Brown algae

The Phaeophyceae or brown algae (singular: alga), are

a large group of mostly marine multicellular algae, including many seaweeds of colder Northern Hemisphere waters. They play an important role in marine environments,
both as food and for the habitats they form. For instance,
Macrocystis, a kelp of the order Laminariales, may reach
60 m (200 ft) in length and forms prominent underwater forests. Another example is Sargassum, which creates
unique habitats in the tropical waters of the Sargasso Sea.
Many brown algae, such as members of the order Fucales,
commonly grow along rocky seashores. Some members
of the class, such as kelp, are used as food for humans.

in Ectocarpus, or of foot-long attened branches resembling a fan, as in Padina.

Regardless of size or form, two visible features set the

Phaeophyceae apart from all other algae. First, members of the group possess a characteristic color that ranges
from an olive green to various shades of brown. The particular shade depends upon the amount of fucoxanthin
present in the alga.[11] Second, all brown algae are
multicellular. There are no known species that exist as
single cells or as colonies of cells,[11] and the brown algae are the only major group of seaweeds that does not
include such forms. However, this may be the result of
Worldwide, about 15002000 species of brown algae classication rather than a consequence of evolution, as
are known.[4] Some species, such as Ascophyllum no- all the groups hypothesized to be the closest relatives of
dosum, are of sucient commercial importance that they the browns include single-celled or colonial forms.
have become subjects of extensive research in their own
right.[5]
Brown algae belong to a very large group, the 1.1
Heterokontophyta, a group of eukaryotic organisms distinguished most prominently by having chloroplasts surrounded by four membranes, suggesting an origin from
a symbiotic relationship between a basal eukaryote and
another eukaryotic organism. Most brown algae contain
the pigment fucoxanthin, which is responsible for the distinctive greenish-brown color that gives them their name.
Brown algae are unique among heterokonts in developing into multicellular forms with dierentiated tissues,
but they reproduce by means of agellated spores and
gametes that closely resemble cells of other heterokonts.
Genetic studies show their closest relatives to be the
yellow-green algae.

Morphology

Visible structures

Two specimens of Laminaria hyperborea, each showing the rootlike holdfast at lower left, a divided blade at upper right, and a
stemlike stipe connecting the blade to the holdfast.

Brown algae exist in a wide range of sizes and forms. The

smallest members of the group grow as tiny, feathery tufts
of threadlike cells no more than a few centimeters long.[6]
Some species have a stage in their life cycle that consists
of only a few cells, making the entire alga microscopic.
Other groups of brown algae grow to much larger sizes.
The rockweeds and leathery kelps are often the most conspicuous algae in their habitats.[7] Kelps can range in size
from the two-foot-tall sea palm Postelsia to the giant kelp
Macrocystis pyrifera, which grows to over 45 m (150 ft)
long[8][9] and is the largest of all the algae. In form, the
brown algae range from small crusts or cushions[10] to
leafy free-oating mats formed by species of Sargassum.
They may consist of delicate felt-like strands of cells, as

Further information: Thallus, Holdfast, Stipe (botany),

Lamina (algae), and Pneumatocyst
Whatever their form, the body of all brown algae is
termed a thallus, indicating that it lacks the complex
xylem and phloem of vascular plants. This does not mean
that brown algae completely lack specialized structures.
But, because some botanists dene true stems, leaves,
and roots by the presence of these tissues, their absence
in the brown algae means that the stem-like and leaf-like
structures found in some groups of brown algae must be
described using dierent terminology.[12] Although not
1

1 MORPHOLOGY

all brown algae are structurally complex, those that are

typically possess one or more characteristic parts.
A holdfast is a rootlike structure present at the base of the
alga. Like a root system in plants, a holdfast serves to anchor the alga in place on the substrate where it grows, and
thus prevents the alga from being carried away by the current. Unlike a root system, the holdfast generally does not
serve as the primary organ for water uptake, nor does it
take in nutrients from the substrate. The overall physical
appearance of the holdfast diers among various brown
algae and among various substrates. It may be heavily
branched, or it may be cup-like in appearance. A single
alga typically has just one holdfast, although some species Species like Fucus vesiculosus produce numerous gas-lled
have more than one stipe growing from their holdfast.
pneumatocysts (air bladders) to increase buoyancy.
A stipe is a stalk or stemlike structure present in an alga.
It may grow as a short structure near the base of the alga
(as in Laminaria), or it may develop into a large, complex structure running throughout the algal body (as in
Sargassum or Macrocystis). In the most structurally differentiated brown algae (such as Fucus), the tissues within
the stipe are divided into three distinct layers or regions.
These regions include a central pith, a surrounding cortex, and an outer epidermis, each of which has an analog
in the stem of a vascular plant. In some brown algae, the
pith region includes a core of elongated cells that resemble the phloem of vascular plants both in structure and
function. In others (such as Nereocystis), the center of the
stipe is hollow and lled with gas that serves to keep that
part of the alga buoyant. The stipe may be relatively exible and elastic in species like Macrocystis pyrifera that
grow in strong currents, or may be more rigid in species
like Postelsia palmaeformis that are exposed to the atmosphere at low tide.

or thick and leathery. In species like Egregia menziesii,

this characteristic may change depending upon the turbulence of the waters in which it grows.[6] In other species,
the surface of the blade is coated with slime to discourage the attachment of epiphytes or to deter herbivores.
Blades are also often the parts of the alga that bear the
reproductive structures.
Gas-lled oats called pneumatocysts provide buoyancy
in many kelps and members of the Fucales. These
bladder-like structures occur in or near the lamina, so that
it is held nearer the water surface and thus receives more
light for photosynthesis. Pneumatocysts are most often
spherical or ellipsoidal, but can vary in shape among different species. Species such as Nereocystis luetkeana and
Pelagophycus porra bear a single large pneumatocyst between the top of the stipe and the base of the blades. In
contrast, the giant kelp Macrocystis pyrifera bears many
blades along its stipe, with a pneumatocyst at the base of
each blade where it attaches to the main stipe. Species
of Sargassum also bear many blades and pneumatocysts,
but both kinds of structures are attached separately to the
stipe by short stalks. In species of Fucus, the pneumatocysts develop within the lamina itself, either as discrete
spherical bladders or as elongated gas-lled regions that
take the outline of the lamina in which they develop.

Many algae have a attened portion that may resemble a

leaf, and this is termed a blade, lamina, or frond. The
name blade is most often applied to a single undivided
structure, while frond may be applied to all or most of an
algal body that is attened, but this distinction is not universally applied. The name lamina refers to that portion
of a structurally dierentiated alga that is attened. It
may be a single or a divided structure, and may be spread
over a substantial portion of the alga. In rockweeds,
for example, the lamina is a broad wing of tissue that
runs continuously along both sides of a branched midrib. 1.2 Growth
The midrib and lamina together constitute almost all of
a rockweed, so that the lamina is spread throughout the The brown algae include the largest and fastest growing
of seaweeds.[6] Fronds of Macrocystis may grow as much
alga rather than existing as a localized portion of it.
as 50 centimetres (20 in) per day, and the stipes can grow
In some brown algae, there is a single lamina or blade,
6 centimetres (2.4 in) in a single day.[13]
while in others there may be many separate blades. Even
in those species that initially produce a single blade, the Growth in most brown algae occurs at the tips of strucstructure may tear with rough currents or as part of mat- tures as a result of divisions in a single apical cell or in a
[7]
uration to form additional blades. These blades may be row of such cells. As this apical cell divides, the new
attached directly to the stipe, to a holdfast with no stipe cells that it produces develop into all the tissues of the
present, or there may be an air bladder between the stipe alga. Branchings and other lateral structures appear when
and blade. The surface of the lamina or blade may be the apical cell divides to produce two new apical cells.
smooth or wrinkled; its tissues may be thin and exible However, a few groups (such as Ectocarpus) grow by a
diuse, unlocalized production of new cells that can oc-

2.1

Fossils

Growth in Dictyota dichotoma occurs at each frond tip, where

new cells are produced.

3
lar ancestors.[18] DNA sequence comparison also suggests that the brown algae evolved from the lamentous Phaeothamniophyceae,[19] Xanthophyceae,[20] or the
Chrysophyceae[21] between 150[1] and 200 million years
ago.[2] In many ways, the evolution of the brown algae parallels that of the green algae and red algae,[22]
as all three groups possess complex multicellular species
with an alternation of generations. Analysis of 5S rRNA
sequences reveals much smaller evolutionary distances
among genera of the brown algae than among genera of
red or green algae,[2][23] which suggests that the brown
algae have diversied much more recently than the other
two groups.

2.1 Fossils
cur anywhere on the thallus.[11]

1.3

Tissue organization

The simplest browns are lamentousthat is, their cells

are elongate and have septa cutting across their width.
They branch by getting wider at their tip, and then dividing the widening.[14]
Aside from lamentous forms, there are two main
types of tissue organization in the brown algae: pseudoparenchymatous (haplostichous) and parenchymatous
(polystichous).[15] The fronds may be multiaxial or
monoaxial.

The occurrence of Phaeophyceae as fossils is rare due to

their generally soft-bodied nature,[24] and scientists continue to debate the identication of some nds.[25] Part
of the problem with identication lies in the convergent
evolution of morphologies between many brown and red
algae.[26] Most fossils of soft-tissue algae preserve only
a attened outline, without the microscopic features that
permit the major groups of multicellular algae to be reliably distinguished. Among the brown algae, only species
of the genus Padina deposit signicant quantities of minerals in or around their cell walls.[27] Other algal groups,
such as the red algae and green algae, have a number
of calcareous members. Because of this, they are more
likely to leave evidence in the fossil record than the soft
bodies of most brown algae and more often can be precisely classied.[28]

The cell wall consists of two layers; the inner layer bears
the strength, and consists of cellulose; the outer wall layer
is mainly algin, and is gummy when wet but becomes hard Fossils comparable in morphology to brown algae are
known from strata as old as the Upper Ordovician,[29]
and brittle when it dries out.[15]
but the taxonomic anity of these impression fossils is
far from certain.[30] Claims that earlier Ediacaran fossils
are brown algae[31] have since been dismissed.[19] While
2 Evolutionary history
many carbonaceous fossils have been described from the
Precambrian, they are typically preserved as attened
Genetic and ultrastructural evidence place the Phaeo- outlines or fragments measuring only millimeters long.[32]
phyceae among the heterokonts (Stramenopiles),[16] Because these fossils lack features diagnostic for idena large assemblage of organisms that includes both tication at even the highest level, they are assigned to
photosynthetic members with plastids (such as the fossil form taxa according to their shape and other gross
diatoms) as well as non-photosynthetic groups (such as morphological features.[33] A number of Devonian fosthe slime nets and water molds). Although some het- sils termed fucoids, from their resemblance in outline
erokont relatives of the brown algae lack plastids in their to species in the genus Fucus, have proven to be inorcells, scientists believe this is a result of evolutionary loss ganic rather than true fossils.[24] The Devonian megafosof that organelle in those groups rather than independent sil Prototaxites, which consists of masses of laments
acquisition by the several photosynthetic members.[17] grouped into trunk-like axes, has been considered a possiThus, all heterokonts are believed to descend from a sin- ble brown alga.[11] However, modern research favors reingle heterotrophic ancestor that became photosynthetic terpretation of this fossil as a terrestrial fungus or fungalwhen it acquired plastids through endosymbiosis of an- like organism.[34] Likewise, the fossil Protosalvinia was
other unicellular eukaryote.[4]
once considered a possible brown alga, but is now thought
[35]
The closest relatives of the brown algae include unicel- to be an early land plant.
lular and lamentous species, but no unicellular species A number of Paleozoic fossils have been tentatively
of brown algae are known. However, most scientists classied with the brown algae, although most have
assume that the Phaeophyceae evolved from unicellu- also been compared to known red algae species.

Phascolophyllaphycus possesses numerous elongate, inated blades attached to a stipe. It is the most abundant of algal fossils found in a collection made from
Carboniferous strata in Illinois.[36] Each hollow blade
bears up to eight pneumatocysts at its base, and the stipes
appear to have been hollow and inated as well. This
combination of characteristics is similar to certain modern genera in the order Laminariales (kelps). Several fossils of Drydenia and a single specimen of Hungerfordia
from the Upper Devonian of New York have also been
compared to both brown and red algae.[26] Fossils of
Drydenia consist of an elliptical blade attached to a
branching lamentous holdfast, not unlike some species
of Laminaria, Porphyra, or Gigartina. The single known
specimen of Hungerfordia branches dichotomously into
lobes and resembles genera like Chondrus and Fucus[26]
or Dictyota.[37]
The earliest known fossils that can be assigned reliably
to the Phaeophyceae come from Miocene diatomite deposits of the Monterey Formation in California.[4] Several
soft-bodied brown macroalgae, such as Julescraneia, have
been found.[38]

CLASSIFICATION

Family Petrodermataceae Silberfeld,

Rousseau & Reviers 2014

Family Ishigeaceae Okamura 1935

Subclass Dictypophycidae Silberfeld, Rousseau
& Reviers 2014

Order Syringodermatales Henry 1984

Family Syringodermataceae Henry
1984

Order Onslowiales Draisma & Prudhomme

van Reine 2008

Family Onslowiaceae Draisma &

Prudhomme van Reine 2001

Order Dictyotales Bory de Saint-Vincent

1828 ex Phillips et al.

Family Dictyotaceae Lamouroux ex

Dumortier 1822 [Scoresbyellaceae
Womersley 1987; Dictyopsidaceae]
Order Sphacelariales Migula 1909
Family Lithodermataceae Hauck 1883
Family Phaeostrophiaceae Kawai et
al. 2005

Family Sphacelodermaceae Draisma,

Prudhomme & Kawai 2010

Classication

3.1

Family

Phylogeny

1922

Family

Sphacelariaceae

Decaisne

1842

Subclass Fucophycidae Cavalier-Smith 1986

Taxonomy

Order Desmarestiales Setchell & Gardner

Further information: List of brown algal genera and

Wikispecies:Phaeophyceae
This is a list of
Phaeophyceae:[39][40]

Oltmanns

Family Cladostephaceae Oltmanns

Based on the work of Silberfeld, Rousseau & de Reviers

2014.[39]

3.2

Stypocaulaceae

1922

the

orders

in

the

class

Class Phaeophyceae Hansgirg 1886 [Fucophyceae;

Melanophycidae Rabenhorst 1863 stat. nov. CavalierSmith 2006]
Subclass Discosporangiophycidae Silberfeld,
Rousseau & Reviers 2014

Order Discosporangiales Schmidt 1937

emend. Kawai et al. 2007

Family Choristocarpaceae Kjellman

1891

Family Discosporangiaceae Schmidt

1937

Subclass Ishigeophycidae Silberfeld, Rousseau

& Reviers 2014

Order Ishigeales Cho & Boo 2004

1925

Family Arthrocladiaceae Chauvin

1842

Family

Desmarestiaceae

(Thuret)

Kjellman 1880

Order Sporochnales Sauvageau 1926

Family Sporochnaceae Greville 1830
Order Ascoseirales Petrov1964 emend.
Moe & Henry 1982

Family Ascoseiraceae Skottsberg 1907

Order Scytothamnales Peters & Clayton
1998 emend. Silberfeld et al. 2011

Family Asteronemataceae Silberfeld

et al. 2011

Family Bachelotiaceae Silberfeld et al.

2011

Family Splachnidiaceae Mitchell &

Whitting 1892 [Scytothamnaceae
Womersley 1987]
Order
Laminariales
Migula
1909
[Phaeosiphoniellales Silberfeld, Rousseau
& Reviers 2014 ord. nov. prop.]

5
Family Phaeosiphoniellaceae Phillips
et al. 2008

Family Akkesiphycaceae Kawai &

Sasaki 2000

Family Pseudochordaceae Kawai &

Kurogi 1985

Family Chordaceae Dumortier 1822

Family Agaraceae Postels & Ruprecht
1840 [Costariaceae]
Family Laminariaceae Bory de SaintVincent 1827 [Arthrothamnaceae
Petrov 1974]
Family Aureophycaceae Kawai &
Ridgway 2013

Family Alariaceae Setchell & Gardner

1925 [Lessoniaceae Setchell & Gardner
1925]
Order Asterocladales T.Silberfeld et al.
2011

Family Asterocladaceae Silberfeld et

al. 2011

Order Ectocarpales Bessey 1907 emend.

Rousseau & Reviers 1999a [Chordariales
Setchell & Gardner 1925; Dictyosiphonales
Setchell & Gardner 1925; Scytosiphonales
Feldmann 1949]
Family ?Sorocarpaceae
Family Adenocystaceae Rousseau et
al. 2000 emend. Silberfeld et al. 2011

[Chordariopsidaceae]
Family Scytosiphonaceae Ardissone
& Straforello 1877 [Chnoosporaceae
Setchell & Gardner 1925]
Family Petrospongiaceae Racault et al.
2009

Family Ectocarpaceae Agardh 1828

emend. Silberfeld et al. 2011

Family Acinetosporaceae Hamel ex

Feldmann 1937 [Pylaiellaceae; Pilayellaceae]
Family Chordariaceae Greville 1830
emend. Peters & Ramrez 2001 [Myrionemataceae]
Order Stschapoviales
Family Stschapoviaceae Kawai 2004
Family Halosiphonaceae Kawai &
Sasaki 2000

Family Cutleriaceae Grith & Henfrey 1856

Order Ralfsiales Nakamura ex Lim & Kawai

2007

Family Mesosporaceae Tanaka & Chihara 1982

Family Neoralfsiaceae Lim & Kawai

2007

Family Ralfsiaceae Farlow 1881 [Heterochordariaceae Setchell & Gardner

1925]
Order Nemodermatales Parente et al. 2008
Family Nemodermataceae Kuckuck
ex Feldmann 1937

Order Fucales Bory de Saint-Vincent 1827

[Notheiales Womersley 1987; Durvillaeales Petrov 1965]
Family Bifurcariopsidaceae Cho et al.
2006

Family Hormosiraceae Fritsch 1945

Family Notheiaceae Schmidt 1938
Family Seirococcaceae Nizamuddin
1987

Family Xiphophoraceae Cho et al.

2006

Family Sargassaceae Ktzing 1843

[Cystoseiraceae De Toni 1891]
Family Durvillaeaceae (Oltmanns) De
Toni 1891

Family Himanthaliaceae (Kjellman)

De Toni 1891

Family Fucaceae Adanson 1763

4 Life cycle
Sexual reproduction may be isogamous, oogamous, or
anisogamous. There is evidence of sex chromosomes.[41]
Union of gametes may take place in water or within the
oogonium (oogamous species). The life cycle shows great
variability from one group to another.
In Laminaria, there is a conspicuous diploid generation.
Meiosis takes place within sporangia, before the spores
are released. As they are haploid, with sex chromosomes,
there are equal numbers of male and female spores.[42]
With the exception of the Fucales, all brown algae have a
life cycle with an alternation between haploid and diploid
forms.

Family Platysiphonaceae
Order Tilopteridales Bessey 1907 emend.
Phillips et al. 2008 [Cutleriales Bessey
1907]
5 Ecology
Family ?Masonophycaceae
Family Tilopteridaceae Kjellman 1890 Brown algae have adapted to a wide variety of marine
Family Phyllariaceae Tilden 1935
ecological niches including the tidal splash zone, rock

REFERENCES

8 See also
Wrack (seaweed)

9 References
[1] Medlin, L. K.; et al. (1997). Phylogenetic relationships
of the 'golden algae' (haptophytes, heterokont chromophytes) and their plastids (PDF). Plant Systematics and
Evolution. Plant Systematics and Evolution. 11: 187
219. doi:10.1007/978-3-7091-6542-3_11. ISBN 978-3211-83035-2. hdl:10013/epic.12690.

Saccharina latissima on a beach.

pools, the whole intertidal zone and relatively deep near

shore waters. They are an important constituent of some
brackish water ecosystems, and four species are restricted
to life in fresh water.[19] A large number of Phaeophyceae
are intertidal or upper littoral,[19] and they are predominantly cool and cold water organisms that benet from
nutrients in up welling cold water currents and inows
from land; Sargassum being a prominent exception to this
generalisation.
Brown algae growing in brackish waters are almost solely
asexual.[19]

Chemistry

Brown algae have a 13 C value in the range of 20.8

to 10.5, in contrast with red algae and greens. This
reects their dierent metabolic pathways.[44]
They have cellulose walls with alginic acid and also contain the polysaccharide fucoidan in the amorphous sections of their cell walls. A few species (of Padina) calcify
with aragonite needles.[19]

[2] Lim, B.-L.; Kawai, H.; Hori, H.; Osawa, S. (1986).

Molecular evolution of 5S ribosomal RNA from red and
brown algae. Japanese Journal of Genetics. 61 (2): 169
176. doi:10.1266/jjg.61.169.
[3] Kjellman, F. R. (1891). Phaeophyceae (Fucoideae)". In
Engler, A.; Prantl, K. Die natrlichen Panzenfamilien. 1.
Leipzig: Wilhelm Engelmann. pp. 176192.
[4] van den Hoek, C.; Mann, D. G.; Jahns, H. M. (1995).
Algae: An Introduction to Phycology.
Cambridge:
Cambridge University Press. pp. 165218. ISBN 0-52131687-1.
[5] Senn, T. L. (1987). Seaweed and Plant Growth. Clemson,
S. C.: T. L. Senn. ISBN 0-939241-01-3.
[6] Connor, J.; Baxter, C. (1989). Kelp Forests. Monterey
Bay Aquarium. ISBN 1-878244-01-9.
[7] Dittmer, H. J. (1964). Phylogeny and Form in the Plant
Kingdom. Princeton, NJ: D. Van Nostrand Company. pp.
115137. ISBN 0-88275-167-0.
[8] Abbott, I. A.; Hollenberg, G. J. (1976). Marine Algae of
California. California: Stanford University Press. ISBN
0-8047-0867-3.
[9] Cribb, A. B. (1953).
"Macrocystis pyrifera (L.)
Ag. in Tasmanian waters. Australian Journal of
Marine and Freshwater Research.
5 (1): 134.
doi:10.1071/MF9540001.
[10] Jones, W. E. (1962). A key to the genera of the British

The photosynthetic system of brown algae is made of

seaweeds (PDF). Field Studies. 1 (4): 132.
a P700 complex containing chlorophyll a. Their plastids also contain chlorophyll c and carotenoids (the most [11] Bold, H. C.; Alexopoulos, C. J.; Delevoryas, T. (1987).
Morphology of Plants and Fungi (5th ed.). New York:
widespread of those being fucoxanthin).[45]
Brown algae produce a specic type of tannin called
phlorotannins in higher amounts than red algae do.

Importance and uses

Harper & Row Publishers. pp. 112131, 174186. ISBN

0-06-040839-1.

[12] Raven, P. H.; Evert, R. F.; Eichhorn, S. E. (2005). Biology of Plants (7th ed.). New York: W. H. Freeman and
Company. pp. 316321, 347. ISBN 0-7167-1007-2.
[13] Round, F. E. (1981). The Ecology of Algae. Cambridge:
Cambridge University Press. p. 103. ISBN 0-521-269067.

The brown algae include a number of edible seaweeds,

and a number of brown algae contains alginic acid that [14] Wynne, M. J. (1981). The Biology of seaweeds. In
is extracted commercially and used in a large number of
Lobban, C. S.; Wynne, M. J. Phaeophyta: Morphology
products. Sargachromanol G, found in brown algae, may
and Classication. Botanical Monographs. 17. University
be anti-inammatory.[46]
of California Press. p. 52. ISBN 978-0-520-04585-9.

[15] Sharma, O. P (1986). Textbook of Algae. Tata McGrawHill. p. 298. ISBN 978-0-07-451928-8.

[30] Speer, B. R.; Waggoner, B. M. (2000). Phaeophyta: Fossil Record.

[16] Adl, S. M.; et al. (2005). The new higher level classication of eukaryotes with emphasis on the taxonomy of
protists (PDF). Journal of Eukaryotic Microbiology. 52
(5): 399451. doi:10.1111/j.1550-7408.2005.00053.x.
PMID 16248873.

[31] Loeblich, A. R. (1974). Protistan Phylogeny as Indicated by the Fossil Record. Taxon. 23 (2/3): 277290.
doi:10.2307/1218707. JSTOR 1218707.

[17] Lane, C. E.; Archibald, J. M. (2008). The eukaryotic tree of life: Endosymbiosis takes its TOL (PDF).
Trends in Ecology and Evolution. 23 (5): 268275.
doi:10.1016/j.tree.2008.02.004. PMID 18378040.
[18] Niklas, K. J. (1997). The Evolutionary Biology of Plants.
Chicago: University of Chicago Press. p. 156. ISBN 0226-58082-2.
[19] Lee, R. E. (2008). Phycology (4th ed.). Cambridge University Press. ISBN 978-0-521-63883-8.
[20] Ariztia, E. V.; Andersen, R. A.; Sogin, M. L.
(1991). A new phylogeny of chromophyte algae using 16S-like rRNA sequences from Mallomonas papillosa (Synurophyceae) and Tribonema aequale (Xanthophyceae)". Journal of Phycology. 27 (3): 428436.
doi:10.1111/j.0022-3646.1991.00428.x.
[21] Taylor, T. N.; Taylor, E. L. (1993). The Biology and Evolution of Fossil Plants. Englewood Clis, NJ: Prentice
Hall. pp. 128131. ISBN 0-13-651589-4.
[22] Dittmer, H. J. (1964). Phylogeny and Form in the Plant
Kingdom. Princeton, NJ: D. Van Nostrand Company. pp.
115137. ISBN 0-88275-167-0.
[23] Hori, H.; Osawa, S. (1987). Origin and evolution of organisms as deduced from 5S ribosomal RNS sequences
(PDF). Molecular Biology and Evolution. 4 (5): 445472.
PMID 2452957.
[24] Arnold, C. A. (1947). An Introduction to Paleobotany.
New York; London: McGraw-Hill. p. 48. ISBN 1-40671861-0.
[25] Coyer, J. A.; Smith, G. J.; Andersen, R. A. (2001).
Evolution of Macrocystis spp. (Phaeophyta) as determined by ITS1 and ITS2 sequences (PDF). Journal
of Phycology. 37 (4): 574585. doi:10.1046/j.15298817.2001.037001574.x.

[32] Hofmann, H. J. (1985). Precambrian Carbonaceous

Megafossils. In D. F. Toomey & M. H. Nitecki. Paleoalgology: Contemporary Research and Applications. Berlin:
Springer-Verlag. pp. 2033.
[33] Hofmann, H. J. (1994). Proterozoic carbonaceous compressions (metaphytes and worms)". In Bengtson,
S. Life on Earth. Nobel Symposium. 84. New York:
Columbia University Press. pp. 342357.
[34] Hueber, F. M. (2001). Rotted wood-alga-fungus: the
history and life of Prototaxites Dawson 1859. Review
of Palaeobotany and Palynology. 116 (1): 123158.
doi:10.1016/S0034-6667(01)00058-6.
[35] Taylor, W. A.; Taylor, T. N. (1987). Spore wall ultrastructure of Protosalvinia" (PDF). American Journal of
Botany. 74 (3): 437433. doi:10.2307/2443819. JSTOR
2443819.
[36] Leary, R. L. (1986). Three new genera of fossil noncalcareous algae from Valmeyeran (Mississippian) strata of
Illinois. American Journal of Botany. 73 (3): 369375.
doi:10.2307/2444080. JSTOR 2444080.
[37] Bold, H. C.; Wynne, M. J. (1978). Introduction to the Algae (2nd ed.). Prentice-Hall. p. 27. ISBN 0-13-4777867.
[38] Parker, B. C.; Dawson, E. Y. (1965). Non-calcareous
marine algae from California Miocene deposits. Nova
Hedwigia. 10: 273295; plates 7696.
[39] Silberfeld, Thomas; Rousseau, Florence; de Reviers, Bruno (2014).
An Updated Classication
of Brown Algae (Ochrophyta, Phaeophyceae)".
Cryptogamie, Algologie. BioOne. pp. 117156.
doi:10.7872/crya.v35.iss2.2014.117. Retrieved 201610-10.
[40] Guiry, M. D.; Guiry, G. M. (2009). AlgaeBase.
National University of Ireland. Retrieved 2012-12-31.

[26] Fry, W. L.; Banks, H. P. (1955). Three new genera of

algae from the Upper Devonian of New York. Journal
of Paleontology. 29: 3744. JSTOR 1300127.

[41] Lewis, R.J. (1996). Chromosomes of the brown algae.

Phycologia. 35 (1): 1940. doi:10.2216/i0031-8884-351-19.1.

[27] Prescott, G. W. (1968). The Algae: A Review. Boston:

Houghton Miin Company. pp. 207231, 371372.
ISBN 3-87429-244-4.

[42] Thomas, D. N. (2002). Seaweeds. London: Natural History Museum, London. ISBN 0-565-09175-1.

[28] Simpson, G. G. (1953). Life of the Past: An Introduction

to Paleontology. New Haven: Yale University Press. pp.
158159.

[43] Maberly, S. C.; Raven, J. A.; Johnston, A. M. (1992).

Discrimination between 12 C and 13 C by marine plants.
Oecologia. 91 (4): 481. doi:10.1007/BF00650320.
JSTOR 4220100.

[29] Fry, W. L. (1983). An algal ora from the Upper Ordovician of the Lake Winnipeg region, Manitoba, Canada.
Review of Palaeobotany and Palynology. 39 (34): 313
341. doi:10.1016/0034-6667(83)90018-0.

[44] Fletcher, B. J.; Beerling, D. J.; Chaloner, W. G. (2004).

Stable carbon isotopes and the metabolism of the terrestrial Devonian organism Spongiophyton. Geobiology. 2
(2): 107119. doi:10.1111/j.1472-4677.2004.00026.x.

10

[45] Subunit organization of PSI particles from brown algae

and diatoms: polypeptide and pigment analysis. Claire
Berkalo, Lise Caron and Bernard Rousseau, Photosynthesis Research, February 1990, Volume 23, Issue 2,
pages 181-193, doi:10.1007/BF00035009
[46] Yoon, WJ.; Heo, SJ.; Han, SC.; Lee, HJ.; Kang, GJ.;
Kang, HK.; Hyun, JW.; Koh, YS.; Yoo, ES. (Aug 2012).
Anti-inammatory eect of sargachromanol G isolated
from Sargassum siliquastrum in RAW 264.7 cells. Arch
Pharm Res. 35 (8): 14211430. doi:10.1007/s12272012-0812-5. PMID 22941485.

10

External links

Monterey Bay Flora

The Monterey Formation of California, University
of California Museum of Paleontology
Phaeophyceae, National University of Ireland, Galway

EXTERNAL LINKS

11
11.1

Text and image sources, contributors, and licenses

Text

Brown algae Source: https://en.wikipedia.org/wiki/Brown_algae?oldid=757214950 Contributors: Josh Grosse, William Avery, Bkonrad,
Andycjp, Bender235, Syp, Vzb83~enwiki, Arcadian, Alansohn, Atlant, Plumbago, Riana, Snowolf, Velella, Stemonitis, Camw, Ashmoo,
BD2412, Rjwilmsi, Vegaswikian, Nihiltres, Maire, Dj Capricorn, YurikBot, Borgx, Anomalocaris, Curtis Clark, Welsh, Imaninjapirate,
Argo Navis, SmackBot, EncycloPetey, Gilliam, Ziiv, Persian Poet Gal, Abyssal, MrPMonday, Thor Dockweiler, SashatoBot, Butko, Mgiganteus1, Smith609, Tawkerbot2, JForget, Friendly Neighbour, Lavateraguy, JohnCD, Tom David, BetacommandBot, Thijs!bot, Osborne,
Headbomb, Sturm55, AntiVandalBot, KP Botany, Rie, JAnDbot, DuncanHill, Db099221, Easchi, Grievous Angel, VoABot II, Adacus12, Nono64, Werothegreat, Wlodzimierz, DorganBot, Idioma-bot, TXiKiBoT, Rei-bot, Corvus cornix, Paucabot, SieBot, YonaBot,
Oxymoron83, ClueBot, NickCT, Plastikspork, Alexbot, Abrech, Muro Bot, Darthvader102938, Leo415, Addbot, DOI bot, Raywil, Wang
lvan, SpBot, Tassedethe, Lightbot, ScAvenger, Jarble, Yobot, AnomieBOT, Jim1138, Materialscientist, Citation bot, Xqbot, RibotBOT,
ChristsAmbassador, FrescoBot, LucienBOT, Citation bot 1, Pinethicket, Fucoidan08, RjwilmsiBot, Gould363, EmausBot, Racerx11,
MadhATTER6, RA0808, Klbrain, Solarra, Wikipelli, Solomonfromnland, Citron, Donner60, DASHBotAV, Kleopatra, Berberisb, ClueBot NG, Ahui21, Widr, Helpful Pixie Bot, Todobo, Videsh Ramsahai, Deuterostome, Achowat, Haminiwiki, YFdyh-bot, Oskoreii, Dexbot,
Sminthopsis84, Cerabot~enwiki, EauOo, BarbBarbBarb, Zorahia, Kiruning, Gabrielaeverett, Hamoudafg, AddWittyNameHere, Buttery
Smith, Jts1882, Monkbot, Algaelover, KasparBot, Caftaric, InternetArchiveBot, Riding69 and Anonymous: 119

11.2

Images

File:Bladder_Wrack_(Fucus_vesiculosus)_-_geograph.org.uk_-_224125.jpg Source:
https://upload.wikimedia.org/wikipedia/
commons/8/80/Bladder_Wrack_%28Fucus_vesiculosus%29_-_geograph.org.uk_-_224125.jpg License: CC BY-SA 2.0 Contributors:
From geograph.org.uk Original artist: Anne Burgess
File:Capo_Gallo_Dicotoma.jpg Source: https://upload.wikimedia.org/wikipedia/commons/0/0b/Capo_Gallo_Dicotoma.jpg License:
CC BY 3.0 Contributors: Own work (personal work) Original artist: albertoromeo@neomedia.it
http://www.romeofotosub.it
File:Caribou_from_Wagon_Trails.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/e0/Caribou_from_Wagon_Trails.
jpg License: CC BY-SA 2.5 Contributors: ? Original artist: ?
File:Commons-logo.svg Source: https://upload.wikimedia.org/wikipedia/en/4/4a/Commons-logo.svg License: PD Contributors: ? Original artist: ?
File:Issoria_lathonia.jpg Source: https://upload.wikimedia.org/wikipedia/commons/2/2d/Issoria_lathonia.jpg License: CC-BY-SA-3.0
Contributors: ? Original artist: ?
File:Laminaria_hyperborea.jpg Source: https://upload.wikimedia.org/wikipedia/commons/0/03/Laminaria_hyperborea.jpg License:
CC BY-SA 3.0 Contributors: My Canon A95 Original artist: Sergey S. Dukachev
File:Rose_Amber_Flush_20070601.jpg Source:
https://upload.wikimedia.org/wikipedia/commons/3/37/Rose_Amber_Flush_
20070601.jpg License: CC BY-SA 3.0 Contributors: Own work Original artist: Georges Seguin (Okki)
File:Saccharina_latissima_NOAA.jpg Source: https://upload.wikimedia.org/wikipedia/commons/c/cd/Saccharina_latissima_NOAA.
jpg License: Public domain Contributors: NOAA Photo Library Original artist: David Csepp, NOAA/NMFS/AKFSC/Auke Bay Lab
File:Wikispecies-logo.svg Source: https://upload.wikimedia.org/wikipedia/commons/d/df/Wikispecies-logo.svg License: CC BY-SA
3.0 Contributors: Image:Wikispecies-logo.jpg Original artist: (of code) cs:User:-xfi File:Yellow.tang.arp.jpg Source: https://upload.wikimedia.org/wikipedia/commons/a/a0/Yellow.tang.arp.jpg License: Public domain
Contributors: Photographed by Adrian Pingstone in December 2005 Original artist: Adrian Pingstone

11.3

Content license

Creative Commons Attribution-Share Alike 3.0