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Abhijit Mitra · Sufia Zaman
Basics of
Marine and
Estuarine
Ecology
Basics of Marine and Estuarine Ecology
Abhijit Mitra • Sufia Zaman
Basics of Marine
and Estuarine Ecology
Abhijit Mitra Sufia Zaman
Department of Marine Science Department of Oceanography
University of Calcutta Techno India University
Kolkata, West Bengal Kolkata, West Bengal
India India
The purpose of this book is to focus the marine and estuarine ecosystem,
which is one of the vital resource reservoirs of the planet Earth. These
resources are not only the abiotic resources like oil, natural gas or minerals,
but also encompass living resources like coastal vegetation, fishes, molluscs,
arthropods and representatives of almost all the animal phyla. We hope this
book will be a useful source of information for those working for a world that
is ecologically sustainable.
This book is not a final work but should be viewed as part of an ongoing
process, of a continuing effort to understand a set of complex interrelated
global issues. It is part of a continuous flow of ecosystem related research
papers and books. Indeed, some of the material in this book have been taken
from our own earlier papers, where research scholars like Dr. Ananda Gupta,
Dr. Amitava Aich, Dr. Kiran Lal Das, Dr. Aftab Alam, Dr. Harekrishna Jana,
Dr. Rajrupa Ghosh, Mr. Prosenjit Pramanick, Mr. Shankhadeep Chakraborty,
Mr. Atanu Roy, Ms. Bulti Nayak, Ms. Suhana Datta, Mr. Deepta
Chakravartty, Ms. Kasturi Sengupta, Mr. Kunal Mondal, Mr. Subhasmita
Sinha, Ms. Mahua Roychowdhury, Mr. Saumya Kanti Ray, Mr. Rahul Bose
and Mr. Saurav Sett contributed their findings through rigorous field work.
Our debt to Dr. Kakoli Banerjee, Subhdra Devi Gadi and Dr. Rajrupa
Ghosh is uncommonly large. Their contribution went far beyond that of
editors as they added texture and colour to our prose. At times it was difficult
to tell when editing ended and writing began.
Each of our colleagues, Dr. Subhra Bikash Bhattacharya and Mr. Tanmay
Ray Chaudhuri, put great effort in updating the book. We are thankful to
Dr. Pardis Fazli of the Department of Biological and Agricultural Engineer-
ing, University Putra, Selangor, Malaysia, for her effort in representing our
data in graphical forms.
We are also indebted to the Ministry of Environment and Forest, Ministry
of Earth Science, Department of Science and Technology (DST), Govern-
ment of India, and IUCN for funding our venture through various projects.
Several innovative programmes that constitute the annexure sections of the
present book may serve as a road map in climate change mitigation and
adaptation process. Few examples of such innovations are mangrove based
fruit products by Mr. Prosenjit Pramanick, carbon content in gastropods by
v
vi Acknowledgement
Ms. Bulti Nayak etc., which are valuable assets of the present knowledge
reservoir.
Finally, Dr. Abhijit Mitra expresses his gratefulness to his wife Shampa,
daughter Ankita and mother Manjulika whose inspirations and
encouragements acted as boosters to complete the manuscript. In many
rigorous field works in Sundarbans, the assistance provided by Ankita has
helped the authors to add case studies of various dimensions. The sacrifice
and inspiration that Dr. Mitra received from his father Late Dhanesh Chandra
Mitra can be considered as the foundation pillar of his effort in learning
ecology and grasping the subject.
Dr. Sufia Zaman expresses her deepest gratitude to her mother Mrs.
Ayesha Zaman for her unconditional love and practical day-to-day support,
and to her father Mr. Salim-uz-Zaman who gave her immense moral support.
Dr. Zaman also acknowledges the support of her beloved husband Dr. Sahid
Imam Mallick. Dr. Zaman wishes to accord her deep sense of gratitude to her
family members including her uncle (Mr. Pradip Kumar Mitra) and aunt
(Late Mrs. Kanika Mitra), younger sister (Ms. Sharmilee Zaman), her in-laws
and beloved grandmother (Mrs. Shibani Dhar) for their encouragement and
inspiration throughout the strenuous period of manuscript preparation.
Contents
vii
viii Contents
3.2 Currents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
3.2.1 Upwelling and Downwelling: Causes
and Significance . . . . . . . . . . . . . . . . . . . . . . . 68
3.2.2 Factors Regulating the Pattern and Distribution
of Currents . . . . . . . . . . . . . . . . . . . . . . . . . . 69
3.2.3 What Happens Practically? . . . . . . . . . . . . . . . 69
3.2.4 El Niño . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Brain Churners . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Annexure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
4 Abiotic Variables of the Marine and Estuarine
Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
4.1 Chemical Components . . . . . . . . . . . . . . . . . . . . . . . . 89
4.1.1 Spatio-temporal Variations of Salinity . . . . . . . 90
4.1.2 Deep-Water Salinity . . . . . . . . . . . . . . . . . . . . 90
4.1.3 Conservative and Non-conservative Ions . . . . . 90
4.1.4 Dissolved Gases in Seawater . . . . . . . . . . . . . 91
4.1.5 Nutrients . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
4.2 Sediment Compartment . . . . . . . . . . . . . . . . . . . . . . . . 96
4.2.1 Transportation of Marine Sediments . . . . . . . . 97
4.2.2 Classification of Marine Sediments . . . . . . . . . 98
4.2.3 Ooze: Definition and Type . . . . . . . . . . . . . . . 100
4.2.4 Sediment Deposition: A Boon or a Curse? . . . . 101
Brain Churners . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Annexure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
5 Producers of the Marine and Estuarine Ecosystems . . . . . . 137
5.1 Phytoplankton Community . . . . . . . . . . . . . . . . . . . . . 139
5.1.1 Diatoms . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
5.1.2 Dinoflagellates . . . . . . . . . . . . . . . . . . . . . . . . 143
5.1.3 Coccolithophores . . . . . . . . . . . . . . . . . . . . . . 143
5.1.4 Blue-Green Algae . . . . . . . . . . . . . . . . . . . . . 143
5.1.5 Green Algae . . . . . . . . . . . . . . . . . . . . . . . . . 143
5.1.6 Classification of Phytoplankton . . . . . . . . . . . . 143
5.2 Macrophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
5.2.1 Seaweed Community . . . . . . . . . . . . . . . . . . . 145
5.2.2 Mangroves . . . . . . . . . . . . . . . . . . . . . . . . . . 152
5.2.3 Seagrass and Salt Marsh Grass . . . . . . . . . . . . 156
Brain Churners . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
Annexure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
6 Consumers of the Marine and Estuarine Ecosystems . . . . . 193
6.1 Zooplankton Community . . . . . . . . . . . . . . . . . . . . . . 194
6.1.1 Zooplankton: Do They Maintain Uniformity
in Their Life Timeline? . . . . . . . . . . . . . . . . . 199
6.2 Vertebrate Community . . . . . . . . . . . . . . . . . . . . . . . . 203
Contents ix
Both ocean and seawater are saline in nature. brought by the rivers is too small and estimated
The salinity exhibits significant spatio-temporal to be two millionth part of the ocean. Some of it
variations. The salt in the marine ecosystem is absorbed by animals and plants to form
originates from various sources. It is stated that skeletons of lime or silica. A certain proportion
from the very beginning, the crust of the Earth is called cyclic salts is swept off the sea towards the
subject to slow dissolving action of water. Ever land by windblown spays and picked up by water
since seawater has been constantly washing the vapour and again brought into the sea. Thus, they
shores and rains falling on the land have been further reduce the gross contribution of salt
draining debris into rivers and seas. Thus, seawa- which the rivers make to the ocean. Moreover,
ter contains all the elements of the Earth’s crust. a great amount of calcium brought by rivers is
The rivers bring about 5.4 108 of the total constantly being withdrawn by marine animals
solids of the ocean. However, it is surprising that like coral polyps, foraminifera, gastropods,
there exists great similarity between river water oysters, etc. Inspite of the above facts, the small
and seawater in terms of chemical composition. amount of salt brought annually by rivers
The latter has chloride in abundance (about increases the salinity of the sea.
75 %), while the former has mostly calcium Salts are also added either due to erosion of
(60 %). There is only 2 % of sodium chloride in the earth crust or by some agencies which add
river waters. The conclusion is naturally that minerals from the obscure sources buried deep
either rivers do not convey enough salts to the within the earth, e.g. volcanic ash brings some
sea which makes it saline or whatever they pour material—especially chlorine, boron, sulphur
in the sea is continually lost. The quantity of salts and iodine.
1.1 Benthic Compartment 3
The marine ecosystem can be divided into devoid of vegetation. These slopes, under per-
benthic (substratum based) and pelagic (aquatic) missible angle of repose, also show development
zones. of eolian ripples and wind ripples. The formation
of dunes follows a basic sequence of dune forma-
tion stages.
1.1 Benthic Compartment During the initial stage of dune formation, the
perennial grass Paspalum vaginatum of the fam-
The benthic compartment of the ocean starts from ily Poaceae forms an arc-shaped train in the sand
the shoreline and extends up to the hadal zone with the convexity of the arc pointed along the
(Fig. 1.1). The zone is vast and complex and is direction of wind. This grass helps trap sand and
characterized by the presence of several interesting initiates the growth of mounds of sand. The
sculptures and features. For the convenience of the growth of the vegetation continues, which further
readers, the benthic zone of the marine ecosystem accelerates the accumulation of sand and which,
is discussed here on the basis of subdivisions like in turn, results in the growth of the dunes.
supralittoral zone, eulittoral zone, sublittoral zone, Hummocks of shrubs like Clerodendrum
continental shelf, continental slope, bathyal zone, inerme may coalesce sometimes to give rise to
abyssal plains and hadal zone. complexities in dune morphology. A typical
dune configuration is finally attained through
the upwind deposition of more sand, away from
1.1.1 Supralittoral Zone the convex front of the arc-shaped hummock
covered with vegetation. As a result, sand grains
The supralittoral zone also known as the splash are deposited on the upper part of the slip face
zone or spray zone is the area above which the and may eventually avalanche. Further drifting
highest tidal water reaches. The organisms of sands along the windward slope creates a steep
inhabiting this zone are adapted to minimum face to the dune structure.
availability of water and include species of Plants play an important role in dune forma-
gastropods and isopods. Some thorny plants tion. The root acts as sand binders and helps
(mangrove variety) like Acanthus sp. are com- stabilize the dunes while the leaves and stems
mon in the supralittoral zone. The thorns are the help trap airborne sand. The vegetation protects
features of xerophytes that reduce the loss of the dune from destructive winds and arrests dune
water from the plant body. migration to a considerable extent. In many des-
Sand dune is a common feature of tropical ert areas, vegetation cover has been found to be
supralittoral zone. A sand dune is formed by a the most effective and permanent mechanism to
mound of sand deposited by winds, which rises to arrest dune migration.
a summit and possesses a slip face. In the The plant communities associated with dunes
Sundarbans, the dunes run parallel to the southern may be separated into two distinct groups,
coastline and have no definite shapes during the namely, those which help in the establishment
initial stages of formation. The eolian sand deposits and growth of dunes and these which arrest dune
make up an area known as the dune field. Wind migration. Some plant species such as grass
sand ripples also characterize many areas of the Paspalum and the herbs Aeluropus lagopoides,
dune field. Sand dunes in the Sundarbans are gen- Cynodon dactylon, Opuntia sp., Salicornia
erally oriented parallel to the relevant wind and are sp. and Suaeda maritima are associated with
barchan or transverse dunes. Barchan dunes are embryonic dunes (pro-dunes) and new dunes.
crescent-shaped formed by unidirectional wind Ipomoea pes-caprae, Launea sp. and Sesuvium
and advance by avalanching of sand on the slop portulacastrum are generally associated with
face. In contrast, transverse dunes are elongated, post-embryonic to mature dunes and arrest dune
nearly straight and regularly spaced sand ridges. migration, to a great extent. The herb
The windward sides of dunes contain S. portulacastrum has extremely long, penetrating
superimposed eolian ripples, wherever they are roots which may reach 2–3 m in length. Ipomoea
a b
Ultraviolet
Pelagic
yellow
green
Neritic Oceanic
blue
red
l 0
ra al
to or
Photic
Lit
Light
zone
b litt
Su
Epipelagic 200 m 100
al
Mesopelagic
hy
1000 m
t
Ba
PELAGIC PROVINCE 200
Epipelagic 0 – 200 m Bathypelagic
2000 m
Dysphotic
Mesopelagic 200 – 1000 m
“Twilight”
zone
Bathypelagic 1000 – 2000 m
l
sa
Abyssalpelagic 2000 – 6000 m
ys
3000 m 300
Ab
Hadalpelagic > 6000 m
Be
Abyssalpelagic
nth
ic
l
da
Bathyal 200 – 2000 m Ha 5000 m
Abyssal 2000 – 6000 m
Hadal > 6000 m 500
6000 m
Aphotic
Hadalpelagic
zone
Dark
7000 m 600
Fig. 1.2 Ipomoea pes-caprae serves as the sand binder in the intertidal zone
1.1 Benthic Compartment 5
Fig. 1.3 View of Launea sp. that forms a dense cover on the dune surface and imparts stability
pes-caprae (Fig. 1.2) has good network of roots In some cases, the azimuthal directions of the
that often penetrate to depth of 1–2 m. The spe- cross-laminations in the same dune showed 180
cies, together with Launea sp (Fig. 1.3), can form reversals. The variability of inclinations of cross-
a dense cover on the dune surface. laminations reflected avalanche faces, variations
Investigations of the internal structure of the in wind directions and vegetation hummocks
dunes revealed that wedge, lenticular and planar occurring in different pats and levels of the
tabular type of cross-laminations in sections cut same dune. The internal laminations of the
both parallel and normal to the lee directions of dunes were often disrupted by the roots of vari-
dunes. The boundary surfaces of the cross- ous types of vegetation.
laminations had, in many cases, been eroded.
These surfaces were either straight or were con-
cave upward. The sets varied in thickness from a 1.1.2 Eulittoral Zone
few cm to 40 cm. The bounding surfaces showed
variable degrees of inclination (5 –40 ) in It is the zone between high tide and low tide and
sections parallel to the lee-slope of the dunes. sustains a wide spectrum of organisms. The zone is
Dune sections cut normal to the lee-slope also also referred to as midlittoral zone and is the dwell-
revealed bounding surfaces with variable ing ground of wide variety of organisms (Fig. 1.4).
inclinations. Sliding of the sands down the lip The eulittoral zone sustains a wide spectrum
face caused high dip of the bounding surfaces, of flora and fauna. In tropical regions, this zone is
while erosion by sourcing on the stoss side was the survival ground of mangroves, which are
responsible for creating upward concave or specialized halophytes with pneumatophores,
straight low dipping bounding surfaces. The supporting root system and possess features like
cross-laminations in between the upper and viviparous germination (Fig. 1.5).
lower bounding surfaces within a set showed The eulittoral zone is the matrix for ecological
dips ranging between 20 and 35 . Laminar succession, and the pH of the substratum
with dips of more than 30 were not uncommon. (Fig. 1.6) associated with the mangrove forest is
High moisture enabled the sand to remain cohe- slightly acidic in nature, with values ranging
sive at high angles of repose. between 5.5 and 6.8 (Fig. 1.7).
6 1 Marine Ecosystem: An Overview
HTL
LTL
Fig. 1.4 Intertidal mudflat in the estuarine ecosystem falls under the eulittoral zone; HTL high tide level, LTL low
tide level
1.1.4 Continental Shelf When the ice melted and sea level rose, these
areas were flooded and sediments built up in
Continental shelves are generally flat areas, aver- areas close to the new shore. Although presently
aging 68 km (40 miles) in width and 130 m submerged, these shelf areas bear the signatures
(430 ft) in depth, which slope gently towards of old riverbeds and glaciers, features they
the bottom of the ocean basin. The width of a acquired when exposed as part of the continent.
continental shelf is frequently related to the slope Some continental shelves are covered with thick
of the land it borders. Mountainous coasts, like deposits of silt, sand and mud derived from the
the West Coast of the United States, usually have land; for example, the Mississippi and Amazon
a narrow continental shelf, whereas low-lying Rivers deposit large amounts of sediments at
land, like the East Coast of the United States, their mouths. Other shelves are bare of
usually has a wide one. Continental shelves are sediments, such as where the Florida Current
the extensions of the continents to which they are sweeps the tip of Florida, carrying the shelf
attached (Fig. 1.8). The criterion for defining the sediments northwards to the deeper water of the
seaward edge of the continental shelf is a marked Atlantic Ocean.
change in slope. The continental shelf has a gra- The repeated emergence and submergence of
dient of about 1:1000. coast have been instrumental in shaping the mor-
The continental shelves are basically part of phology of the continental shelves. A study
the landmasses, and during geologic past, they conducted on this dynamicity in the Indian sub-
witnessed rise and fall of the sea level. When the continent (East Coast) is highlighted as
sea level was low during, the ice ages (periods of Annexure 1A.
increased ice on land), erosion deepened valleys, The continental shelves (Fig. 1.9) act as the
waves eroded previously submerged land and reservoir of several commercially important
rivers left their sediments far out on the shelf. items like sand, petroleum, natural gas and
dry land
60 M
continental shelf
idge
sl
op
r
arine
e e
is
lr abyssal
subm
continental crust a
nt plain
i ne
foot of slope nt
(max. change of gradient) co
sediment
1% sediment thickness ocean crust
Outer limit of CS
Fixed points (lat, long)
Fig. 1.9 Continental shelf: a zone for resource exploration and utilization
different categories of shellfish, finfish, rapidly from 200 m (600 ft) to 3000 m
seaweeds, etc. These areas are also treated as (10,000 ft) or it may drop as far as 8000 m
the bin of civilization, as several wastes of com- (26,000 ft) into a deep-sea trench, as it does
plex nature are dumped on the continental off the West Coast of South America. The con-
shelves. As the continental shelves are extremely tinental slope may show rocky outcroppings,
important in terms of commercial valuation and and it is often relatively bare of sediments
ecosystem services, therefore some regulations because of its steepness. Some continental
have been imposed on the issue of commercial slopes possess submarine canyons that are simi-
exploration and valuation. In the domain of laws lar to canyons found on land. Many of these
and policies, continental shelves as mentioned in submarine canyons are aligned with river
Article 76 encompasses prolongation of the land systems on land and were probably formed by
territory of all the maritime states that remains in these rivers during periods of low sea level. The
submerged state including the seabed and subsoil Hudson River canyon on the East Coast of the
and extends up to a distance of some 200 nautical United States is an example of this. Other sub-
miles. marine canyons have ripple marks on the floor,
and at the ends of the canyons sediments fan out,
suggesting that they were formed by moving
1.1.5 Continental Slope sediments and water. Oceanographers believe
that these canyons were formed by turbidity
Seawards from the continental shelf is the con- currents. Turbidity currents are swift avalanches
tinental slope, which extends to a depth of of sediment and water that erode a slope as they
1.6–3.2 km. The continental slope has a gradient sweep down and pick up speed. At the end of the
between 1:2 and 1:40. The angle and extent of slope, the current slows and the sediments fan
the slope may be of varying nature. In some out. Turbidity currents can be caused by
oceans, the angles may be low, whereas in earthquakes or the accumulation of large
some other cases, the angles may be high that amounts of sediments on steep slopes that over-
causes steepness. The depth may increase load the slope’s capacity to hold them.
10 1 Marine Ecosystem: An Overview
1.1.6 Bathyal Zone from the continental rise carried down by the
turbidity currents. Abyssal hills and seamounts
The bathyal zone lying between 200 m and are scattered across the seafloor. Submerged flat-
2000 m is characterized by absolute darkness topped seamounts, known as guyots, are found
and low temperature, making the survival of most often in the Pacific Ocean. They may vary
organisms highly difficult. In the absence of from low hills to mountains as high as 1525 m.
solar radiation, photosynthesis is practically nil The example of Bermuda rise is very prominent
in the bathyal zone except in some regions of in this context on which the Bermuda Islands are
tropics where a small amount of solar radiation formed. Seamounts are isolated peaks that rise
can penetrate up to some 600 m. The speed of several thousand metres above the seafloor.
water current is extremely slow in bathyal zone, These guyots are 1000–1700 m (3300–5600 ft)
and in some regions, water is almost stagnant. below the surface, with many at the 1300
The biodiversity is extremely poor in bathyal (4300 ft) depth. Many of these guyots exhibit
zone and encompasses few species of suspension the remains of shallow marine coral reefs and
feeders and mud scavengers. In sub-Arctic to the evidence of wave erosion at their summits.
equatorial regions, cold water bathyal corals are This indicates that at one time they were surface
widely distributed. features and that their flat tops are the result of
The sediments in the bathyal zone are terres- past coral reef growth, wave erosion or both.
trial, pelagic or authigenic (formed in place) in
nature. Terrestrial (or land-derived) sediments
are predominantly clays and silts and are com- 1.1.8 Hadal Zone
monly coloured blue because of accumulated
organic debris as well as bacterially produced The hadal zone represents the deepest part of the
ferrous iron sulphides. Coarser terrigenous ocean. This zone is found from a depth of around
sediments are also brought to the bathyal seafloor 6000 m (20,000 ft) to the bottom of the ocean.
by sporadic turbidity currents originating in The hadal zone has extremely poor biodiversity
shallower areas. Where supplies of terrigenous due to stressful environmental conditions in
materials are scarce, microscopic shells of phy- terms of temperature, pressure and light avail-
toplankton (coccolithophorids) and zooplankton ability. In the absence of any solar radiation, life
(foraminiferous and pteropods) fall through the in the hadal zone is triggered by the process of
water grain by grain, accumulating as white cal- chemosynthesis.
careous ooze deposits. Authigenic sediments Trenches are long narrow depressions in the
result from the interaction of clay, feldspar, and hadal zone that are over 6100 m deep. The
volcanic-glass particles with seawater, forming deepest known trench in the ocean compartment
the minerals glauconite, chlorite, phillipsite and is the Mariana Trench of the western North
palagonite. These sediments are characteristi- Pacific that is about 11,000 m deep. The
cally green because of their chlorite and glauco- Peru–Chile Trench extends for over 6120 km
nite contents. (3600 miles) along the coast of South America
and is the longest of the ocean trenches. The Java
Trench extends for a distance of almost 4760 km
(2956 miles) along the coast of the islands of
1.1.7 Abyssal Plains Indonesia. By comparison, there are only two
relatively short trenches in the Atlantic, namely,
The term abyssal plain indicates the flatness of the South Sandwich Trench and the Puerto
the surface. Abyssal plains are found at the base Rico–Cayman Trench. Trenches are invariably
of the continental rise and are relatively flat associated with the systems of active volcanoes
plains having a gradient less than 1:1000. They and are believed to be caused by down wrapping
are formed due to even deposition of sediments of the oceanic crust beneath the continental crust.
1.2 Aquatic Compartment 11
Continental Continental
shelf slope
Figure 1.10 highlights some major trenches of relatively calm recesses of the ocean, and
the world. changes are mainly attributed to sedimentation,
It is interesting to note from the above discus- uplifting and subsidence.
sion that the ocean basin has unique designs and The structures and features of ocean basin are
sculptures similar to the land features above it summarized in Fig. 1.11.
(Fig. 1.11). Mountain ranges, canyons, valleys
and vast plains are all the important components
of the underwater landscape. These physical 1.2 Aquatic Compartment
features of the ocean bottom are called
bathygraphic features, and unlike their counter- The entire water mass above the ocean floor
part topographic features on land, they change at (benthic substratum) with all the diverse types
relatively slow pace. Erosion is slow in the of organisms is referred to as the pelagic zone
12 1 Marine Ecosystem: An Overview
and may be differentiated into neritic and oce- and (2) aphotic zone. The photic zone is
anic zones. Neritic zone is the shallow-water restricted up to that layer where light is available
zone that extends from the intertidal zone to the and usually ranges up to 200 m from the surface.
edge of the continental shelf and includes the The productivity is more in this zone and the
coastal waters having an average depth of producers include many species of large algae
200 m. The water masses beyond the continental thriving in extensive beds (Fig. 1.13), as well as
shelf and overlying the deep abyssal plain con- huge populations of phytoplankton that serve as
stitute the oceanic zone. The depth of this zone is nutrient base for the rest of the consumers.
more than 6 Km. The pelagic zone is divided The aphotic zone is the deeper lightless zone
vertically into sub zones like epipelagic zone of the ocean usually below 200 m. Apart from the
(up to 200 m), mesopelagic zone (200–1000 m), above two zones, there is another zone known as
bathypelagic zone (1000–4000 m) and the dysphotic zone, sometimes referred to as the
abyssopelagic zone (4000–6000 m) (Fig. 1.12). twilight zone. This zone has very low levels of
illumination because the water above absorbs
more than 95 % of the sunlight. Photosynthesis
1.2.1 Classification on the Basis under these dim conditions is not possible.
of Light Penetration In the euphotic zone, there is plenty of light
to perform the process of photosynthesis by
On the basis of distribution/penetration of light, macrophytes and macrophytes due to which
the oceanic compartment may be divided into organic material is produced. This production
two distinct zones, namely, the (1) photic zone exceeds the loss of organic matter through the
1.2 Aquatic Compartment 13
process of plant respiration. The amount of light themselves against UV radiation (Ylognen
required for photosynthetic productions to just et al. 2005). Larvae of several benthic organisms
balance the respiratory losses in plants is and deep-dwelling shrimp species escape the
referred to as the ‘compensation light intensity’, adverse impact of UV radiation by changing
and the depth at which the photosynthetic pro- their dwelling place (Frank and Widder 1994;
duction is balanced by plant respiration is Adams 2001). Copepod (Martin et al. 2000;
known as ‘compensation depth’. Basically com- Rhode et al. 2001) and cladoceran (Johnsen and
pensation depth defines the lower boundary of Widder 2001) species avoid UV stress by vertical
the euphotic zone. migrations. Some crustaceans have evolved bio-
Light has immense role in regulating the chemical methods of avoiding the UV-induced
behaviour of marine organisms. Light has regu- stress, including pigmentation (Rhode
latory role on the behaviour of marine and estua- et al. 2001); more coloured shrimps tend to
rine organisms particularly on zooplankton occur in deeper water layers (Vestheim and
(Clarke 1934; Duval and Geen 1976; Pagano Kaartvedt 2009). Different responses of cladoc-
et al. 1993; Atkinson et al. 1996). Ultraviolet eran species to UV exposure in various freshwa-
(UV) radiation, especially with a wavelength of ter lakes have been documented (Leech and
280–315 nm, has adverse impact on many zoo- Williamson 2000; Leech et al. 2005). Light-
plankton species (Hunter et al. 1981; dependent behaviour of the abundant zooplank-
Kouwenberg et al. 1999). Some animals, for ton species inhabiting the White Sea (situated
example, larvae of Coregonidae fishes, use skin adjacent to North Polar circle) was studied exper-
pigmentation and avoidance behaviour to protect imentally by Daria et al. (2010).
14 1 Marine Ecosystem: An Overview
1.2.2 How to Estimate the Age constant through seasons and years and (iii) physi-
of Ocean Water? cal processes like wave action, turbulence, upwell-
ing, etc., have been ignored in this calculation.
The age of ocean water refers to the time or
period since the water mass was last at the sur-
face and in contact with the atmosphere. The 1.2.4 Decay Rate of 14C Method
computation of age of ocean water has great
relevance. It throws light on the rate of overturn This method was first reported by Kulp
of ocean water. This has important implications et al. (1952). According to this estimation, the
in terms of dumping of conservative pollutants atmosphere at the sea surface was only source of
like radioactive wastes and also replenishment of 14
C to ocean waters. Away from the surface, the
nutrients. If the average time of overturn is much 14
C content would not be replenished and would
less than the half-life of such materials, it would decay with its half-life of about 5600 years. The
be dangerous to dump them in the ocean because early measurements suggested an age of the
they would be brought to the surface while still order of 2000 years for water at 2000–5000 m
active and might be picked up by fish and so in the North Atlantic, but this was subsequently
conveyed back to man. shown to be too high on account of contamina-
There are several methods to estimate the age tion in the chemical processing.
of ocean water, which are discussed here.
90
1.2.3 Dissolved Oxygen 1.2.5 Sr Content Method
(DO) Consumption Method
Measurement of the 90Sr content of ocean water
This method was devised by Worthington in has revealed significant amounts at depths to
1954. He observed in the deep water of North 1000 m. As the only source of this isotope is
Atlantic Ocean (at depths of 2500 m) the average presumed to be the residue from atom bombs,
dissolved oxygen content decreased by 0.3 ppm starting in 1954, this indicates that the rate of
between 1930 and 1950. Considering the con- vertical mixing in the upper waters may be
sumption of oxygen to be constant, the rate of quite rapid.
decrease of DO is 0.3 ppm/20 ¼ 0.015 ppm/
year. Now this data can be used to calculate the
age of the ocean in the following way: 1.2.6 Hydrogen Isotope Method
Suppose, initial DO in the surface water of Bay
Radioactive tracer like tritium (3H) with a half-
of Bengal ¼ 7:6 ppm ðin 2014Þ:
life of about 12 years is also used to estimate the
Assume final DO ðat a certain depthÞ in Bay of
age of ocean water. It occurs in the upper layers
Bengal ¼ 5:8 ppm
of the ocean at concentrations of the order of only
Change in DO ¼ 1:8 ppm
one tritium atom for 1017 or 1018 ordinary hydro-
Considering the rate of decrease of
DO ¼ 0:015 ppm=year, the age of the water is gen atoms (1H), but with new techniques, it can
then 120 years: be measured quantitatively for age
determinations of seawater which have been rou-
This implies that the present parcel of water was tinely measured in the systematic redetermina-
last at the surface of Bay of Bengal during 1894. tion of the distribution of the main elements in
This calculation is based on several the world oceans in the GEOSECS (Geochemical
assumptions as, for example, (i) the consumption Ocean Sections Study). This was a multinational,
of oxygen occurs by chemical combination with multi-institutional study of the main oceans and
detritus, (ii) the consumption rate of oxygen is whose objective was the study of the
Annexure 1A: Oscillation of Seafloor in the Gulf of Mannar 15
geochemical properties of the oceans with West Coast. The shelf on the West Coast is broad
respect to large-scale circulation problems. with thin layer of sediment, while the shelf in the
East Coast is narrow with thick layer of sedi-
ment. The case studies of the area of Gulf of
Brain Churners
Mannar, extending from Tuticorin to
1. What causes the water salty in the
Rameswaram Island, are very pertinent in this
ocean?
context. The area lies between 8 470 and
2. Why the environment is stressful in
9 150 N latitudes to 78 50 and 79 300 E
supralittoral zone?
longitudes and sustains several types of habitats
3. How the salts of the ocean water are
like seagrasses, seaweeds and coral reefs
used by the organisms?
(Fig. 1A.1).
4. Why continental shelves are highly
productive compared to continental
slopes and abyssal plains?
2. Materials and Methods
5. Why the length of the food chain and
the number of trophic levels are more
The main components used for this study are
in the continental shelves and low in
Naval Hydrographic Chart (1975), Topographic
the hadal zone?
Sheet of Survey of India (SOI), Eco-Sounder and
6. Why photic zone of the ocean has
Global Positioning System (GPS). The bathyme-
greater fishery production compared
try map was prepared on the basis of Naval
to dysphotic or aphotic zone?
Hydrographic (1975) chart data and was subse-
7. Why seamounts are flat topped?
quently digitized into ARC-INFO to prepare a
8. How submarine canyons are formed
digital elevation model using ERDAS imagine
on the continental slopes?
software. A survey on the bathymetry of the
9. How trenches are formed at the bottom
area was carried out during April 1999 using
of the ocean?
eco-sounder and GPS along Mandapam and
10. Why the potential fishing zones mostly
Tuticorin coastal area in the Gulf of Mannar.
coincide with the zone of upwelling?
The depth values were recorded with respect to
chart datum. The measured depths were tide
corrected with respect to time and then converted
with respect to chart datum.
north-east and north directions. Whereas in the shore of Vilangu Shuli Island, whereas west of
west, southwest and northwestern parts of the the seafloor tends to slope very steeply. An
island, the seafloor slope is very steep. On the extensive well-developed fringing reef has been
north-eastern, eastern and southern sides of the identified around Vilangu Shuli and Karia Shuli
island, the seafloor is covered by fringing coral Islands extending to 3 m depth.
reef, extending down to 2 m depth with an In the shore between Vaippar and
approximate distance of 1.50 km from the coast Muttiapuram, the seafloor is at a depth of 3 m,
of the island. Between the depths of 3 m and 4 m, which extends to an approximate distance of
there is another island called Koswari Island, 3.51 km from the shore. In the area near the
which is located at 8 520 200 N–78 130 2200 E, mouth of Vembar, the seafloor topography is
with a distance of 6.09 km from Tharuvaikulam plain with a depth of 2 m. Between 2 m and
coast. In between 5 m and 10 m depths, the 20 m, the seafloor slopes very gently and extends
seafloor slopes moderately. From the depth of to a distance of 19.75 km. A sudden steep slope is
10 m to 20 m, the seafloor slopes gently, having encountered at 20 m depth; this area is the
a distance of 16.65 km between them. At a depth starting point of the continental slope and it
of 20 m, the seafloor falls suddenly with a very extends to the depth of 30 m. The width of the
steep slope (continental slope) extending till slope has been calculated as 5.45 km, and the
30 m depth. The width of this slope has been width of the continental shelf has been calculated
calculated as 7.27 km. The total width of conti- as 19.75 km. In the area from Terkku Narippeyur
nental shelf in Tuticorin region has been calcu- to Gundar River, the shelf is found at a depth of
lated as 26.75 km. In the coast from Veppalodai 3 m, extending to 0.36 km south and southeast of
to Sippikkulam, between 4 m and 6 m depths, Narippeyur coast and 0.88 km south of Gundar
elevated islands, namely, Kariya Shuli and River-mouth. Between the depths of 7 and 8 km,
Vilangu Shuli Islands, are situated at a distance the seafloor is considerably broad and the slope is
of 4.77 km and 6.56 km from Sippikulam coast very gentle. An elevated island is found exposed
respectively. The seafloor is found to be sloping above sea level, viz., Uppu Tanni Island situated
gently towards north, east and south from the at 6.72 km from Gundar River-mouth. The
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and tempers, the more Cicely compassionated the state of mind
which gave rise to them.
“It must be so terrible to feel that one has been false and
deceitful,” thought Cicely with a shudder, crediting, as was natural for
her to do, remorse with a far larger share in Geneviève’s
wretchedness than it really deserved. And she was marvellously
patient with the wayward girl; but yet in her very patience, in her
quiet kindness, there was a something against which Geneviève
instinctively rebelled.
“Why does she look at me so? I have done no wrong; it is not my
fault that Mr. Fawcett likes me best,” she would say to herself with a
species of childish defiance that was one of her characteristics when
roused to anger. “It was all that she was rich; but now that she is no
longer rich, how will it be now?” and a gleam of hope would shoot
across her for an instant, to be as quickly succeeded by misgiving
and despair. “He said, he promised, he would tell her he could no
longer marry her,” she repeated to herself a dozen times a day. “Why
has he not done so? Two, three days are past since her father’s
funeral, and he has not yet come; he has never come since the day
she would not see him. And Cicely does not seem surprised. What
can it be? Perhaps he has gone away!”
At last one morning, Geneviève in a fit of restless dreariness, set
off for a walk by herself. It was the same morning on which Mrs.
Methvyn and Cicely were talking together in the library, and it was on
her return from her walk that Geneviève, entering the room,
interrupted their conversation.
“So you have been out, my dear?” said Mrs. Methvyn kindly.
“Have you had a nice walk?”
“It is very cold,” replied the girl, shivering a little, and going nearer
to the fire.
She still had her hat and cloak on, and the light in the room was
not very bright. But now, something in her voice struck both Cicely
and her mother as unusual. It sounded faint and toneless.
“You have not caught cold, I hope?” said Mrs. Methvyn anxiously.
She was conscious that she had not given much attention to her
cousin’s daughter of late, and a touch of self-reproach made itself
felt.
“No, thank you; I have not caught cold,” said Geneviève. Then
she came a step or two nearer to where her aunt and cousin were
sitting, and they, looking at her, saw that she was very pale, and that
her eyes were red and swollen with crying.
“Aunt,” she said suddenly, and with a something of dignity in her
manner, new to her. “Aunt, you have been very good for me. I thank
you much, very much, for your kindness. I shall always thank you.
But I want you to let me go home now, home to Hivèritz, to my
mother. Please let me go; I can make the voyage by myself alone,
perfectly well. Please let me go. To-morrow, or in two or three days
at the latest.”
Mrs. Methvyn looked at her in astonishment.
“Geneviève, what is the matter?” she exclaimed. “What has
happened to put such an extraordinary idea into your head? Go
home alone! Nonsense, you know such a thing is impossible. You
must be reasonable, my dear, and tell me what has made you
unhappy. I can see you have been crying.”
“Nothing has happened,” replied Geneviève. “It is only quite
simply that I want to go home.”
“But you cannot go home all of a sudden in that way,” persisted
Mrs. Methvyn. “If there were no other reason against it, the
appearance of it at such a time would be an objection. You should
consider that, my dear. I have a great many troubles just now,
Geneviève. I think you should try not to add to them. And it is plain
that something has put you out this morning.”
Geneviève felt that Cicely’s eyes were fixed upon her with what
she imagined to be reproach, and she hardened her heart.
“Nothing has put me out,” she repeated. “I am not happy, that is
all. I do not love England; I want to go home.”
“But I cannot allow you to go home unless I am shown a good
reason for it,” said Mrs. Methvyn firmly. “When I brought you away
from your mother, Geneviève, it was with the wish and intention of
making you happy with us. If I have not succeeded, I regret it very
much; but still that does not free me from the responsibility I
undertook. I cannot possibly let you go home as you propose. You
do not really mean what you are saying—you are put out about
something, and afterwards you will be sorry.”
Mrs. Methvyn leant back wearily in her chair. Geneviève stood
before her, her eyes fixed on the ground.
“No,” she said, after a little pause, “no; I shall not be sorry
afterwards. I am sorry now,” she glanced up for a moment, “I am
sorry to trouble you. But I shall not be sorry for asking to go home. I
must go home. If I write and ask my mother, and if she consents, you
will let me go then?”
“I cannot prevent your writing home what you choose,” said Mrs.
Methvyn, as if tired of the discussion, “but, of course, it is very painful
to me that my plans for your welfare should end so, and I know it will
disappoint your mother.” She was silent for a moment, then she
suddenly looked at her niece with a new suspicion. “Geneviève,” she
said, speaking with an effort, “can it be that the reason you want to
leave us is, that you have heard any talk about our not being as rich
as we were?”
The blood rushed to Geneviève’s white face.
“No; oh, no!” she cried. “Indeed, it is not that. I am not so—so—
what do you call it?—so mean. No, it is not that.”
“But you might have some mistaken idea about it without being
mean,” replied Mrs. Methvyn, speaking more kindly. “You might have
some notion that it would be difficult now for me to do what before
was quite easy—that you would be an additional burden upon me.
But things are not as bad as all that, my dear. I shall be very glad to
have you with me, and I shall be quite able to manage comfortably. If
I saw you happy, I should be more pleased even than before to have
you with me, when—when I am quite alone—when Cicely has to
leave us.”
Her voice faltered a little as she glanced at her daughter, who all
this time had sat perfectly silent, neither by word nor look taking part
in the discussion. Once or twice during the conversation Cicely had
been tempted to interfere, but on reflection she refrained from doing
so. “It is better that mother should be prepared for something,” she
thought, “even this ill-timed request of Genevieve’s may pave the
way for what I must tell her.”
Geneviève’s eyes followed her aunt’s, but again something in
Cicely’s expression roused her latent obstinacy and defiance.
“I am sorry,” she said slowly. “I am sorry, but it must be. I cannot
stay here. Give me leave then, my aunt, to write to my mother about
my return home.”
“I told you before, you must write what you choose,” said Mrs.
Methvyn coldly.
And Geneviève left the room without saying more.
“Do you understand her, Cicely?” said Mrs. Methvyn when she
was again alone with her daughter. “Do you in the least understand
what has put this into her head? She is evidently very unhappy.
Surely,” she went on as a new idea struck her, “surely it cannot have
anything to do with Mr. Guildford?”
“No,” replied Cicely, almost, in spite of herself, amused at her
mother’s recurrence to her favourite scheme; “no. I am perfectly
certain it has nothing whatever to do with him.”
“Then, what can it have to do with?”
“She is certainly not happy,” answered Cicely, evasively. “I am
sorry for her.”
“Do you think you could find out more, if you saw her alone?” said
Mrs. Methvyn uneasily.
“I will go up and speak to her if you like,” said Cicely.
She rose from her chair as she spoke. As she passed her mother,
she stooped and kissed Mrs. Methvyn’s soft pale face—the lines had
grown much deeper and more numerous on it of late—the
roundness and comeliness were fast disappearing.
“Don’t worry yourself about Geneviève, dear mother,” she said.
“Even if she leaves you, you have me, haven’t you?”
“Yes, dear,” answered her mother. “I should not want her if I could
always have you! But, of course, it is not a question of wanting her. It
is so vexing to think of poor Caroline’s disappointment; it is so utterly
unexpected. I do not understand the child at all; she is not the least
like her mother.”
Cicely made her way up to her cousin’s room. Geneviève was
already seated at her little writing-table—pens, paper, and ink,
spread out before her.
“Geneviève,” said Cicely. “You have made my mother very
uneasy. She is most sorry on your mother’s account. The letter you
are going to write will distress Madame Casalis very much. I want
you not to send it—at least not to-day.”
“But I will send it,” said Geneviève angrily. “Why should you
prevent it? It is best for me to go, I tell you,” her voice softened a
little. “You don’t know—” she went on, “and if you did, you, so cold,
so réglée, how could you understand?”
Cicely looked at her with a strange mixture of pity and contempt.
“No,” she said, “perhaps I could not. But still Geneviève, for my
mother’s sake—I am determined to spare her all the annoyance I
can—I ask you not to write that hasty letter about going home, to
your mother to-day.”
“Why should I not?” said Geneviève.
“Because I tell you it is better not,” replied Cicely. “And you know I
always have spoken the truth to you, Geneviève.”
Geneviève looked cowed and frightened.
“Very well,” she said, “I will not write it. Not to-day.”
Cicely saw that she had gained her point. She left the room
without saying any more. And no letter was written by Geneviève
that afternoon. She sat in her room crying till it grew dark, and by
dinner-time had succeeded in making herself as miserable looking a
little object as could well be imagined, so that poor Mrs. Methvyn
said in her heart, that if it were not for the disappointment to
Caroline, her daughter’s absence would hardly be a matter of regret.
Cicely had no time to spare for crying; and tears, she was
beginning to find, are, for the less “med’cinable griefs,” a balm by no
means so easy of attainment as for slighter wounds.
“I think my tears are all frozen,” she said to herself with a sigh, as
she folded and sealed the last of her letters. She sat for a moment or
two gazing at the address before she closed the envelope, as if the
familiar words had a sort of fascination for her.
“I wonder if it is the last time I shall ever write to him,” she said to
herself. “When—when he is Geneviève’s husband, there can surely
never be any necessity for our coming in contact with each other. Yet
people grow accustomed to such things I have heard, and my
suffering cannot be unprecedented. Ah, what a sad thing life
becomes when one’s trust is broken! Far, far sadder than death!”
And after all, two or three large tears rolled slowly down her cheeks
and dropped upon the white paper.
This was the letter.
“Greystone,
“October 25th.
“My dear Trevor,—I should like to see you alone to-morrow. Will
you call here between two and three in the afternoon? I have
deferred asking you to come till now, because I thought it best that
you should thoroughly understand that I, in what I have determined
to do, am not acting hastily or impulsively.
“Your affectionate cousin,
“CICELY MAUD METHVYN.”
“It will prepare him to some extent,” she said to herself. The note,
simple as it was, had a certain formality about it, very different from
the girlishly off-hand letters she had been accustomed to send him.
“Will he feel it all relief?” she said to herself, as she thought how best
and most clearly she must put into words the resolution she had
come to. “Or will it be pain too? However he loves her, he did love
me, and he cannot have changed so entirely as to give no thought to
me.”
And again some tears blistered the smooth surface of the black-
bordered envelope in her hand.
CHAPTER V.
“HOW LITTLE YOU UNDERSTAND.”