Algae 1

Algae

Algae

Laurencia, a genus of red algae from HawaiiScientific classification

Domain: Eukaryota

Included groups

•• Archaeplastida• Chlorophyta (Green algae)• Rhodophyta (Red algae)•• Glaucophyta

• Rhizaria, Excavata•• Chlorarachniophytes•• Euglenids

• Chromista, Alveolata•• Heterokonts

• Bacillariophyceae (Diatoms)•• Axodine•• Bolidomonas•• Eustigmatophyceae• Phaeophyceae (Brown algae)• Chrysophyceae (Golden algae)•• Raphidophyceae•• Synurophyceae• Xanthophyceae (Yellow-green algae)

•• Cryptophyta•• Dinoflagellates•• Haptophyta

Excluded groups

•• Cyanobacteria• Plantae

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The lineage of algae according to Thomas Cavalier-Smith. The exactnumber and placement of endosymbiotic events is currently unknown,so this diagram can be taken only as a general guide[1][2] It represents

the most parsimonious way of explaining the three types ofendosymbiotic origins of plastids. These types include the

endosymbiotic events of cyanobacteria, red algae and green algae,leading to the hypothesis of the supergroups Archaeplastida,

Chromalveolata and Cabozoa respectively. However, the monophyly ofCabozoa has been refuted and the monophylies of Archaeplastida and

Chromalveolata are currently strongly challenged. Endosymbioticevents are noted by dotted lines.

Algae (/ˈældʒiː/ or /ˈælɡiː/; singular alga /ˈælɡə/,Latin for "seaweed") are a very large and diversegroup of simple, typically autotrophic organisms,ranging from unicellular to multicellular forms, suchas the giant kelps that grow to 65 meters in length.They are photosynthetic like plants, and "simple"because their tissues are not organized into the manydistinct organs found in land plants. The largest andmost complex marine forms are called seaweeds.

Though the prokaryotic cyanobacteria (commonlyreferred to as blue-green algae) were traditionallyincluded as "algae" in older textbooks, many modernsources regard this as outdated[3] as they are nowconsidered to be bacteria.[4] The term algae is nowrestricted to eukaryotic organisms.[5] All true algaetherefore have a nucleus enclosed within a membraneand plastids bound in one or more membranes.[3][6]

Algae constitute a paraphyletic and polyphyleticgroup,[3] as they do not include all the descendants ofthe last universal ancestor nor do they all descendfrom a common algal ancestor, although theirplastids seem to have a single origin.[1] Diatoms arealso examples of algae.

Blue-green algae are found in the fossil record datingback to approximately 3 billion years in thePrecambrian.

Algae exhibit a wide range of reproductive strategies,from simple, asexual cell division to complex forms of sexual reproduction.[7]

Algae lack the various structures that characterize land plants, such as phyllids (leaves) and rhizoids in nonvascularplants, or leaves, roots, and other organs that are found in tracheophytes (vascular plants). Many arephotoautotrophic, although some groups contain members that are mixotrophic, deriving energy both fromphotosynthesis and uptake of organic carbon either by osmotrophy, myzotrophy, or phagotrophy. Some unicellularspecies rely entirely on external energy sources and have limited or no photosynthetic apparatus.

Nearly all algae have photosynthetic machinery ultimately derived from cyanobacteria, and so produce oxygen as aby-product of photosynthesis, unlike other photosynthetic bacteria such as purple and green sulfur bacteria.Fossilized filamentous algae from the Vindhya basin have been dated back to 1.6 to 1.7 billion years ago.[8]

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Etymology and study

Title page of Samuel Gottlieb Gmelin, HistoriaFucorum, dated 1768.

The singular alga is the Latin word for a particular seaweed andretains that meaning in English.[9] The etymology is obscure.Although some speculate that it is related to Latin algēre, "becold",[10] there is no known reason to associate seaweed withtemperature. A more likely source is alliga, "binding, entwining."[11]

Since Algae has become a biological classification, alga can alsomean one classification under Algae, parallel to a fungus being aspecies of fungi, a plant being a species of plant, and so on.

The ancient Greek word for seaweed was φῦκος (fūkos or phykos),which could mean either the seaweed, probably Red Algae, or a reddye derived from it. The Latinization, fūcus, meant primarily thecosmetic rouge. The etymology is uncertain, but a strong candidatehas long been some word related to the Biblical פוך (pūk), "paint" (ifnot that word itself), a cosmetic eye-shadow used by the ancientEgyptians and other inhabitants of the eastern Mediterranean. Itcould be any color: black, red, green, blue.[12]

Accordingly the modern study of marine and freshwater algae iscalled either phycology or algology. The name Fucus appears in anumber of taxa.

Classification

False-color Scanning electron micrograph of the unicellular coccolithophore,Gephyrocapsa oceanica.

While Cyanobacteria have beentraditionally included among the Algae,recent works usually exclude them due tolarge differences such as the lack ofmembrane-bound organelles, the presenceof a single circular chromosome, thepresence of peptidoglycan in the cell walls,and ribosomes different in size and contentfrom those of the Eukaryotes.[13][14] Ratherthan in chloroplasts, they conductphotosynthesis on specialized infoldedcytoplasmic membranes called thylakoidmembranes. Therefore, they differsignificantly from the Algae despiteoccupying similar ecological niches.

By modern definitions Algae are Eukaryotesand conduct photosynthesis withinmembrane-bound organelles calledchloroplasts. Chloroplasts contain circularDNA and are similar in structure to

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Cyanobacteria, presumably representing reduced cyanobacterial endosymbionts. The exact nature of the chloroplastsis different among the different lines of Algae, reflecting different endosymbiotic events. The table below describesthe composition of the three major groups of Algae. Their lineage relationships are shown in the figure in the upperright. Many of these groups contain some members that are no longer photosynthetic. Some retain plastids, but notchloroplasts, while others have lost plastids entirely.Phylogeny based on plastid[15] not nucleocytoplasmic genealogy:

Cyanobacteria

Cyanelles

Rhodoplasts

Rhodophytes

Heterokonts

Cryptophytes

Haptophytes

Chloroplasts

Euglenophytes

Chlorophytes

Charophytes

Higher plants (Embryophyta)

Chlorarachniophytes

Supergroupaffiliation

Members Endosymbiont Summary

Primoplantae/Archaeplastida

•• Chlorophyta•• Rhodophyta•• Glaucophyta

Cyanobacteria These Algae have primary chloroplasts, i.e. the chloroplasts are surrounded by twomembranes and probably developed through a single endosymbiotic event. Thechloroplasts of Red Algae have chlorophylls a and c (often), and phycobilins, whilethose of Green Algae have chloroplasts with chlorophyll a and b. Higher plants arepigmented similarly to Green Algae and probably developed from them, and thusChlorophyta is a sister taxon to the plants; sometimes they are grouped asViridiplantae.

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Excavata andRhizaria

•• Chlorarachniophytes•• Euglenids

Green Algae These groups have green chloroplasts containing chlorophylls a and b.[13] Theirchloroplasts are surrounded by four and three membranes, respectively, and wereprobably retained from ingested Green Algae.

Chlorarachniophytes, which belong to the phylum Cercozoa, contain a smallnucleomorph, which is a relict of the algae's nucleus.

Euglenids, which belong to the phylum Euglenozoa, live primarily in freshwater andhave chloroplasts with only three membranes. It has been suggested that theendosymbiotic Green Algae were acquired through myzocytosis rather thanphagocytosis.

Chromista andAlveolata

•• Heterokonts•• Haptophyta• Cryptomonads•• Dinoflagellates

Red Algae These groups have chloroplasts containing chlorophylls a and c, and phycobilins.Theshape varies from plant to plant. they may be of discoid, plate-like, reticulate,cup-shaped, spiral or ribbon shaped. They have one or more pyrenoids to preserveprotein and starch. The latter chlorophyll type is not known from any prokaryotes orprimary chloroplasts, but genetic similarities with the Red Algae suggest arelationship there.

In the first three of these groups (Chromista), the chloroplast has four membranes,retaining a nucleomorph in Cryptomonads, and they likely share a commonpigmented ancestor, although other evidence casts doubt on whether the Heterokonts,Haptophyta, and Cryptomonads are in fact more closely related to each other than toother groups.[2][16]

The typical dinoflagellate chloroplast has three membranes, but there is considerablediversity in chloroplasts within the group, and it appears there were a number ofendosymbiotic events.[1] The Apicomplexa, a group of closely related parasites, alsohave plastids called apicoplasts. Apicoplasts are not photosynthetic but appear to havea common origin with Dinoflagellate chloroplasts.[1]

W.H.Harvey (1811—1866) was the first to divide the Algae into four divisions based on their pigmentation. This isthe first use of a biochemical criterion in plant systematics. Harvey's four divisions are: Red Algae (Rhodophyta),Brown Algae (Heteromontophyta), Green Algae (Chlorophyta) and Diatomaceae.[17]

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Relationship to higher plantsThe first plants on earth probably evolved from shallow freshwater algae much like Chara some 400 million yearsago. These probably had an isomorphic alternation of generations and were probably filamentous. Fossils of isolatedland plant spores suggest land plants may have been around as long as 475 million years ago.[18][19]

Morphology

The kelp forest exhibit at the Monterey Bay Aquarium. Athree-dimensional, multicellular thallus.

A range of algal morphologies are exhibited, andconvergence of features in unrelated groups is common.The only groups to exhibit three dimensional multicellularthalli are the reds and browns, and some chlorophytes.[20]

Apical growth is constrained to subsets of these groups: theflorideophyte reds, various browns, and the charophytes.[20]

The form of charophytes is quite different to those of redsand browns, because have distinct nodes, separated byinternode 'stems'; whorls of branches reminiscent of thehorsetails occur at the nodes.[20] Conceptacles are anotherpolyphyletic trait; they appear in the coralline algae and theHildenbrandiales, as well as the browns.[20]

Most of the simpler algae are unicellular flagellates oramoeboids, but colonial and non-motile forms havedeveloped independently among several of the groups.Some of the more common organizational levels, more thanone 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 tissuesIn three lines even higher levels of organization have been reached, with full tissue differentiation. These are thebrown algae,[21]—some of which may reach 50 m in length (kelps)[22]—the red algae,[23] and the green algae.[24]

The most complex forms are found among the green algae (see Charales and Charophyta), in a lineage thateventually led to the higher land plants. The point where these non-algal plants begin and algae stop is usually takento be the presence of reproductive organs with protective cell layers, a characteristic not found in the other algagroups.

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Symbiotic algaeSome species of algae form symbiotic relationships with other organisms. In these symbioses, the algae supplyphotosynthates (organic substances) to the host organism providing protection to the algal cells. The host organismderives some or all of its energy requirements from the algae. Examples are as follows.

Lichens

Rock lichens in Ireland.

Lichens are defined by the International Association for Lichenology tobe "an association of a fungus and a photosynthetic symbiont resultingin a stable vegetative body having a specific structure."[25] The fungi,or mycobionts, are from the Ascomycota with a few from theBasidiomycota. They are not found alone in nature but when theybegan to associate is not known.[26] One mycobiont associates with thesame phycobiont species, rarely two, from the Green Algae, except thatalternatively the mycobiont may associate with the same species ofCyanobacteria (hence "photobiont" is the more accurate term). Aphotobiont may be associated with many specific mycobionts or liveindependently; accordingly, lichens are named and classified as fungal

species.[27] The association is termed a morphogenesis because the lichen has a form and capabilities not possessedby the symbiont species alone (they can be experimentally isolated). It is possible that the photobiont triggersotherwise latent genes in the mycobiont.[28]

Coral reefs

Floridian coral reef

Coral reefs are accumulated from the calcareous exoskeletons ofmarine invertebrates of the Scleractinia order; i.e., the Stony Corals. Asanimals they metabolize sugar and oxygen to obtain energy for theircell-building processes, including secretion of the exoskeleton, withwater and carbon dioxide as byproducts. As the reef is the result of afavorable equilibrium between construction by the corals anddestruction by marine erosion, the rate at which metabolism canproceed determines the growth or deterioration of the reef.

Algae of the Dinoflagellate phylum are often endosymbionts in thecells of marine invertebrates, where they accelerate host-cell

metabolism by generating immediately available sugar and oxygen through photosynthesis using incident light andthe carbon dioxide produced in the host. Endosymbiont algae in the Stony Corals are described by the termzooxanthellae, with the host Stony Corals called on that account hermatypic corals, which although not a taxon arenot in healthy condition without their endosymbionts. Zooxanthellae belong almost entirely to the genusSymbiodinium.[29] The loss of Symbiodinium from the host is known as coral bleaching, a condition which unlesscorrected leads to the deterioration and loss of the reef.

Algae 8

Sea spongesGreen 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 50to 80% of sponge growth in some species.[30]

Life-cycleRhodophyta, Chlorophyta and Heterokontophyta, the three main algal Phyla, have life-cycles which showtremendous variation with considerable complexity. In general there is an asexual phase where the seaweed's cellsare diploid, a sexual phase where the cells are haploid followed by fusion of the male and female gametes. Asexualreproduction is advantageous in that it permits efficient population increases, but less variation is possible. Sexualreproduction allows more variation, but is more costly. Often there is no strict alternation between the sporophyteand also because there is often an asexual phase, which could include the fragmentation of the thallus.[22][31][32]

Numbers

Algae on coastal rocks at Shihtiping in Taiwan

The Algal Collection of the US National Herbarium (located in theNational Museum of Natural History) consists of approximately320,500 dried specimens, which, although not exhaustive (noexhaustive collection exists), gives an idea of the order of magnitude ofthe number of algal species (that number remains unknown).[33]

Estimates vary widely. For example, according to one standardtextbook,[34] in the British Isles the UK Biodiversity Steering GroupReport estimated there to be 20000 algal species in the UK. Anotherchecklist reports only about 5000 species. Regarding the difference ofabout 15000 species, the text concludes: "It will require many detailedfield surveys before it is possible to provide a reliable estimate of the total number of species ...."

Regional and group estimates have been made as well:• 5000–5500 species of Red Algae worldwide• "some 1300 in Australian Seas"[35]

• 400 seaweed species for the western coastline of South Africa,[36] and 212 species from the coast ofKwaZulu-Natal.[37] Some of these are duplicates as the range extends across both coasts, and the total recorded isprobably about 500 species. Most of these are listed in List of seaweeds of South Africa. These excludephytoplankton and crustose corallines.

• 669 marine species from California (US)[38]

• 642 in the check-list of Britain and Ireland[39]

and so on, but lacking any scientific basis or reliable sources, these numbers have no more credibility than the Britishones mentioned above. Most estimates also omit the microscopic Algae, such as the phytoplankta, entirely.

DistributionThe topic of distribution of algal species has been fairly well studied since the founding of phytogeography in themid-19th century AD.[40] Algae spread mainly by the dispersal of spores analogously to the dispersal of Plantae byseeds and spores. Spores are everywhere in all parts of the Earth: the waters fresh and marine, the atmosphere,free-floating and in precipitation or mixed with dust, the humus and in other organisms, such as humans. Whether aspore is to grow into an organism depends on the combination of the species and the environmental conditions ofwhere the spore lands.

Algae 9

The spores of fresh-water Algae are dispersed mainly by running water and wind, as well as by living carriers.[41]

The bodies of water into which they are transported are chemically selective. Marine spores are spread by currents.Ocean water is temperature selective, resulting in phytogeographic zones, regions and provinces.[42]

To some degree the distribution of Algae is subject to floristic discontinuities caused by geographical features, suchas Antarctica, long distances of ocean or general land masses. It is therefore possible to identify species occurring bylocality, such as "Pacific Algae" or "North Sea Algae". When they occur out of their localities, it is usually possibleto hypothesize a transport mechanism, such as the hulls of ships. For example, Ulva reticulata and Ulva fasciatatravelled from the mainland to Hawaii in this manner.Mapping is possible for select species only: "there are many valid examples of confined distribution patterns."[43]

For example, Clathromorphum is an arctic genus and is not mapped far south of there.[44] On the other hand,scientists regard the overall data as insufficient due to the "difficulties of undertaking such studies."[45]

Locations

Phytoplankton, Lake Chuzenji

Algae are prominent in bodies of water, common in terrestrialenvironments and are found in unusual environments, such as onsnow and on ice. Seaweeds grow mostly in shallow marine waters,under 100 metres (unknown operator: u'strong' ft); howeversome have been recorded to a depth of 360 metres (unknownoperator: u'strong' ft).[46]

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 foodchains. In very high densities (algal blooms) these algae maydiscolor the water and outcompete, poison, or asphyxiate other lifeforms.

Algae are variously sensitive to different factors, which has made them useful as biological indicators in theBallantine Scale and its modification.

Uses

Harvesting Algae

Agar

Agar, a gelatinous substance derived from red algae, hasa number of commercial uses.[47] It is a good medium forbacteria.

Alginates

Between 100,000 and 170,000 wet tons of Macrocystisare harvested annually in California for alginateextraction and abalone feed.[48][49]

Algae 10

Energy sourceTo be competitive and independent from fluctuating support from (local) policy on the long run, biofuels shouldequal or beat the cost level of fossil fuels. Here, algae based fuels hold great promise, directly related to the potentialto produce more biomass per unit area in a year than any other form of biomass. The break-even point foralgae-based biofuels is estimated to occur in about ten to fifteen years.[50]

Fertilizer

Seaweed is used as a fertilizer.

For centuries seaweed has been used as a fertilizer;George Owen of Henllys writing in the 16th centuryreferring to drift weed in South Wales:[51]

This kind of ore they often gather and lay ongreat heapes, where it heteth and rotteth, andwill have a strong and loathsome smell;when being so rotten they cast on the land,as they do their muck, and thereof springethgood corn, especially barley ... Afterspring-tydes or great rigs of the sea, theyfetch it in sacks on horse backes, and cariethe same three, four, or five miles, and castit on the lande, which doth very much betterthe ground for corn and grass.

Today Algae are used by humans in many ways; for example, as fertilizers, soil conditioners and livestock feed.[52]

Aquatic and microscopic species are cultured in clear tanks or ponds and are either harvested or used to treateffluents 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.

Nutrition

Seaweed gardens on Inisheer.

Naturally growing seaweeds are an important source offood, especially in Asia. They provide many vitaminsincluding: A, B1, B2, B6, niacin and C, and are rich iniodine, potassium, iron, magnesium and calcium.[53] Inaddition commercially cultivated microalgae, includingboth Algae and Cyanobacteria, are marketed asnutritional supplements, such as Spirulina,[54] Chlorellaand the Vitamin-C supplement, Dunaliella, high inbeta-carotene.

Algae are national foods of many nations: Chinaconsumes more than 70 species, including fat choy, acyanobacterium considered a vegetable; Japan, over 20

species;[55] Ireland, dulse; Chile, cochayuyo.[56] Laver is used to make "laver bread" in Wales where it is known asbara lawr; in Korea, gim; in Japan, nori and aonori. It is also used along the west coast of North America fromCalifornia to British Columbia, in Hawaii and by the Māori of New Zealand. Sea lettuce and badderlocks are a saladingredient in Scotland, Ireland, Greenland and Iceland.

Algae 11

Dulse, a food.

The oils from some Algae have high levels of unsaturated fattyacids. For example, Parietochloris incisa is very high inarachidonic acid, where it reaches up to 47% of the triglyceridepool.[57] Some varieties of Algae favored by vegetarianism andveganism contain the long-chain, essential omega-3 fatty acids,Docosahexaenoic acid (DHA) and Eicosapentaenoic acid(EPA), in addition to vitamin B12. The vitamin B12 in algae isnot biologically active. Fish oil contains the omega-3 fattyacids, but the original source is algae (microalgae inparticular), which are eaten by marine life such as copepodsand are passed up the food chain.[58] Algae has emerged inrecent years as a popular source of omega-3 fatty acids forvegetarians who cannot get long-chain EPA and DHA fromother vegetarian sources such as flaxseed oil, which onlycontains the short-chain Alpha-Linolenic acid (ALA).

Pollution control•• Sewage can be treated with algae, 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.• Aquariums and ponds can be filtered using algae, which absorb nutrients from the water in a device called an

Algae scrubber, also known as an "ATS".[59][60][61][62]

Agricultural Research Service scientists found that 60-90% of nitrogen runoff and 70-100% of phosphorus runoffcan be captured from manure effluents using an algal turf scrubber (ATS). Scientists developed the ATS, which areshallow, 100-foot raceways of nylon netting where algae colonies can form, and studied its efficacy for three years.They found that algae can readily be used to reduce the nutrient runoff from agricultural fields and increase thequality of water flowing into rivers, streams, and oceans. The enriched algae itself also can be used as a fertilizer.Researchers collected and dried the nutrient-rich algae from the ATS and studied its potential as an organic fertilizer.They found that cucumber and corn seedlings grew just as well using ATS organic fertilizer as they did withcommercial fertilizers.[63]

PigmentsThe natural pigments produced by algae can be used as an alternative to chemical dyes and coloring agents.[64]

Stabilizing substancesCarrageenan, from the red alga Chondrus crispus, is used as a stabilizer in milk products.

PlasticsAlgae has been implemented in the production of biodegradable plastics by Cereplast, Inc. An agreement has alsobeen reached with the US Military to introduce more biodegradable plastics as it attempts to move away frompetroleum based plastics and utilize more environmentally friendly alternatives.[65]

Algae 12

Notes[1] Patrick J. Keeling (2004). "Diversity and evolutionary history of plastids and their hosts" (http:/ / www. amjbot. org/ cgi/ content/ full/ 91/ 10/

1481). American Journal of Botany 91 (10): 1481–1493. doi:10.3732/ajb.91.10.1481. PMID 21652304. .[2] Laura Wegener Parfrey, Erika Barbero, Elyse Lasser, Micah Dunthorn, Debashish Bhattacharya, David J Patterson, and Laura A Katz

(December 2006). "Evaluating Support for the Current Classification of Eukaryotic Diversity". PLoS Genet. 2 (12): e220.doi:10.1371/journal.pgen.0020220. PMC 1713255. PMID 17194223.

[3] Nabors, Murray W. (2004). Introduction to Botany. San Francisco, CA: Pearson Education, Inc. ISBN 0-8053-4416-0.[4] Ed. Guiry, M.D., John, D.M., Rindi, F and McCarthy, T.K. 2007. New Survey of Clare Island Volume 6: The Freshwater and Terrestrial

Algae. Royal Irish Academy. ISBN 978-1-904890-31-7[5] Allaby, M ed. (1992). "Algae". The Concise Dictionary of Botany. Oxford: Oxford University Press.[6][6] Round (1981)[7] Smithsonian National Museum of Natural History; Department of Botany. http:/ / botany. si. edu/ projects/ algae/ introduction. htm[8] Bengtson, S.; Belivanova, V.; Rasmussen, B.; Whitehouse, M. (May 2009). "The controversial "Cambrian" fossils of the Vindhyan are real

but more than a billion years older". Proceedings of the National Academy of Sciences of the United States of America 106 (19): 7729–7734.Bibcode 2009PNAS..106.7729B. doi:10.1073/pnas.0812460106. ISSN 0027-8424. PMC 2683128. PMID 19416859.

[9] "alga, algae". Webster's Third New International Dictionary of the English Language Unabridged with Seven Language Dictionary. 1.Encyclopædia Britannica, Inc. 1986.

[10] Partridge, Eric (1983). "algae". Origins.[11] Lewis, Charlton T.; Charles Short (1879). "alga" (http:/ / www. perseus. tufts. edu/ cgi-bin/ ptext?layout. reflang=la;layout.

refdoc=Perseus:text:1999. 04. 0059;layout. reflookup=Alga;layout. refcit=;doc=Perseus:text:1999. 04. 0059:entry=#1812). alga. Oxford:Clarendon Press. ISBN 0-19-864201-6. .

[12] Cheyne, Thomas Kelly; John Sutherland Black (1899–1903). "Paint". Encyclopædia Biblica–A Dictionary of the Bible. 3. New York:Macmillan Co.. pp. 3524–3525. Downloadable Google Books (http:/ / books. google. com/ books?id=GccVAAAAYAAJ&pg=RA1-PA3525& dq=puk+ paint& lr=& ei=Ye9RSYPEAYjcygSSyrW3BA#PPA2689,M1).

[13] Losos, Jonathan B.; Mason, Kenneth A.; Singer, Susan R. (2007). Biology (8 ed.). McGraw-Hill. ISBN 0-07-304110-6.[14] Jochem, Frank J. "Botany 4404 Lecture Notes" (http:/ / www. jochemnet. de/ fiu/ bot4404/ BOT4404_12. html). Florida International

University (FIU). Archived (http:/ / web. archive. org/ web/ 20081225083828/ http:/ / www. jochemnet. de/ fiu/ bot4404/ BOT4404_12. html)from the original on 25 December 2008. . Retrieved 2008-12-20.

[15] Bhattacharya, D.; Medlin, L. (1998). "Algal Phylogeny and the Origin of Land Plants" (http:/ / www. plantphysiol. org/ cgi/ reprint/ 116/ 1/9. pdf). Plant Physiology 116 (1): 9–15. doi:10.1104/pp.116.1.9. .

[16] Burki F, Shalchian-Tabrizi K, Minge M, Skjæveland Å, Nikolaev SI, et al. (2007). Butler, Geraldine. ed. "Phylogenomics Reshuffles theEukaryotic Supergroups". PLoS ONE 2 (8: e790): e790. doi:10.1371/journal.pone.0000790. PMC 1949142. PMID 17726520.

[17] Dixon, P S (1973). Biology of the Rhodophyta. Edinburgh: Oliver & Boyd. p. 232. ISBN 0-05-002485-X.[18] Ivan Noble (18 September 2003). "When plants conquered land" (http:/ / news. bbc. co. uk/ 1/ hi/ sci/ tech/ 3117034. stm). BBC. .[19] Wellman, C.H.; Osterloff, P.L.; Mohiuddin, U. (2003). "Fragments of the earliest land plants". Nature 425 (6955): 282–285.

doi:10.1038/nature01884. PMID 13679913.[20] Xiao, S.; Knoll, A.H.; Yuan, X.; Pueschel, C.M. (2004). "Phosphatized multicellular algae in the Neoproterozoic Doushantuo Formation,

China, and the early evolution of florideophyte red algae" (http:/ / www. amjbot. org/ cgi/ content/ full/ 91/ 2/ 214). American Journal ofBotany 91 (2): 214–227. doi:10.3732/ajb.91.2.214. PMID 21653378. .

[21] Waggoner, Ben (1994–2008). "Introduction to the Phaeophyta: Kelps and brown "Algae"" (http:/ / www. ucmp. berkeley. edu/ chromista/phaeophyta. html). University of California Museum of Palaeontology (UCMP). Archived (http:/ / web. archive. org/ web/ 20081221171218/http:/ / www. ucmp. berkeley. edu/ chromista/ phaeophyta. html) from the original on 21 December 2008. . Retrieved 2008-12-19.

[22] Thomas, D N (2002). Seaweeds. London: The Natural History Museum. ISBN 0-565-09175-1.[23] Waggoner, Ben (1994–2008). "Introduction to the Rhodophyta, The red "algae"" (http:/ / www. ucmp. berkeley. edu/ protista/ rhodophyta.

html). University of California Museum of Palaeontology (UCMP). Archived (http:/ / web. archive. org/ web/ 20081218211021/ http:/ / www.ucmp. berkeley. edu/ protista/ rhodophyta. html) from the original on 18 December 2008. . Retrieved 2008-12-19.

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