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Photosynthetic organisms inhabiting the water column
are called
phytoplankton; they include
microscopic unicellular algae and photosynthetic
bacteria. There are a great number of them in the water.
At the end of the summer - beginning of
autumn, when surface water of Black Sea is the
warmest, there are can be between thousands to ten
millions of phytoplankton cells in 1 liter of seawater
at Caucasian coast. Since these creatures are very small
(their size ranging from several to hundreds microns)
this large amount of phytoplankton corresponds to a very
small weight: one million cells of Black Sea
phytoplankton weighs only one half of a gram.
There can be ten, even hundred times more
phytoplankton cells in water in the Western part of the
Black Sea, which is well fertilized by large rivers,
such as the Danube. |
 |
Putting together the biomass of all the phytoplankton in
Black Sea in a usual day of August gives an astronomic number
of about six millions ton. This photosynthesizing mass (called
standing crop) is growing (primary production), and - at the
same time - disappearing due to grazing by zooplankton.
This great mass is hard to visualize, but the value itself
may help understanding the role of planktonic algae and
bacteria in the Sea life: it's the main role. The ecology of
the Black Sea is primarily plankton ecology.
Ceratium tripos reaching
400 µm
(almost half of a millimeter, still
invisible without microscope) is one of the largest
phytoplankton species
in the Black Sea, it's
a giant microalgae |
This is not only
true for the Black Sea but also for the ocean in
general.
The words 'marine plants' usually make us remember
grass-like or bush-like seaweeds. They grow only at
relatively shallow depth near the sea shore, since they
require a hard substrate to settle on, and at the same
time require light. Because of this marine
macrophytes inhabit the bottom slope to the depth of
40-50m in the Black Sea where water transparency is
relatively low (Key Features
of the Black Sea), and to the depth of 100
meters in more transparent waters.
Microscopic phytoplankton inhabit surface waters of
the whole sea: from coast to coast, to the depth where
light is sufficient for algal photosynthesis to prevail
over algal respiration (they breathe too).
That sunlit upper layer of the sea is called the
photic zone.
In the Black Sea it is no deeper
then 100m; usually most of the phytoplankton is found in
the upper 0-60m layer of the Sea.
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On the other hand, phytoplankton cells can grow and
proliferate very fast. Many species of planktonic
microalgae and bacteria are capable of doubling their
biomass during 24 hours.
Because of this, very small, but abundant and
fast-growing phytoplankton are the most important marine
plants and is the base for Black Sea life.
Phytoplankton captures light that the sun is
generously pouring into the Sea. The power of this
light channeled through the photosynthetic machine of
the chloroplasts, turns non-living matter (salts, water,
carbon dioxide) of the environment into the new living
matter of algal cells.
In terms of ecology all this is called primary
production. Zooplankton feeds on phytoplankton
- and add its living matter to its own mass, growing and
proliferating in its turn, using the same energy of
sunlight stored in organic molecules made by
phytoplankton. This is a part of overall secondary
production, growth of the whole pelagic
community. Then small zooplankters are being
devoured by the larger ones and by planktivourous fish,
and so on, along the pelagic food chain all the way to
the large predatory fish, dolphins, marine birds, and
fishermen. And then inevitably the turn of
reduction comes: everything that is born, grow,
and live - eventually die, and the remains of pelagic
living beings, both of plankton and nekton, sink to
deeper water, turning into detritus.
The remains of marine life become food for marine
saprophytic bacteria. In a special case of the Black Sea
absolute majority of saprophytic bacterioplankton
is represented by anaerobic bacteria inhabiting anoxic
zone of the Sea below 200m depth. Saprophytic bacteria
decompose organic remains, returning the matter back to
inorganic state, making it available again for the use
by phytoplankton. Again, in the particular case of the
Black Sea, it takes a very long time - many years
- before the regenerated inorganic nutrients can
become available to phytoplankton, because of the
impeded vertical circulation in the distinctly
stratified Black Sea.
Most of the Black Sea ecosystem production takes
place above the picnocline, in its less saline, sunlit
upper water layer, whereas most of bacterial reduction
occur under the picnocline (50-100m) - in the dark,
anoxic, denser water of deep Black Sea (Key Features of the Black
Sea). |
| Phytoplankton includes
photosynthesizing bacteria as well as algae; most
of them are cyanobacteria.
Earlier they were also called blue-green algae,
but they are bacteria, prokaryotes: there are no
nuclei in their cells.
Cyanobacteria occur mostly in coastal waters of
Black Sea, particularly near river estuaries;
there are more cyanobacteria in the brackish,
eutrophied Sea of Azov. Many cyanobacteria are
capable of producing toxins harmful to marine
fauna. |
Colonies of planktonic
cyanobacteria;
nearshore sample, Caucasian
coast of Black Sea;
a scanning electron microscope
image.
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All planktonic autotrophs are fragile unicellular
organisms, and there are so many agile predators
swimming around in seawater; environmental conditions
sometimes become very rough for phytoplankton. Then
how do they manage to survive? The answer is that
they are not able to securely protect themselves,
but they do prolong their living enough
to have time to reproduce. When the environmental
conditions become unfavorable, many phytoplankton
species form hibernating cells - cysts, spores.
Many planktonic algae are mobile, which helps them to
escape zooplankton grazers: they have flagella, some
have one (e.g. Euglenophycean algae), some, like
dinoflagellates, have two, and green Prasinophycean
algae may have up to eight flagella. Those unicellular
plants
rush about their small world almost as vividly as
animals hunting them.
Also many of phytoplankton algae bear hard armor,
exoskeleton called theca, which support the cell shape
and provide them a protection from grazers and parasites
in some instances. Of course theca is of no help when
algal cell is swallowed by a fish or absorbed by a
bivalve mollusk.
Black Sea phytoplankton includes no less then 600
species; here we will look at the planktonic
algae that are most important for the life in the
Black Sea, or - just interesting ones. More
attention - to the species that can be viewed using
simple light microscope, like those used in schools.
These algae include species of the following taxa:
Dinoflagellates, Class Dinophyceae (armored
plant,
Greek). Many of these algae are large unicellular organisms
easily discernible even at low magnification of light
microscope. Dinoflagellates possess two flagella, laid
in the two perpendicular grooves of the theca (scanning electron microscopic
images of Black Sea dinoflagellates): one
flagellum girdle the cell around, another one is projected
forward. Dinoflagellate's flagella are twisted
and work as propellers. Due to the two flagella movement,
algal cell is turning around, and at the same time swimming
forth along the spiral trajectory - as if it's
screwing into the water. Flagella are very thin
and barely visible (mostly due to their rapid movement)
with the use of the light microscope.
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Ceratium furca; above -
a large Protoperidinium species
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Most species of Dinoflagellates possess outer carcass
called theca made of cellulose; it consists of
many plates located beneath the cell membrane. Their
shape of the thecal plates is used by researchers for
species identification. Some dinoflagellates lack
a hard theca; there are many small naked dinophycean
algae, most of them belong to the numerous genus
Gymnodinium. The largest dinophycean algae of
genus Noctiluca are also naked ones. Noctiluca
scintillans, reaching macroscopic size of
1mm, sometimes creates red tides in the coastal
Black Sea waters. When that happens, patches of sea
surface really acquire pinky-red color during the
daytime, and in the night those algae are the cause of
noticeable luminescence in the water.
Dinoflagellate cells may have most fantastical shape
- just look at Ceratium or Dinophysis
species. These are several dinoflagellate species common
in Black Sea summer phytoplankton; they are easy to
study with the use of most simple light microscope:
Prorocentrum micans, Ceratium furca
(furca means a fork in Latin - look at the shape
of the cell), Scrippsiella trochoidea, and
Goniaulax spinifera. The grooves hosting
the two flagella are well seen
in the sculptured theca of the latter.
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Prorocentrum micans
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Scrippsiella trochoidea
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Gonyaulax spinifera
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This microscopic pincushion is a cyst, thick-walled
dormant cell of dinophycean microalgae of genus
Gonyaulax. When cold autumn storms
come to Black Sea in November, many dinoflagellates
transform their cell shape, switch their physiology into
the hibernating mode, and fall to the bottom. Thick cell
wall of the cyst is necessary for the protection from
benthic predators, scavengers, and saprophytic bacteria.
Cell wall of Gonyaulax cyst is additionally
protected by the bristle of long spines. Dinophycean cysts are
even capable of going through the digestive system of
crustaceans or deritivourous fish like mullet and remain
intact; they can spend years buried in bottom sediments
and stay capable of germination. If
currents raise cysts from the bottom, and the water
temperature is high enough, germination ( excystment )
takes place: vegetative algal cell breaks off the
cracked cyst's shell. |

Gonyaulax cyst
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Alexandrium
excystment,
Utrish, Caucasian coast,
Feb. 2002 |
Massive excystment of dinoflagellate of genus
Alexandrium occurred in Utrish near Anapa in
February 2002. The sea storm disturbed bottom at the 30m
depth, and apparently raised cysts resting in the silt.
The weather was unusually warm for the winter and
Alexandrium cells concentration in seawater
reached 4000 cells per liter in several days. The
coat of the cyst seen at this micrograph is already very
thin, it is going to break, and vegetative algal cell is
coming out. This way dinoflagellate bloom can occur due
to the massive cyst germination, not to algal cells'
proliferation as it happen usually.
Alexandrium species are of special importance
for us for many of them can produce potent toxins of
saxitoxin group (like in scorpions, and
in tetraodontid fishes). Mussels feeding on
the phytoplankton containing Alexandrium cells
accumulate toxins, and consumption of such shellfish can
cause a possibly lethal disease called paralytic
shellfish poisoning. |
Dinoflagellates are unusual algae: many of them can
feed like animals, absorbing dissolved organic substances from
seawater, or even ingesting detritus particles and plankton
cells. Some of those dinophyceans retain photosynthetic
activity, they are called myxotrophs, that is the case
of the Ceratium species. There are dinoflagellates that
do not have chloroplasts at all, they are complete
heterotrophs; among such algae-animals are large
species of genera Dinophysis and
Protoperidinium. Huge dinoflagellates of genus
Noctiluca are even accounted as zooplankton. The
size of Noctiluca cells reaching 1.5 mm allows it
devouring even larvae of animals.
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Heterotrophic
Protoperidinium have a special part
of the antapical cell surface acting as gullet turning
out to swallow food, smaller cells and organic
particles. A predatory algae.
According to their
genetic relationships, these organisms are plants;
according to their ecological niche they are similar to
animals. Still during
marine ecological studies most heterotrophic
dinoflagellates are accounted as phytoplankton.
There are also heterotrophic dinoflagellates that are
parasites of other algae. |

Protoperidinium granii
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Dinoflagellates appear in plankton net samples from
Black Sea in early spring, and reach their highest
concentration in August and September; they almost
completely disappear from plankton to the end of autumn.
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Achnantes brevipes
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Diatoms, class
Bacillariophyceae. Cells of these algae
are coated in ornate silicon frustules, like boxes with
covers (diatom is consisting of two parts in Greek).
When diatom cell divides, each daughter cell retain one
half of the parent outer carcass. This is a side view
of the colony of a large diatom Achnantes,
it is possible to discern the two parts of the frustule
even with the average light microscope. Achnantes
is a benthic diatom living on underwater surfaces to the
depth where sunlight reaches; currents and waves
disturbing bottom sediments and macroalgae, often raise
it into plankton community. |
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Here are several other common benthic diatoms of the
Black Sea coastal waters; they also are usual temporary
participants of the plankton community near shore. The
shape and intracellular distribution of chloroplasts
visible through the thin frustule, as well as the shape
and ornamentation of the frustule itself, are the
identification characters used in the light microscopic
studies of phytoplankton. |
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Licmophora gracilis
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Grammatophora marina
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Pleurosigma elongatum
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Thalassionema costatum
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There are many pores - slots and holes - can be seen
in the large frustules of Achnantes and
Licmophora: they provide the exchange of
dissolved matter between the cell and its
environment. Because of the massive perforation, the
diatom silicon thecae can be
more correctly compared to lattice cages, then boxes.
Even better these well ordered apertures can be seen in
the scanning electron microscopic images of marine
diatom frustules:

Left: benthic diatom
Achnantes;
Right: large planktonic diatom
Coscinodiscus;
Middle: A portion of the
frustule of Pseudosolenia calcar-avis, the
largest planktonic diatom in the Black
Sea. |
|
This is another,
very nice bentho-planktonic diatom common in the Black Sea: Striatella.
Its radiant chloroplast, the antenna for
capturing the sunlight, itself
looks like sun. |

Striatella unipunctata
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The most common and abundant marine planktonic diatoms are
species of the genus Chaetoceros. They can be found in
any part of the World Ocean, and they almost always represent
a substantial (often - the dominant) portion of the
phytoplankton in the Black Sea. Chaetoceros cells bear
four long spines
called chaeta,
and usually are associated in chain-type colonies.
Chaeta form defense for Chaetoceros cells, they are
effective even against big animals. Mussels
in Black sea were observed to close their valves and stop
feeding when Chaetoceros concentration in seawater was
very high (tens of thousand cells or more), thus protecting
their internal organs from sharp chaeta.
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Chaetoceros compressus
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Chaetoceros laciniosus can be
distinguished by terminal pairs of chaetae reminding
forceps
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Chaetoceros curvisetus - most
common diatom in the Black Sea
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Pseudosolenia calcar-avis
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Pseudosolenia calcar-avis is the
largest planktonic diatom of the Black Sea, its
length may exceed 1 mm. Sharp curved ends of the
frustule prompted botanists to give this algae its
species name: bird's spur - calcar-avis.
Despite its very large size and the silicon frustule
weight, Pseudosolenia does not sink fast: due to
the high surface-to-volume ratio it sails the currents
in the water.
Planktonic diatoms having no flagella, and incapable
of active locomotion are left the only way of struggling
sinking: reducing the volume to surface ratio -
increasing the windage of
their cells. That is reached
by an elongated (like in Pseudosolenia or
Nitzschia) or flattened (Coscinodiscus)
cell shape, frustule projections (chaeta of
Chaetoceros), and colony formation. In most
Chaetoceros species the daughter cells emerging
after the mother cell division remain conjunct - this
way the colony chain grows. Chaetoceros socialis
cells excrete mucous mass that holds ball-shaped colony
of this species together. |
Chain colonies are most common in diatoms. Hemiaulus builds
ladder-like chains, each cell division adds another step to
this ladder:
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Hemiaulus hauckii colonies
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Colonies of Pseudonitzschia
pseudodelicatissima
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Pseudonitzschia also build colonies: needle-shaped
cells are connected at the ends forming long threads. This
small diatom is a very illustrative example of an
opportunistic species, it is capable of explosive
proliferation resulting in algal bloom when environmental
conditions are most unfavorable for the majority of the other
phytoplankton species. It can produce population outbursts in
the middle of the winter, or during the summer depression of
phytoplankton. Using minimal available resources, this tiny -
1-2 μm thick and 20 μm long - diatom grows and divides very
quickly.
The smaller is the cell, the larger is its surface as
related to its volume, and consequently higher the rate
of matter exchange
between the cell and its environment. It means a higher
rate of mineral nutrient consumption, and higher rate of
excretion, and this is a secret both of higher growth rate,
and faster proliferation of the smallest cells of phyto- and
bacterioplankton.
Because of that, the smallest phytoplankton cells make
main contribution to renewal of the living matter in the Sea.
They are cells less then 20 μm in size, and they are
further divided into the two size groups:
nanoplankton -
their cells' size ranges from 2 to 20 μm, and
picoplankton,
cell size < 2 μm.
Nano- and
picophytoplankton cells contain more chlorophyll per mass unit
then larger phytoplankton cells, which adds to their
photosynthetic activity and correspondingly growth rate. They
are barely visible using conventional light microscopes, and
only when they are alive. When dead - they are impossible to
tell from detritus particles. They are also impossible to
catch using a plankton net: they pass trough the most dense
10μm gauze. Because of those technical reasons the role of the
nanophytoplankton was underestimated for a long time;
researchers paid more attention to the larger
microplankton (cells >20μm) which includes
most diatoms and dinoflagellates. Nanophytoplankton in
the Black Sea includes, among others, coccolithophores and
silicoflagellates which are referred to below.
There are very few algae in the winter phytoplankton in
Black Sea coastal waters. As the spring comes with the growing
amount of sunlight and growing water temperature, the Sea
start to enliven.
First - the smallest algae of nanoplankton appear: tiny
naked flagellates, coccolithophores, and small diatoms (often
Pseudonitzschia pseudodelicatissima is starting the
annual diatom succession).
further - Chaetoceros species follow, and larger
diatoms;
later - turn of large dinoflagellates comes;
then - zooplankton consume almost all phytoplankton.
From mid-May to mid-July phytoplankton proliferation is
calming down, Chaetoceros species dominate
microphytoplankton. This is a time when a new, summer-autumn
phase of phytoplankton succession in the Black Sea starts.
Like in the spring, this second cycle of the Black Sea
planktonic life often begins with the proliferation of the
smallest diatoms (Pseudonitzschia) and/or
coccolithophores.
Most pronounced and prolonged phytoplankton peak takes
place in August-September. With the autumn cooling of water
phytoplankton concentration falls, and as November storms come
the annual cycle of the phytoplankton succession in the Black
Sea finishes. This is a way of phytoplankton succession common
to the Central and Eastern Black Sea; marine life in the
eutrophied Western Black Sea coastal waters is more dependent
on the Danube, Dnieper and other large rivers' desalinating
discharge, and its nutrient load.
It was noticed long ago that warm winters usually are
followed by a more pronounced spring phytoplankton rise in the
Black Sea. Cold winters, and even more - sudden cooling in
spring, inhibit phytoplankton succession, sometimes erasing
spring phytoplankton peak and decreasing by times the summer
phytoplankton concentration.
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Coccolithophores (bearing round
stones in Greek) are very
small (cell size 5-10 μm) - nanoplankton algae
possessing two flagella and covered by round limestone
plates called coccoliths.
They belong to the algal division Haptophyta (=
Prymnesiophyta). They are so small that pass through
the most dense mesh used in plankton nets; special
filters with pores of 1-3 μm diameter are needed to
concentrate them. Due to their small size coccolithophores
are barely seen through the light microscope,
still coccoliths can be seen - they make cell surface
looking curly.
Emiliana huxleyi is the most ubiquitous coccolithophore
in the World Ocean. That very small
algae is famous due to its ability
for explosive proliferation, and causing blooms over
vast areas of water; for example almost whole surface of
Indian ocean can be engulfed in
the Emiliana bloom. |

Coccolithophore
Syracosphaera sp.
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Coccolithophore bloom,
Black Sea,
13 June 2000 |
Summer blooms of Emiliana huxleyi and other
coccolithophores occur in Black Sea almost each summer,
particularly during the June to July period. This NASA
satellite image taken on 13 June 2000 shows white
coccolithophores bloom visualizing the pattern of surface circulation in the Black Sea
at that moment. When the bloom is over, coccoliths fall
to the bottom, forming layers of the limestone
deposits.
Geologists found out that first layers of coccoliths
on Black Sea bottom were deposited ca. 3500 years ago,
when salinity in the Sea reached 11‰ after the connection of the Black
Sea to Mediterranean basin was reestablished;
coccolithophore blooms in Black Sea started at that
time
( History of the Black Sea
Ecosystem ) |
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Meringia sp
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Silicoflagellates or Dictyochophyñeaens (class Dictyochophyñeae ). Usually they are less abundant
then diatoms or dinoflagellates. During the well expressed
spring phytoplankton peak in the Black Sea, there can
be many nice little cells with numerous sharp spicules
- it's Dictyocha: unicellular
algae with silicic carcass. Look at it using the scanning
electron microscope. Siliceous framework
of dictyochophyceans is intracellular, in contrast to
the diatom's outer skeleton. Silicoflagellates can swim,
they have 2 flagella. Here
is another silicoflagellate from the Black Sea: Meringia.
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Euglenas, (class Euglenophyceae ),
relatives of green algae have no hard outer casing, they
are protected only by layer of mucous excretion;
euglenas swim using flagella. They appear in the
Black Sea coastal waters only when the environmental
conditions are specially favorable for them:
decreased salinity and higher nutrients' concentration.
Their population grows fast, then they disappear as fast
as appeared - zooplankton grazes them. Those survived
cover themselves by hard capsule and sink to the bottom;
cysts will rest before the coincidence of the conditions
favorable for them occur again. Euglenas possess simple
photoreceptor eye, tiny red organelle within their
cells, directing the algae to the surface water. This
small green worm-like euglenophycean algae often
appearing at our coast is Eutreptia
lanowii. |

Eutreptia lanowii
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Four-flagellated cells of Prasinophycean
algae from the near shore bloom north of Tuapse,
Black Sea; SEM picture
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Prasinophytes ( class Prasinophyceae,
division Chlorophyta - green algae ) -
small picoplankton cells having 1 to 8 flagella, covered by
protective scales. Sometimes they cause algal blooms in
coastal waters of the Black Sea, after heavy rains when
water becomes brackish and eutrophied. Their role in the
general ecology of the Black Sea is poorly investigated
since they are almost impossible to identify and study
with the use of the light microscope.
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Another, much larger algae in our net sample: it's of
brown-greenish color, it obviously lacks hard theca, and
it's multicellular. This is a seedling of brown
macroalgae, one of those that form bushy macrophyte belt
at the rocky coasts of Black Sea; may be it's a
beginning of an underwater 'tree' of Cystoseira
barbata, the dominant brown algae in the Black Sea.
At the moment it consists of no more then dozen cells,
it lives within plankton, and currents and waves can
throw it on shore, it can be eaten by crustaceans.
Reaching the age and size of recruitment, it needs to
settle on the hard bottom nearshore; if it settles on
the sandy bottom, it dyes. If the settlement stage of
development is reached when the seedling is at the depth
over 30 meters, it dies, for there is no light
sufficient for brown algae growth at that depth in the
Black Sea. Of thousands of such seedlings only several
will settle, grow, and reach the age of
reproduction. |

Brown algae seedling
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Studying plankton samples from the Black Sea we find
something new all the time: species new to Black Sea
phytoplankton occur each year. This is the result of
both the continuing invasions of algal species from other
parts of Ocean (Evolution of the Black Sea
Ecosystem), and progress in
research techniques, development of new methods and
instruments - now we can see many details that were
impossible to discern some 50 years ago.
Subtle details of astonishing architecture of phytoplankton
cells can be discovered with use of the scanning electron
microscope. Cells of dinoflagellates and diatoms, having hard
outer skeleton are most beautiful, and they are better
preserved during the preparation procedure.
Black Sea phytoplankton - scanning
electron microscope images
The following pages are available
only in Russian:
Black Sea
Marine Life - sandy bottom habitats - fishes, crabs,
mollusks...
Black
Sea Marine Life - sandy bottom habitats - let's think about it
Black Sea
Marine Life - submarine rocks - near
the surfline
Black
Sea Marine Life - submarine rocks -
deeper
Black
Sea Marine Life - submarine rocks -
even deeper
Oil pollution, coastal deforestation, etc.
Aegean Sea - compare to Black Sea
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