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Confirming Proofs
Chapters
Are organized into four parts and written as short, readily absorbed
units to increase instructor flexibility in selecting topics.
CHAPTER
Marine Animals
Without
a Backbone
Part Two
The Organisms of the Sea
FPO
The crab Trapezia flavopunctata
opu
and its coral host, Pocillopora.
M
CHAPTER
The Microbial
World
ost species
es of multicellular organisms inhabiting our
planet aree animals
a
(kingdom Animalia). In contrast
to photosynthetic
syn
organisms like algae and plants, we
animals cannot manufacture
our own food and must therefore
nuf
obtain it from others.
s. The
T need to eat has resulted in the evolution
of myriad ways of obtaining
bta
and processing food, as well as equally
diverse ways of avoiding
din being eaten.
The colorful crab in the photo on this page is a good
example. It inhabits
ts rreef-building corals, relying on them for
food and shelter. Th
The crabs feed on mucus, which the coral
produces to keep its
ts ssurface free from debris. The coral is also
an animal, though it may
m not look like one. It gets some of its
food from zooxanthellae
hel
that live in its tissues. The coral also
eats small planktonic
ic organisms that it captures by using stinging structures in its te
tentacles. Though an absent-minded crab is
occasionally captured
ed by a fish or an octopus, the crabs are usually safe among the coral
co branches. The crabs repay the favor by
using their claws to drive away other animals that have a taste
for coral tissue.
Our survey of the many kinds of marine animals follows the
traditional classification into two major groups: the vertebrates,
which have a backbone (a row of bones called vertebrae), and the
invertebrates, animals without a backbone.
At least 97% of all species of animals are invertebrates. All
major groups of invertebrates have marine representatives, and
many are exclusively marine. Only a few groups have successfully
invaded land. Were it not for one of these groups, the insects, we
could boast without hesitation that most invertebrate species, and
therefore most animals, are marine.
CHAPTER
Marine Fishes
Zooxanthellae Dinoflagellates (single-celled algae) that live within animal tissues.
• Chapter 5, p. 00; Figure 14.1
Bluestriped snapper
per (L
((Lutjanus
Lutjanus
tj
kkasmira)
i ) on an Indian
I di Ocean
O
cor
coral
all reef.f
F
ishes were
re the first vertebrates, appearing more than
500 million
on years ago. The first fishes probably evolved from
an invertebrate
ebrate chordate not much different from the lancelets
or the tadpole larvae
arvae of sea squirts that still inhabit the oceans.
Fishes soon
n made their presence felt and have had a tremendous impact on the marine environment. They feed on nearly all
types of marinee organisms. Some of the organisms discussed in
Chapters 5 through
ough 7, from bacteria to crustaceans, use fishes as
their home. Many
any other animals eat them.
Fishes are the most economically important marine organisms, and marine
ne fishes are a vital source of protein for millions of
people. Some are
re ground up for fertilizer or chicken feed. Leather,
glue, vitamins, and other products are obtained from them. Many
marine fishes are
re chased by sportfishing enthusiasts. Others are
kept as pets and have brought the wonders of ocean life intoC H A P T E R
many homes.
Marine Reptiles,
Birds,
and Mammals
VERTEBRATES: AN INTRODUCTION
Vertebrates (subphylum Vertebrata) share four fundamental
characteristics of the phylum Chordata with protochordates,
the invertebrate chordates like lancelets and sea squirts. Vertebrates
differ from these other chordates in having a backbone,
also called the vertebral column or spine, which is a dorsal
row of hollow skeletal elements, usually bone, called vertebrae.
The vertebrae enclose and protect the nerve cord, also called
Male frigatebird (Fregata magnificens), Galápagos Islands.
V
ertebrates originated in the ocean and have thrived there
ever since. Roughly 350 million years ago, vertebrates
invaded the land as well, an event that changed life on
earth forever. Descended from bony fishes, land vertebrates had
to adapt to the harsher conditions ashore. They lost the structural
support that water provides and had to develop ways of crawling
or walking to get around. They evolved from fish-like vertebrates
with two pairs of limbs as an adaptation for “walking” on the bottom or on land between pools of water (see “When Fishes Stepped
on Land,” p. 00). Because of this, land-dwelling vertebrates—even
snakes—are called tetrapods, meaning “four-footed.”
Living on land also means having to breathe air. Tetrapods
evolved from fishes that had lungs, internal air sacs that allow
absorption of oxygen directly from air. Tetrapods had to evolve ways
to keep from drying out. The delicate egg is especially vulnerable,
Key Concept Summaries
Highlight the most important terms
and ideas presented in preceding
paragraphs.
In-Text Glossary
Briefly explains key terms and concepts from other chapters.
Chapter and page references point students to more detailed
information. The extensive glossary in the back of the book provides complete definitions and often refers to illustrations or
other key terms that help explain a concept.
swimmers simplify things by just opening their mouths to force
water into the gills.
Structure of the Gills Fish gills are supported by cartilaginous or bony structures, the gill arches (Fig. 8.17b). Each gill
arch bears two rows of slender, fleshy projections called gill filaments. Gill rakers project along the inner surface of the gill arch.
They prevent food particles from entering and injuring the gill
filaments or may be specialized for filtering the water in filterfeeding fishes.
The gill filaments have a rich supply of capillaries (Fig. 8.17c),
the oxygenated blood of which gives them a bright red color. Each
gill filament contains many rows of thin plates or disks called
lamellae, which contain capillaries. The lamellae greatly increase
the surface area through which gas exchange can take place. The
number of lamellae is greater in active swimmers, which need large
supplies of oxygen.
Gas Exchange Oxygen dissolved in the water diffuses into the
capillaries of the gill filaments to oxygenate the blood. Diffusion
will take place only if oxygen is more concentrated in the water
than in the blood. This is usually true because the blood coming to the gills has already traveled through the rest of the body
and is depleted of oxygen (Fig. 8.15). As oxygen diffuses from
the water to blood in the capillaries, the amount of oxygen in
and the first land tetrapods, the amphibians (class Amphibia), never
really solved this problem. Represented today by frogs, salamanders,
and their relatives, amphibians must keep themselves moist, and
most lay their eggs in water. None of them are strictly marine.
Other groups of tetrapods solved the problem of water
loss and truly adapted to life on land. Reptiles (class Reptilia;
Fig. 9.1) evolved from now-extinct amphibians and for a long
time were the dominant land vertebrates. The birds (class Aves)
and mammals (class Mammalia) both evolved from different
groups of now-extinct reptiles.
Having adapted to the land, various groups of reptiles,
birds, and mammals turned around and reinvaded the ocean. This
chapter deals with these marine tetrapods. Some, like sea turtles,
have not fully made the transition and still return to land to lay
their eggs. Others, like whales, spend their entire lives at sea.
By the time the blood has flowed most of the way through the
gill, picking up oxygen, it encounters water that is just entering
the gill chamber and is rich in oxygen. Thus, the oxygen content
of the water is always higher than that of the blood. This system
makes the gills very efficient at extracting oxygen. Without this
countercurrent system, blood returning to the body would have
less oxygen.
The blood disposes of its carbon dioxide using the same
mechanism. Blood flowing into the gills from the body has a high
concentration of carbon dioxide, a product of respiration. It easily
diffuses out into the water.
Gas exchange in the gills of fishes is highly efficient. The surface area
of gills is greatly increased by lamellae, and the flow of water through
them is in a direction opposite to that of blood.
Gas Exchange The absorption of oxygen to be used in respiration (the
breakdown of glucose to release energy) and the elimination of carbon
dioxide that results from the same process.
• Chapter 7, p. 000
Diffusion The movement of molecules from areas of high concentration
to areas of low concentration.
• Chapter 4, p. 00
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Confirming Proofs
BALEEN WHALES
TOOTHED WHALES
(k) Spinner Dolphin
Males - 2.4m (7.8 ft)
Females - 2.0 m (6.5 ft)
(b) Minke whale
Males and females
9 m (30 ft)
(j) Common Dolphin
Males - 2.6 m (8.5 ft)
Females - 2.3 m (7.5 ft)
(m) Vaquita
Males - 1.4m (4.5 ft)
Females-1.5 m (4.9 ft)
(l) Dall's porpoise
Males - 2.4 m (7.8 ft)
Females - 2.2 m (7.2 ft)
(n) Harbor porpoise
Males and
Females—2 m (6.5 ft)
(a) Northern right whale
Males and females
17–18 m (56–59 ft)
(c) Humpback whale
Males and females
11–16 m (36–53 ft)
(d) Bryde’s whale
Males and females
15 m (49 ft)
(o) Beluga whale
Males - 5.5 m (18 ft)
Females - 4.1 m (13.4 ft)
(p) Killer whale
Males - 9.8 m (32 ft)
Females - 8.5 m (28 ft)
(q) Narwhal
Males - 4.7 m (15.4 ft)
Females - 4.2 m (13.7 ft)
(f) Fin whale
Males and females
22–24m (73–79 ft)
(e) Bowhead whale
Males - 18m (59 ft)
Females - 20 m (66 ft)
(r) Pilot whale
Males - 6.1 m (20 ft)
Females - 5.5 m (18 ft)
(g) Blue whale
Males and females
23–27 m (76–89 ft)
(t) Cuvier’s beaked whale
Males and
Females—5.5 m (18 ft)
(h) Gray whale
Males and females
11–15 m (36–49 ft)
(s) Sperm whale
Males - 18 m (59 ft)
Females - 12 m (40 ft)
(i) Sei whale
Males and females
18 m (59 ft)
EVOLUTIONARY PERSPECTIVE
Symbiotic Bacteria—the Essential Guests
S
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Illustrations and photographs have been carefully designed and
selected to complement and reinforce the text. The eighth edition
contains many new illustrations and photographs.
Table 8.1
by some fishes, squids, octopuses, and other
animals of the deep (see “Bioluminescence,”
p. 000). The bacteria are usually sheltered in
special light-producing organs, or photophores.
These deep-sea animals, which live in darkness,
use light to communicate with other members
of their species, lure prey, blend with the light
that filters from the surface, and perform other
functions. Flashlight fishes (Anomalops), which
live in shallow tropical waters, lodge their symbiotic bacteria in an organ beneath each eye.
A shutter mechanism controls the emission of
light, so that the fish can “blink” at night. Groups
of fish blink in synchrony, a behavior that is
involved in attracting prey.
Chemosynthetic bacteria symbiotic with mussels, clams, and tubeworms that live around deepsea hydrothermal vents have a very particular
role: manufacturing organic matter from CO2
and the abundant hydrogen sulfide (H2S) from
the vents. The symbiotic bacteria live in a special
organ, the feeding body, of the giant hydrothermalvent tubeworm Riftia (see Fig. 16.29).
Marine symbiotic bacteria can also affect
human health. Pufferfishes store a toxin that is
deadly to any predators (including humans) that
eat them. The fish, or fugu, is a delicacy in Japan.
Licensed chefs must prepare the fish; otherwise,
the toxin (tetrodotoxin) will kill anyone feasting on fish that has been improperly prepared.
Tetrodotoxin is a deadly neurotoxin (that is, it
affects the nervous system). In fact, it is one of
the most powerful poisons known, and there is
no antidote. The deadly toxin is stored mostly
in the liver and gonads of the fish, so the internal
organs must be expertly removed. Mistakes do
happen and numerous deaths (including the suicides of disgraced cooks) take place every year
in Japan. Cooks may still be guilty, but not the
puffers: the toxin is not produced by puffers but
apparently by symbiotic bacteria. The fish are
actually resistant to tetrodotoxin as a result of
a genetic mutation that changes the site where
the toxin binds to nerve cells.
Tetrodotoxin and very similar toxins have
also been found in a variety of marine organisms, including flatworms, snails, crabs, sea
stars, and several species of fishes. It has
also been found in the blue-ringed octopuses,
notoriously toxic animals. Tetrodotoxin of an
unknown source has also been found in dead
sea urchins and is suspected as their cause
of death. We are not yet sure if in all these
animals the toxin is produced by symbiotic
bacteria or if it is accumulated from their food.
In arrow worms, however, symbiotic bacteria
in the mouth do produce tetrodotoxin that is
used to paralyze prey.
Being a neurotoxin, tetrodotoxin may
block pain signals in humans. A derivative being
tested as a pain reliever for cancer and other
diseases would have the advantage of not being
addictive, as are morphine and similar drugs.
The production of powerful toxins that are
used by other organisms is just one example of
the amazing abilities of bacteria—invisible to the
eye but powerful giants when it comes to their
role in the environment.
Most Important Characteristics of Marine Fishes
Distinguishing
Features
Skeleton
Feeding
Reproduction
Hagfishes
No paired fins; no scales;
exposed gill slits;
marine
Cartilaginous skull;
no vertebrae,
no jaws
Suction by round,
muscular mouth
with teeth
Oviparous
Predators of dead or dying
fishes and bottom invertebrates
Lampreys
Two dorsal fins only;
no scales;
exposed gill slits;
fresh-water or anadromous
Cartilaginous skull;
no vertebrae,
no jaws
Suction by round,
muscular mouth
with teeth
Oviparous
Suckers of fish blood or
predators of
bottom invertebrates
Rays, skates
Paired fins;
placoid scales;
pectoral fins greatly
expanded;
five ventral gill slits;
mostly marine
Cartilaginous
Grinding plates to
feed on bottom
animals or gill rakers
to filter plankton
Oviparous,
viviparous
Predators of bottom
animals or filter feeders
Sharks
Paired fins;
placoid scales;
exposed, 5–7 lateral
gill slits; mostly marine
Cartilaginous
Teeth in jaws to
capture prey or
gill rakers to filter
plankton
Oviparous,
ovoviviparous,
viviparous
Predators or
filter feeders
Ratfishes
Paired fins;
placoid scales;
one pair of gill slits covered
by flap of tissue;
deep water
Cartilaginous
Grinding plates to
feed on bottom
invertebrates
Oviparous
Predators of bottom
invertebrates
Coelocanth
Paired fins;
large scales;
gills covered by operculum;
deep water
Bony
Teeth in jaws
to capture prey
Ovoviviparous
Predators
Bony fishes
Paired fins;
cycloid or ctenoid scales
(absent in some);
gills covered by operculum;
marine and fresh water
Bony
Teeth in jaws
to capture prey
or graze or gill
rakers to filter
plankton
Oviparous,
ovoviviparous,
viviparous
Predators, grazers, and
filter feeders
Group
ome bacteria have evolved to live in close,
or symbiotic, associations with other
marine organisms. Some symbiotic bacteria are parasites that may cause a disease.
Others, on the other hand, benefit their hosts.
These symbiotic bacteria began their association by increasing the chances of survival of the
host and evolved to become essential—hosts
could not survive without them. In many cases
the symbiotic bacteria are even sheltered in
special tissues or organs that evolved in the
host, an example of coevolution, in which
two species evolve in response to each other.
All eukaryotic organisms, including humans,
shelter bacteria without which they could not live.
The chloroplasts and mitochondria of eukaryotic
cells evolved from symbiotic bacteria (see “From
Snack to Servant: How Complex Cells Arose,”
p. 00). These bacteria have become an integral
part of all complex cells.
There are many cases of symbiotic bacteria
among marine organisms. Symbiotic bacteria,
for example, are involved in digesting the wood
that is ingested by shipworms (Teredo), which
happen to be bivalve molluscs, not worms.
Like all wood-eating animals, shipworms lack
cellulase, the enzyme needed to digest cellulose, the main component of wood. Wood is
a surprisingly common habitat in many marine
environments, so everything from driftwood
to boat bottoms is exploited by shipworms,
thanks to their symbiotic bacteria.
Symbiotic bacteria are also responsible for
the light, or bioluminescence, that is produced
Significance in the
Marine Environment
A Japanese pufferfish, or fugu (Takifugu niphobles).
Boxed Essays present interesting supplemental information on
varied subjects such as deep-water coral communities, tsunamis,
and red tides. A number of boxes have been highlighted as
“Evolutionary Perspective” boxes to emphasize the central role of
an evolutionary perspective in biology.
New! Illustrated tables at the end of each chapter in Part Two
provide a consolidated review of major taxonomic groups and
their characteristics.
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Coral Reef Seaweeds
Rolling the Dice:
Climate Change
The earth is getting warmer. Global temperatures have steadily increased over the last 150
years (Fig. 1), rising by about 0.6°C (1.1°F) during the 20th century. As of 2006, the six warmest years on record occurred during the
previous decade, the three warmest were
since 1998, and 19 of the 20 warmest occurred
since 1980. We already know, however, that
climate goes through regular cycles, not just
over the tens or hundreds of thousands of
years it takes for ice ages to come and go, but
also over centuries. Viking and other histories
recount a “Medieval Warm Period” from about
AD 850–1250, overlapping a period of high
solar activity. That was followed by the “Little
Ice Age,” a time of bitterly cold weather in
⬚C
14.4
Global average temperature
⬚F
57.9
14.2
57.6
14.0
57.2
13.8
1998 temperature
0.5
56.5
1880
1960
1920
Year
2000
Results of field work in central Pacific
coral reefs, particularly in the subtropical
Northwestern Hawaiian Islands, showed that
fleshy macroalgae were dominant in many of
the healthy reefs of the region, often forming
expansive meadows spanning acres and serving
as critical habitats for numerous invertebrates
and juvenile fishes. Macroalgae occupied as
much as or even more surface area than live
reef corals in 46% of the sites that were studied. Some of the macroalgae even proved to be
species new to science. Coralline algae, once
thought to be slow growing, are now known
to grow fast, and probably playing previously
unknown roles. The algae have been found, for
instance, to produce substances that stimulate
the settlement of the larvae of corals and
other invertebrate animals. Ongoing research
by the NOAA team is also geared towards
understanding the effect of grazers, nutrients,
and other factors on the species diversity and
relative abundance of macroalgae over time,
and, most of all, seeking answers to one critical
question: Are these changes good environmental indicators of the reef health?
For more information, explore the links provided on
the Marine Biology Online Learning Center.
0.0
-0.5
56.8
13.6
Figure 1
1.0
Thermometer measurements (1902 -1999)
Reconstructed average annual temperature (1000-1980)
Reconstructed 40-year average (1000-1980)
Long-term trend (1000-1900)
the first to colonize any vacant surfaces on the
reef. Coralline algae, the second group, are red
algae that produce a hard calcareous skeleton
(see “Red Algae,” p. 000). Some reef biologists
believe that coralline algae literally hold coral
reefs together. The third group belongs to
macroalgae, typically fleshy seaweeds that are
larger in size and therefore more obvious to
the observer than those in the first two groups.
Halimeda (see Fig. 14.10) is a calcareous green
macroalga that together with the coralline algae
plays an important role in the deposition of
calcium carbonate in coral reefs.
Understanding the role of algae in healthy
coral reefs is what stimulated some of the
recent research conducted by NOAA in the
central Pacific. The distribution of algae on
reefs was studied using techniques such as
video-transects, photoquadrats, and toweddiver surveys. Hundreds of species of tropical
macroalgae are common components of coral
reef ecosystems, yet they remain among the
least-understood and least-studied of all coral
reef inhabitants. Their abundance can vary,
increasing with rising water nutrients, which
are typically very low in tropical regions.
Grazers, like sea urchins, molluscs, and fishes,
keep their numbers down.
S
Europe from roughly 1500 to 1850. It could be,
then, that the 20th-century rise in temperat ure
is just another natural fluctuation.
It is important to understand the difference between climate and weather. “Climate”
means average conditions over decades or
longer, whereas “weather” refers to shortterm conditions over days or months. Cold
weather in Miami—a frost, say—doesn’t mean
Miami has a cold climate. A snap of cold
weather for a few weeks, or even an unusually
cold winter, doesn’t tell us anything about
how climate might be changing, nor is one
heat spell proof of global warming. To judge
the 20th-century temperature rise we need to
know how climate has fluctuated in the past.
Reliable, widespread measurements
with thermometers go back only to about
1850, but there are natural temperature records of past climates, including ocean
sediments (see “The Record in the
Sediments,” p.••), corals, ice cores (Fig. 2),
tree rings, pollen, boreholes in continental
rock, and historical records such as the extent of Arctic sea ice or how far north grapes
Figure 2 Annual bands of ice, like these from Péru’s
could be grown. One such reconstruction,
Quelccaya ice cap, are one way scientists can tell what
past climates were like.
called the “hockey stick” graph (Fig. 3),
Mean global temperature relative to 1961-1990
Our planet never stops changing, in cycles
that range from the hundreds of millions of
years over which continents drift apart and
reform to the annual change of seasons and
even the daily cycle of night and day. These
cycles are earth’s natural rhythms, but humans
are changing our planet in ways and at rates
unprecedented not just in recorded history
but on geological timescales. Where once we
weren’t aware of our impacts on “spaceship
earth,” we now know that some of our activities are changing the entire planet. It could be
that everything will be all right—anything is
possible. By continuing these activities unabated, however, humanity has embarked on
a vast global experiment, gambling that the
fundamental processes that provide our most
basic needs—
air, food, water, and shelter—
will stand up to whatever we do.
eaweeds are not often regarded as important components of healthy coral reefs,
mostly because reefs impacted by pollution and other detrimental factors very often
become overrun by seaweeds (see Fig. 14.14).
Recent research by the Coral Reef Ecosystem
Division (CRED) of the United States’ Ocean
and Atmospheric Administration (NOAA) has
shown, however, that some highly diverse,
healthy coral reefs in pristine environments from
the relatively isolated central Pacific Ocean are
dominated by seaweeds rather than by corals
or other types of marine organisms. Such a surprising finding has challenged marine biologists
to revise their views on the role of seaweeds
in coral reefs, while still recognizing that reefs
are built by colonial reef animals, their symbiotic zooxanthellae, and coralline algae (see “The
Organisms That Build Reefs,” p. 000).
Hundreds of species of coral reef algae are
known, each with their own unique morphological and functional attributes. This vast algal
diversity is typically reduced by reef biologists
into three major functional groups. Turf algae
consist of a group of small, filamentous seaweeds that cover essentially all non-living hard
surfaces on the reef, including dead coral and
spaces between live coral colonies. They are
-1.0
1000
1100
1200
1300
1400
1500
1600
1700
1800
1900
2000
Year
Figure 3 Recent temperatures (red) and past temperatures (blue) estimated from tree rings, ice cores, corals, and
historical records, The grey shading indicates the range of uncertainty. Even with this uncertainty, 1998 was the
warmest year in a millennium.
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hnology in the field of marine biology. Sample topics include the
evolution of tetrapods, the use of ocean observations in forensic
science, common ground between ecology and economics, and
deep-sea exploration in the Cayman Trench.
New Feature! A major highlight, introduced in the seventh edition,
is an insert located after Chapter 10, Special Report: Our Changing
Planet. This current and informative insert highlights several key
aspects of global change, including global warming, ocean acidification,
eutrophication, hypoxic zones, and stratospheric ozone depletion. The
global depletion of fisheries, and loss of critical habitats.
Each chapter ends with an Interactive Exploration to be used in
conjunction with the Marine Biology Website. Students are encouraged to visit www.mhhe.com/castrohuber8e for access to chapter
quizzing, interactive chapter summaries, key term flashcards, marine
biology video clips, and web links to chapter-related material.
Part One
360
Structure and Function of Marine Ecosystems
Interactive
Exploration
reduce climate change by fertilizing the oceans, whether they will
work, and what the effects on the ocean might be.
Whynott, D., 2001. Something fishy about this robot. Smithsonian, vol.
31, no. 5, August, pp. 54–60. In the hope of designing more efficient
vessels, scientists and engineers try to copy the bluefin tuna.
Wray, G. A., 2001. A world apart. Natural History, vol. 110, no. 2,
March, pp. 52–63. The larvae of marine invertebrates have many 6/29/09
adaptations for life in the plankton.
The Marine Biology Online Learning Center is a great place to check your
understanding of chapter material. Visit www.mhhe.com/castrohuber8e
for access to interactive chapter summaries, chapter quizzing, and more!
Further enhance your knowledge with videoclips and weblinks to chapterrelated material.
Critical Thinking
cas24166_Ch06_102_114.indd 105
1. Plankton are unable to swim effectively and drift about at the mercy
of the currents. You might think that the currents would scatter
planktonic organisms throughout the oceans, but many species are
restricted to particular regions. What mechanisms might allow a species to maintain its characteristic distribution?
2. Spiny species of diatoms are found in both warm subtropical waters
and colder areas. Because warm water is less dense than cold water,
would you predict any differences between the spines of warmwater and cold-water individuals? Why?
Anderson, D. M., J. M. Burkholder, W. P. Cochlan, P. M. Glibert,
C. J. Gobler, C. A. Heil, R. M. Kudela, M. L. Parsons, J. E. Rensel,
D. W. Townsend, V. L. Trainer, and G. A. Vargo, 2008. Harmful algal
blooms and eutrophication: Examining linkages from selected coastal
regions of the United States. Harmful Algae, vol. 8, pp. 39–53.
Arrigo, K. R., 2005. Marine microorganisms and global nutrient cycles.
Nature, vol. 437, no. pp. 349–355.
Dybas, C. L., 2006. On a collision course: Ocean plankton and climate
change. Bioscience, vol. 56, no. 8, August, pp. 642–646.
Fonteneau, A., P. Pallares, J. Sibert, and Z. Suzuki, 2002. The effect
of tuna fisheries on tuna resources and offshore pelagic ecosystems.
Ocean Yearbook, vol. 16, pp. 142–170.
Heisler, J., P. M. Glibert, J. M. Burkholder, D. M. Anderson, W. Cochlan,
W. C. Dennison, Q. Dortch, C. J. Gobler, C. A. Heil, E. Humphires,
A. Lewitus, R. Magnien, H. G. Marshall, K. Sellner, D. A. Stockwell,
D. K. Stoecker, and M. Suddleson, 2008. Eutrophication and harmful
algal blooms: A scientific consensus. Harmful Algae, vol. 8, pp. 3–13.
Hofmann, E. E., P. H. Wiebe, D. P. Costa, and J. J. Torres, 2004. An overview of the Southern Ocean Global Ocean Ecosystems Dynamics program. Deep Sea Research Part II, vol. 51, nos. 17–19, pp. 1921–1924.
Johnsen, S., 2001. Hidden in plain sight: The ecology and physiology of
organismal transparency. Biological Reviews, vol. 201, pp. 301–318.
Katz, M. E., Z. V. Finkel, D. Grzebyk, A. H. Knoll, and P. G. Falkowski,
2004. Evolutionary trajectories and biogeochemical impacts of marine
eukaryotic phytoplankton. Annual Review of Ecology, Evolution, and
Systematics, vol. 35, pp. 523–556.
Pearre, S., Jr., 2003. Eat and run? The hunger/satiation hypothesis in
vertical migration: History, evidence and consequences. Biological
Reviews, vol. 78, pp. 1–79.
Schmidt, D. N., D. Lazarus, J. R. Young, and M. Kucera, 2006.
Biogeography and evolution of body size in marine plankton. EarthScience Reviews, vol. 78, nos. 3–4, pp. 239–266.
Wiebe, P. H. and M. C. Benfield, 2003. From the Hensen net toward
four-dimensional biological oceanography. Progress in Oceanography,
vol. 56, no. 1, January, pp. 7–136.
Wilhelm, S. W. and C. A. Suttle, 1999. Viruses and nutrient cycles in the
sea. BioScience, vol. 49, no. 10, October, pp. 781–788.
Worm, B., M. Sandow, A. Oschlies, H. K. Lotze, and R. A. Meyers,
2005. Global patterns of predator diversity in the open oceans.
Science, vol. 309, no. 5739, pp. 1365–1369.
For Further Reading
Some of the recommended reading may be available online. Look for live
links on the Marine Biology Online Learning Center.
General Interest
Critical thinking questions challenge students to think more
deeply about the chapter material and also help stimulate class
discussion.
Amato, I., 2004. Plankton planet. Discover, vol. 25. no. 8, August,
pp. 52–57. Descriptions and beautiful images of some of the
phytoplankton that support the food web.
Falkowski, P. G., 2002. The ocean’s invisible forest. Scientific American,
vol. 287, no. 2, August, pp. 54–61. Phytoplankton help control the
earth’s climate. Should we risk tinkering with them?
Glibert, P. M., D. M. Anderson, P. Gentien, E. Granéli, and K. G. Sellner,
2005. The global, complex phenomena of harmful algal blooms.
Oceanography, vol. 18, no. 2, June, pp. 136–147.
Holland, J. S., 2007. Small wonders, the secret life of marine microfauna.
National Geographic, vol. 212, no. 5, November, pp. 96–111. Very
nice photos of small zooplankton.
Johnsen, S., 2000. Transparent animals. Scientific American, vol. 282, no. 2,
February, pp. 80–89. Becoming invisible requires a bag of tricks.
Klimley, A. P., J. E. Richert and S. J. Jorgensen, 2005. The home of
blue water fish. American Scientist, vol. 93, no. 1, January/February,
pp. 42–49. A look at the habitat and spectacular migrations of epipelagic
fishes.
Leslie, M., 2001. Tales of the sea. New Scientist, vol. 169, issue 2275,
27 January, pp. 32–35. Biological detectives deduce which unseen
marine microbes live in the ocean, and how, from fragments of their
genetic material.
Lippsett, L., 2000. Beyond El Niño. Scientific American Presents, vol. 11,
no. 1, Spring, pp. 76–83. El Niño is just one of several regular oscillations in ocean circulation that influences our climate on land.
McClintock, J., 2002. The sea of life. Discover, vol. 23, no. 3, March, pp. 46–53.
A new look at the Sargasso Sea reveals secrets about all the oceans.
Should we fertilize the ocean to reduce greenhouse gases? 2008. Oceanus,
vol. 46, no. 1, January. This issue of Oceanus explains proposals to
For Further Reading lists “General Interest” articles in publications such as Scientific American, Discover, and National Geographic,
which are appropriate for students with limited backgrounds in
science and “In Depth” readings for students who want to study
particular topics in detail.
11:25:43 PM
In Depth
The World Ocean
ARCTI C
S O UTHE RN
G
T
PA
CI
FIC
BA S
IN
DG
E
RI
A
NA
ZC
P A C I F I C
7,939 m
324 m
5,089 m
KJA
NES
RIDGE
ve
r
5,664 m
RIO GRANDE
RISE
S O U T H
ATLANTIC
O C E A N
5,183 m
Equator
0°
Ascension
W
3,915 m
Gulf of
Guinea
5,130 m
6,434 m
R I DG E
Ri
BRAZIL
5,955 m
20°
Tropic of Capricorn
R
A
L
V
ID
G
E
NAMIBIA
SOUTH
AFRICA
IS
536 m
CAPE OF
GOOD HOPE
Tristan da
Cunha
2,274 m
40°
6,115 m
1,506 m
Falkland Islands
SOUTH INDIAN BASIN
O CE AN
Antarctic Circle
Strait of
Magellan
4,799 m
Auckland
Islands
PA
C
IF
IC
-A
NT
AR
CT
IC
R
G
ID
E
S O UTHE R N
O CE AN
LING
B EL
SHAU
S C OTI A South
Georgia
SEA
CAPE HORN
5,158 m
492 m
S OU T H SAN
LAIN
SEN P
WE
WEDDELL
SEA
120°
150°
180°
150°
120°
90°
DDE
I
DW
AT
LA
IC
NT
D
-IN
CH
LA
LL P
T
IAN RIDGE
60°
827 m
IN
Antarctic Circle
4,754 m
80°
A N T A R C T I C A
90°
N CH
TASMAN
PLATEAU
RE
R ID G E
60°
on
ID
- A
T L
A N T
IC
R I
S
E
ES
REY
Be
A
KE
HW
NEW
ZEALAND
Easter
Island
E A S T
H
C
TRE N C H
DEC
R MA
S O UTH
PAC I F I C
O CE AN
UT
80°
cas24166_FM.indd xiv
ID
T
NA
A
E
YA
P
EN
CH
TR
H
EAST INDIAM
A
RIDGE
NG A
R I D G E
Y E A S T
N I N E T
W
TH
U
O
ENDERBY PLAIN
SO
20°
A F R I C A
M
ri ng S tra it
H
JA
R E PAN
NC
H
N
CAS
P IA
EAU
S- LACCADIVE PLAT
C HAGO
E
DG
a
AR
Ch
Mozambi que
M A DAG A S C
RI
DI
IN
ES
T
IAN
KERGUELEN
PLATEAU
5,993 m
60°
TA SMAN
5,184 m
SEA
SOUTH AUSTRALIAN
BASIN
IND
10,711 m
A m az
S O U T H
A M E R I C A
PERÚ
H
N
E
C
S
MO
EN
U N TAI N
TR
LE
PERÚ -CHI
ST
4,135 m
4,763 m
Tropic of Cancer
Cape Verde
Islands
7,177 m
S
EA
O
C
ID
R Panamá
Canal
S
CO
Canary
Islands
E
R
ES T
5,220 m
9,988 m
5,581 m
TH
CARIBBEAN 5,335 m
SEA
TR
EN
CH
3,573 m
40°
2,281 m
2,041 m
M ED I T ER R A N EA N
SEA
3,124 m
SPAIN
Madeira
D
OU
7,565 m
AM
ER
ICA
Azores
E
N
D
New
Caledonia
Great
Australian
Bight
2,937 m
DL
E
5,399 m
French
Polynesia
Tahiti
60°
IC
E U R O P E
A
AN
a
Kerguélen
ni
S
S
NORTH 6,885 m
AMERICAN
We
RICO
BASIN
TO
st
CH
ER
I nd
i e s PU TREN
or
RI
40°
l if
R
A U S T R A L I A
PERTH
BASIN
N R ID G
E
4,402 m
6,863 m
xiv
HE B
OK E
M ID
5,630 m
Marquesas
Islands
Fiji
LT
4,619 m
PORTUGAL
G
Bermuda
SARGASSO
SEA
5,376 m
3,960 m
Kiribati
OU8,999 m
VI
T Y Tuvalu
GA
AZ
IN
H
T R E N CH
VI
LL E TR EN C
CORAL
SEA
R
IE
R
RE
EF
BA
234 m
EN NORTH
GL SEA
AN
D
3,359 m
153 m
NORT H
ATLANTIC
GRAND BANKS
OF NEWFOUNDLAND O C E A N
6,210 m
CAPE
HATTERAS
Gulf of
Mexico
MEXICO
Galapagos
Islands
B
BA
E
BR
Hawai’i
Islands
4,733 m
4,392 m
New
Guinea
7,333 m
NCH
N
INDIAN
OCEAN
6,199 m
1,727 m
N R
IDGE
Marshall
Islands
NE W
TR
E
Bay of
Fundy
CAPE COD
ip p i R i v e r
Sulawesi
Krakatoa Island
M
ss
A
IA
N O R T H
A M E R I C A
1,040 m
RI
G R EA T
V
AI
NORWEGIAN
BASIN
2,461 m
Newfoundland
UNITED
STATES
Midway
Island
W
a
C o l u mbi
R i v er
a
fC
fo
J
6,235 m
Cocos
Islands
Tropic of Capricorn
6,299 m
MA
CH
ra
A
6,139 m
10,861 m
NO RTH
PAC I F I C
O CE AN
HA
INDONESIA
at
l
ne
Maldive
Islands
DG
RI
AN
DI
IN
P LAT E AU
NE
E
AR
n
5,161 m
um
SC
D-
Comoro
Islands
20°
MI
MA
A foldout map at the end of text provides quick reference to the
World Ocean and the major coastal marine ecosystems and marine
protected areas of North America.
TaiwanPHILIPPINE
C
SEA
H
Philippine Mariana
Islands
Islands
10,322 m
S
5,035 m
Equator
KENYA
Gulf of
Thailand
RE N
AFRICA
SOUTH
CHINA
SEA
Bay of
Bengal
3,456 m
NORTHWEST
PACIFIC
BASIN
CANADA
5,174 m
ul
ARABIAN
SEA
IN E T
ILIPP
PH
n
R
Gulf of
Alaska
TR
I AN
AL E UT
8,501 m
EN
S EA
Ade
of
U
T
BERING
SK
A
AL A SUL
SEA
IN
s
CH
i a n I s l a n d P EN
EN
LA LABRADOR
BR
AD S E A
Strait of
OR
Belle Isle
G
B O N IN T R
R ED
Tropic of Cancer
G u lf
0°
I ZU TRE NCH
EAST
10,211 m
CHINA
SEA
7,389 m
Persia n
Gulf
20°
K
SEA OF
JA PAN
KOR EA
JAPAN
CHINA
Suez
Canal
I L
R
N O RW EG I A N
SEA
Arctic Circle
ICELAND
296 m
si
40°
10,367 m
ARAL
SEA
eut
7,973 m
T R E NC H
S
A S I A
EA
C
80°
3,632 m
Hudson
B ay
is
BLAC K
SEA
N
E M P E RO R S E A MO U N T S
Al
E
0°
GREENLAND
Baffin
Bay
A
SEA OF
OKHOTSK
RUS S IA
30°
2,102 m
BEAU FORT SEA
SE
MEIJI
SEAMOUNT
TO
R
60°
90°
O CE AN
C A N A DA
BASIN
3,740 m
AL AS KA
TR E N C
E
B
I
S
60°
120°
150°
180°
CHUKCHI
SEA
R
150°
EAST SIBERIAN
SEA
A
-ATL A NT
MI D
IC
120°
O CE AN
LAPTEV SEA
I
M O
U N
TA
I N
S
90°
Arctic Circle
A N
D E
S
60°
ARCTI C
80°
BAR ENTS
SEA
SEA
374 m
RA
KA
60°
30°
0°
8/6/09 12:22:31 AM