Hardy, F G and Guiry, M D (2006) A Check-list and Atlas of the Seaweeds of Britain and Ireland. British Phycological Society, London. ISBN 3 906166 35 X
- Cullinane, J P (1973) Phycology of the South Coast of Ireland. The Cork University Press, University College Cork.
Northumberland and Durham (England)
- Hardy, F G and Aspinall, R J (1988). An Atlas of the Seaweeds of Northumberland and Durham. Northumberland Biological Records Centre. The Hancock Museum. The University Newcastle upon Tyne. Special publication: 3. ISBN 0 9509680 5 6
- Morton, O (1994) Marine Algae of Northern Ireland. Ulster Museum, Belfast. ISBN 0 900761 28 8
Republic of Ireland: County Donegal
- Morton, O. The marine macroalgae of County Donegal, Ireland. Bull. Ir. biogeog. Soc. 27:3 - 164.
Isle of Man
- Knight, M and Park, M W (1931) Manx algae. An algal survey of the south end of the Isle of Man. L.M.B.C. Mem. Typ. Br. Mar. Pl. 390: 1 - 155.
- Kjellman, F R (1883) The algae of the Arctic Sea. K. sevenka. VetenskAkad. Handl. 20: 1 - 350.
- Lund, S (1959) The Marine Algae of East Greenland. I. Taxonomical part. Meddr. Grønland 156: 1 - 247.
- Borgesen, F (1903) Marine Algae, pp.339 - 532. In, Warming, E. (Ed.), Botany of the Faröes Based Upon Danish Investigations. Part II. Copenhagen. [reprint 1970]
- Cabioc'h, J; Floc'h, J Y; Le Toquin, A; Boudouresque, C F; Meinesz, A and Verlaque, M (1992) Guide des algues des mers d'Europe. Delachaux et Niestlé, Switzerland.
- Gayral, P (1958) Algues de la Côte Atlantique Marocaine. Rabat.
- Gayral, P (1966) Algues des Côtes Françaises. Paris.
- Borgesen, F (1925) Marine algae from the Canary Islands, especially from Tenerife and Gran Canaria. I. Chlorophyceae. Biol. Meddr 5: 1 - 113.
- Borgesen, F (1926) Marine algae from the Canary Islands especially from Tenerife and Gran Canaria. II. Phaeophyceae. Biol. Meddr 6: 1 - 112.
- Borgesen, F (1927) Marine algae from the Canary Islands. III. Rhodophyceae. Part I, Bangiales and Nemalionales. Biol. Meddr 6: 1 - 97.
- Borgesen, F (1929) Marine algae from the Canary Islands. III Rhodophyceae. Part II. Cryptonemiales, Gigartinales and Rhodymeniales. Biol. Meddr 8: 1 - 97.
- Borgesen, F (1930) Marine algae from the Canary Islands. III Rhodophyceae. Part II. Cryptonemiales, Gigartinales and Rhodymeniales. Biol. Meddr 9: 1 - 159.
- Taylor, W R (1957) Marine Algae of the Northeastern Coast of North America. University of Michigan Press, Ann Arbor.
- Abbott, I A and Hollenberg, G J (1976) Marine Algae of California. Stanford University Press, California.
- Wehr, J D and Sheath, R G (2003) Freshwater Algae of North America: Ecology and Classification. Academic Press, USA.
- Stegenga, H Bolton, J J and Anderson, R J (1997) Seaweeds of the South African West Coast. Bolus Herbarium Number 18, Publication jointly financed by the Fourcade Bequest and the Research Committee of the University of Cape Town and the Foundation for Research Development.
- Lindauer, V W; Chapman, V J and Aiken, M (1961) The Marine Algae of New Zealand. Part II. Phaeophyta. Nova Hedwigia 3: 129 - 350.
- Chapman, V J (1969) The Marine Algae of New Zealand. Part III issues 1. Lehre: J. Cramer, 1 - 113.
- Chapman, V J and Dromgoole, F I (1970) The Marine Algae of New Zealand. Part III issues 2. Lehre: J.Cramer, 115 - 154.
- Chapman, V J and Parkinson, P G (1974) The Marine Algae of New Zealand. Part III issues 3. Lehre: J.Cramer,155 - 278.
- Chapman, V J (1979) The Marine Algae of New Zealand. Part III issues 4. Lehre: J.Cramer, 279 - 420.
Uses of algae
For centuries seaweed has been used as a fertilizer; Orwell writing in the 16th century referring to drift weed in South Wales: "This kind of ore they often gather and lay in heaps where it heats and rots, and will have a strong and loathsome smell; when being so rotten they cast it on the land, as they do their muck, and thereof springeth good corn, especially barley" and "After spring tides or great rigs of the sea, they fetch it in sacks on horse brackets, and carry the same three, four, or five miles, and cast it on the lande, which doth very much better the ground for corn and grass" (Chapman p.35).
Algae are used by humans in many ways. They are used as fertilizers, soil conditioners and are a source of livestock feed. Because many species are aquatic and microscopic, they are cultured in clear tanks or ponds and either harvested or used to treat effluents pumped through the ponds. Algaculture on a large scale is an important type of aquaculture in some places.
Maerl is commonly used as a soil conditioner, it is dredged from the sea floor and crushed to form a powder. It is still harvested around the coasts of Brittany in France and off Falmouth, Cornwall (also extensively in western Ireland) and is a popular fertilizer in these days of organic gardening investigated Falmouth maerl and found that L. corallioides predominated down to 6 m and P. calcareum from 6-10 m (Blunden et al., 1981).
Chemical analysis of maerl showed that it contained 32.1% CaCO3 and 3.1% MgCO3 (dry weight).
- Algae can be grown to produce biohydrogen. In 1939 a German researcher named Hans Gaffron, while working at the University of Chicago, observed that the algae he was studying, Chlamydomonas reinhardtii (a green-alga), would sometimes switch from the production of oxygen to the production of hydrogen. Gaffron never discovered the cause for this change and for many years other scientists failed to repeat his findings. In the late 1990s professor Anastasios Melis, a researcher at the University of California at Berkeley, discovered that if the algae culture medium is deprived of sulfur it will switch from the production of oxygen (normal photosynthesis), to the production of hydrogen. He found that the enzyme responsible for this reaction is hydrogenase, but that the hydrogenase lost this function in the presence of oxygen. Melis found that depleting the amount of sulfur available to the algae interrupted its internal oxygen flow, allowing the hydrogenase an environment in which it can react, causing the algae to produce hydrogen. Chlamydomonas moeweesi is also a good strain for the production of hydrogen. Scientists at the U.S. Department of Energy’s Argonne National Laboratory are currently trying to find a way to take the part of the hydrogenase enzyme that creates the hydrogen gas and introduce it into the photosynthesis process. The result would be a large amount of hydrogen gas, possibly on par with the amount of oxygen created.
- Algae can be used to make biodiesel (see algaculture), bioethanol and biobutanol and by some estimates can produce vastly superior amounts of vegetable oil, compared to terrestrial crops grown for the same purpose.
- Algae can be used in oil production which could replace the petrol and other gas products in the near future.
- Algae can be grown to produce biomass, which can be burned to produce heat and electricity.
- Algae are used in wastewater treatment facilities, reducing the need for greater amounts of toxic chemicals than are already used.
- Algae can be used to capture fertilizers in runoff from farms. When subsequently harvested, the enriched algae itself can be used as fertilizer.
- Algae Bioreactors are used by some powerplants to reduce CO2 emissions. The CO2 can be pumped into a pond, or some kind of tank, on which the algae feed. Alternatively, the bioreactor can be installed directly on top of a smokestack. This technology has been pioneered by Massachusetts-based GreenFuelTechnologies.
, (probably confused with Mastocarpus stellatus
, common name: Irish moss), is also used as "carrageen
". The name carrageenan comes from the Irish Gaelic for Chondrus crispus
. It is an excellent stabiliser in milk products - it reacts with the milk protein caesin, other products include: petfoods, toothpaste, ice-creams and lotions etc. Alginates in creams and lotions are absorbable through the skin.
Seaweeds are an important source of food, especially in Asia; They are excellent sources of many vitamins including: A, B1
. They are rich in iodine
Algae is commercially cultivated as a nutritional supplement. One of the most popular microalgal species is Spirulina (Arthrospira platensis), which is a Cyanobacteria (known as blue-green algae), and has been hailed by some as a superfood. Other algal species cultivated for their nutritional value include; Chlorella (a green algae), and Dunaliella (Dunaliella salina), which is high in beta-carotene and is used in vitamin C supplements.
In China at least 70 species of algae are eaten as is the Chinese "vegetable" known as fat choy (which is actually a cyanobacterium). Roughly 20 species of algae are used in everyday cooking in Japan.
Certain species are edible; the best known, especially in Ireland is Palmaria palmata (Linnaeus) O. Kuntze, also known as Rhodymenia palmata (Linnaeus) Kuntze, common name: dulse). This is a red alga which is dried and may be bought in the shops in Ireland. It is eaten raw, fresh or dried, or cooked like spinach. Similarly, Durvillaea antarctica is eaten in Chile, common name: cochayuyo.
Porphyra (common name: purple laver), is also collected and used in a variety of ways (e.g. "laver bread" in the British Isles). In Ireland it is collected and made into a jelly by stewing or boiling. Preparation also involves frying with fat or converting to a pinkish jelly by heating the fronds in a saucepan with a little water and beating with a fork. It is also collected and used by people parts of Asia, specifically China, Korea (gim) and Japan (nori) and along most of the coast from California to British Columbia. The Hawaiians and the Maoris of New Zealand also use it.
One particular use is in "instant" puddings, sauces and creams. Ulva lactuca (common name: sea lettuce), is used locally in Scotland where it is added to soups or used in salads. Alaria esculenta (common name: badderlocks or dabberlocks), is used either fresh or cooked, in Greenland, Iceland, Scotland and Ireland.
The oil from some algae have high levels of unsaturated fatty acids. Arachidonic acid (a polyunsaturated fatty acid), is very high in Parietochloris incisa, (a green alga) where it reaches up to 47% of the triglyceride pool (Bigogno C et al. Phytochemistry 2002, 60, 497).
Some varieties of algae are a vegetarian / vegan / plant based source of long chain essential omega-3 fatty acids Docosahexaenoic acid (DHA) and Eicosapentaenoic acid (EPA) in addition to vitamin B12. Fish oil contains the omega-3 fatty acids, but the original source is algae, which are eaten by marine life such as copepods and passed up the food chain.
There are also commercial uses of algae as agar.
The natural pigments produced by algae can be used as an alternative to chemical dyes and coloring agents. Many of the paper products used today are not recyclable because of the chemical inks that they use, paper recyclers have found that inks made from algae are much easier to break down. There is also much interest in the food industry into replacing the coloring agents that are currently used with coloring derived from algal pigments. Algae can be used to make pharmaceuticalsSewage can be treated with algae as well
Some cosmetics can come from microalgae as well.
In Israel, a species of green algae is grown in water tanks, then exposed to direct sunlight and heat which causes it to become bright red in color. It is then harvested and used as a natural pigment for foods such as Salmon.
Between 100,000 and 170,000 wet tons of Macrocystis
are harvested annually in California
for alginate extraction and abalone feed.
Further references to the uses
- Guiry, M D and Blunden, G (Eds) (1991) Seaweed Resources in Europe: Uses and Potential. John Wiley & Sons. ISBN 0-471-92947-6
- Mumford, T F and Miura, A (1988) 4. Porphyra as food: cultivation and economics. p.87 — 117. In Lembi, C.A. and Waaland, J.R. (Ed.) Algae and Human Affairs. 1988. Cambridge University Press. ISBN 0 521 32115 8
Collecting and preserving specimens
Seaweed specimens can be collected and preserved for research. Such preserved specimens will keep for two or three hundred years. Those of Carl von Linné
(1707 — 1778) are still available for reference, and are used. Specimens may be collected from the shore; those below low tide must be collected by diving or dredging. The whole algal specimen should be collected, that is the holdfast
, stipe and lamina. Specimens of algae reproducing will be the more useful for identification and research. When collected the details of the location and site should be noted. They can then be preserved pressed on paper or in a preserving liquid such as alcohol or solution of 5 per cent formalin/seawater. However, formalin is reported to be carcinogenic.
Biological Exposure Scale
The ecology of the shores of the British Isles, including a discussion of the different shores from sheltered to exposed along with an exposure scale, is given by Lewis (1964). An exposure scale of five stages is given:- Very Exposed Shores; Exposed Shores; Semi-exposed Shores; Sheltered Shores and Very Sheltered Shores. Factors indicating the differences between these exposure scales are detailed. Very Exposed Shores have a wide Verrucaria
zone entirely above the upper tide level, a Porphyra
zone above the barnacle level and Lichina pygmaea
is locally abundant. The eulittoral zone is dominated by barnacles
with a coralline belt in the very low littoral along with other Rhodophyta and Alaria
in the upper sublittoral. Exposed shores show a Verrucaria
belt mainly above the high tide, with Porphyra
and Lichina pygmaea
. The mid shore is dominated by barnacles, limpets and some Fucus
. Some Rhodophyta. Himanthalia
and some Rhodophyta such as Mastocarpus
are found in the low littorral with Himanthalia
and Laminaria digitata
dominant in the upper sublittoral. The semi-exposed shores show a Verrucaria
belt just above high tide with clear Pelvetia
in the upper-littoral and Fucus serratus
in the lower-littoral. Limpets, barnacles and short Fucus vesiculosus
midshore. Fucus serratus
with Rhodophyta, (Laurencia
, Mastocarpus stellatus
and Saccorhiza polyschides
and small algae common in the sublittoral. The sheltered shores show a narrow Verrucaria
zone at high water and a full sequence of fucoids: Pelvetia
, Fucus spiralis
, Fucus vesiculosus
, Fucus serratus
, Ascophyllum nodosum
. Laminaria digitata
is dominant the upper sublittoral. The very sheltered shores show a very narrow zone of Verrucaria
, the dominance of the littoral by a full sequence of the fucoids and Ascophyllum
covering the rocks. Laminaria saccharina
and or Furcellaria
- Abbott, I.A. and Hollenberg, G.J. 1976. Marine Algae of California. Stanford University Press, California. ISBN 0-8047-0867-3
- Brodie, J.A. and Irvine, L.M. 2003. Seaweeds of the British Isles. Volume 1 Part 3B. The Natural History Museum, London. ISBN 1 898298 87 4
- Burrows, E.M. 1991. Seaweeds of the British Isles. Volume 2. British Museum (Natural History), London. ISBN 0-565-00981-8
- Christensen, T. 1987. Seaweeds of the British Isles. Tribophyceae. Volume 4. British Museum (Natural History), London. ISBN 0-565-00980-X
- Dixon, P.S. and Irvine, L.M. 1977. Seaweeds of the British Isles. Volume 1. Part 1. Introduction, Nemaliales, Gigartinales. British Museum (Natural History), London. ISBN 0 565 00781 5
- Greeson, Phillip E. 1982. An annotated key to the identification of commonly occurring and dominant genera of Algae observed in the Phytoplankton of the United States [Geological Survey Water-Supply Paper 2079]. Washington, D.C.: United States Government Printing Office.
- Irvine, L.M. 1983. Seaweeds of the British Isles. Volume 1, Part 2A. British Museum (Natural History), London. ISBN 0-565-00871-4
- Irvine, L.M. and Chamberlain, Y.M. 1994. Seaweeds of the British Isles. Volume 1 Part 2B. The Natural History Museum, London. ISBN 0 11 310016 7
- Fletcher, R.L. 1987. Seaweeds of the British Isles. Volume 3 Part 1. British Museum (Natural History), London. ISBN 0-565-00992-3
- John, D.M., Whitton, B.A. and Brook, J.A. (Eds.) 2002. The Freshwater Algal Flora of the British Isles. Cambridge University Press, UK. ISBN 0 521 77051 3
- Stegenga, H., Bolton, J.J. and Anderson, R.J.1997. Seaweeds of the South African west coast. Boltus Herbarium, University of Cape Town. ISBN 0-7992-1793-x
- Taylor, W.R. 1957. Marine algae of the north-eastern coasts of North America. Revised edition. University of Michigan Press. Ann Arbor.
- Chapman, V.J. 1950.p.36. Seaweeds and their Uses. Methuen & Co. Ltd., London.
- Guiry, M.D. and Blunden, G. (Eds) 1991. Seaweed Resources in Europe: Uses and Potential. John Wiley & Sons. ISBN 0-471-92947-6
- Lembi, C.A. and Waaland, J.R. (Eds.) 1988. Algae and Human Affairs. Cambridge University Press, Cambridge. ISBN 0-521-32115-8