Introduction To Marine Biology

An Independent Learning Course
3 credit hours
Developed and Graded by
Susan DeLisa
smdelisa@alaska.edu
University of Alaska Fairbanks
Copyright 2012 University of Alaska Fairbanks (revised 2.28.13)
The University of Alaska Fairbanks is an affirmative action/equal opportunity employer and educational institution.
BIOL F150
Introduction to Marine Biology
Course Guide
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Table of
Contents
Preface Welcome ............................................................................................................................................ v
Introduction Course Outline and Requirements..................................................................................................... v
Part One Setting The Stage: Principles of Marine Science
Lesson 1 The Scope, History and Practice of Marine Biology .........................................................................3
Lesson 2 The Benthic Environment .................................................................................................................. 7
Lesson 3 The Pelagic Environment ................................................................................................................ 19
Lesson 4 Biological Principles, Cells, Organisms & Ecosystems .................................................................... 37
Part Two The Players: A Survey of Marine Organisms
Lesson 5 Microorganisms and the Tree of Life: A Huge Cast of Tiny Characters ......................................... 53
Lesson 6 The Producers: Marine Autotrophs and Primary Production ........................................................ 67
Request for Exam #1 ...................................................................................................75
Lesson 7 Marine InvertebratesThe Lower Phyla ...................................................................................... 79
Lesson 8 Marine InvertebratesThe Higher Phyla ..................................................................................... 87
Lesson 9 Marine Fishes ................................................................................................................................. 97
Lesson 10 Marine Reptiles and Birds ........................................................................................................... 105
Lesson 11 Marine Mammals .......................................................................................................................... 115
Request for Exam #2 .................................................................................................123
Part Three Putting It All Together: Marine Ecosystems
Lesson 12 Life At The Top: The Epipelagic Realm ......................................................................................... 129
Lesson 13 The Netherworld: The Mesopelagic and Deep-Sea Realms ......................................................... 137
Lesson 14 On the Shelf: Continental Shelf Communities ............................................................................... 141
Lesson 15 Between Two Worlds: The Intertidal Zone and Estuaries ............................................................. 145
Lesson 16 The Human Factor ........................................................................................................................ 151
Request for Final Exam ..............................................................................................157
Appendix ................................................................................................................................................ 161
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WELCOME
Welcome to Biol 150, an Introduction to Marine Biology, the scientific study of life in the sea. This simple definition can be mislead-
ing, since marine biology is not really a separate science, but rather encompasses all aspects of biology as it applies to marine
organisms and their environments. Marine environments cover nearly three quarters of the planet and are extremely varied,
ranging from mangrove swamps and tidal pools, to volcanic vents and trenches lying miles below the surface of the sea. The
variety of life inhabiting these environments is equally diverse. We depend on the ocean and its life for food and raw materials,
transportation and recreation, and even for the oxygen we breathe and the regulation of Earth's climate. Yet, only a tiny fraction
of this huge realm has been explored. Amazingly, we have better maps of the moon and other planets in our solar system, than
we have of the ocean floor! Marine biology is an exciting field, with new discoveries being made every day. At the same time,
there is growing concern about the health of the oceans, and how that affects the health of the planet. The future of the oceans,
and indeed our own future, depend on how well we study this complex system, and how wisely we use the knowledge we gain.
Course Organization and Requirements
This course is divided into three parts, each building on the last, synthesizing information into a more complex and complete pic-
ture. Part One includes an overview of the scope and history of marine biology and the scientific method, a review of basic marine
geology, physics and chemistry, and basic biological and ecological principles necessary for the study of marine organisms and
their environments. In Part Two, we are introduced to the marine organisms, from the smallest bacteria, to the largest whales. In
Part Three, we visit these organisms in their homes, the major marine habitats where organisms interact with each other and their
surroundings to create unique ecosystems. Finally, we step back for a critical look at our own interactions with the marine world.
The course includes 16 lessons, each with a reading assignment and a written assignment. You will send your an-
swers to the written assignments to UAF eLearning and Distance Education, according to the instructions found at
their website or in your "Survival Handbook." Usually, your work will be graded and returned to UAF within one week. There
are also 3 exams, one at the end of each of the 3 parts of the course, and instructions for taking the exams are inserted
at the appropriate places in this Course Guide. Exams are open-book format, with a time limit of 4 hours. The only
materials that you may have to consult during exams will be this Course Guide and your textbook. Although the exams are
open-book, remember that it is important to be well-prepared, in order to comfortably complete the exam in the allotted time.
Because of the vast scope, complexity and interdisciplinary nature of marine biology, even at an introductory level, this
course covers a huge amount of information. Reading assignments are extensive and there may be many new concepts,
and much new vocabulary to master. So, it is critical that you understand the course requirements and plan a schedule
that will be manageable for you, and at the same time will allow you to complete the course on time. To help with this,
the written assignment for Lesson 1 includes a "Planning Schedule" to be filled out in duplicate, one copy to be sent
in with your completed assignment, and the other to be kept by you, so you can record and monitor your progress.
It also helps to have a systematic approach to studying. To get oriented, look at the list of Objectives at the beginning of each
lesson, and at the questions in the written assignment at the end of each lesson. Then, read the material in the Course Guide
for that lesson, followed by the reading assignment in the textbook. As you read, keep the objectives and questions for that
lesson in the back of your mind. Study the material until you have a good understanding. Then answer the questions. To show
your understanding of the material, you must answer clearly and completely, and in your own words, not those of the textbook,
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Course Guide or other references. If you need help, you may contact the instructor, Susan DeLisa, at smdelisa@alaska.edu
or at 907-474-4011 between 8:00am and 8:00pm Alaska Time. Submit all work to UAF eLearning and Distance Education.
Grading System
Your grade is calculated as the percentage of points earned, out of a maximum of 1000 points. Written assignments account
for 55% of your grade, exams for 45% (each exam is worth up to 150 points). Final grades will be assigned according to the
following scale:

98-100% = A+
9297% = A
9091% = A
8889% = B+
8287% = B
8081% = B
7879% = C+
7277% = C
7071% = C
6869% = D+
6267% = D
5961% = D
below 59% = F
Extra Credit
If a student's grade is borderline passing, then it may be possible to earn extra credit by writing a paper, upon approval by the
instructor. This is NOT a required (or usual) assignment for this class.
Extensions and Incompletes
All students must complete all coursework within one year from enrollment and will receive a grade based on work completed.
Semester-based students may apply for an incomplete IF they meet the requirements described in the "Survival Handbook"
(generally, must have at least half the work done and a passing grade). No Incomplete grades or additional extensions will be
given after one year from enrollment.
Required Textbooks
The texts required for this course are this Course Guide and Marine Biology 9th Ed., by Peter Castro and Michael Huber (ISBN
978-0-07-352420-7). The publishers (McGraw Hill) also offer a Marine Biology Website at www.mhhe.com/castrohuber9e
with chapter summaries and quizzes, other learning aids such as flashcards, and video clips and links to related sites. I strongly
recommend that you use this site, if possible, especially the videoclips, which can make the organisms and environments de-
scribed in this course "come alive."
About the instructor
Susan DeLisa received a B.S. in Biology from Antioch College, and M.S. and PhD. degrees in Biology from UAF.
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Other Important Information
Off-Campus Library Services through the UAF Rasmuson Library can be arranged by calling 1-800-478-5348. This unit
was set up to serve UAF students who do not have access to appropriate information resources in their town or village.
UAF Writing Center and Computer Lab offers free writing tutoring to any student in any subject via telephone and fax or
over the Internet. Students can call 907-474-5314 for information on how to fax a paper and have it tutored over the
telephone, or engage in an interactive Internet session. Both services are free.
Planning Schedule
Complete the Planning Schedule on the following pages and submit one copy (with your name and contact information) with
your first lesson. Put the appropriate deadline for course completion at the bottom of the schedule. If you are a graduating
senior, check with your counselor to find out the date all grades are due. If you are graduating or have any other personal
deadlines, it is imperative that you write to your instructor on your first lesson and inform him/her of your needs. Although the
staff and faculty will do everything possible to help you, a lack of planning on your part does not constitute an emergency for
the staff and faculty! Please plan ahead!
Opposite final exam, enter a date at least two weeks before your planned date of course completion. If you are a graduating
senior, plan to take the final exam at least one month before graduation. When entering planned dates for submitting each
written assignment, be realistic and try to pace yourself. Unless otherwise arranged with your instructor, up to two lessons per
week should be submitted.
YEAR-LONG: Remember, we assume you will take up to a year to finish this course. At a bare minimum, it should take three
months to complete a course. Instructor permission (in writing) must be obtained to finish in an accelerated time frame.
SEMESTER-BASED: If you are a semester-based student, follow the semester time requirements provided with your book
purchase. Remember, lessons cannot be accepted if turned in all at once.
Goals of this Course
The goals of this course are that:
1) you will experience the fascination of marine life and marine environments and appreciate their amazing diversity and
complex interrelationships
2) you will grasp the basic principles underlying this diversity and complexity (and be able to apply these principles for a
better understanding of your world beyond this course)
3) you will recognize how human activities influence marine life and marine environments, how the sea influences our world,
and the great need for a better understanding, and better management, of these relationships
Let's begin to explore marine biology!
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Planning Schedule
(Keep this copy.)
START: I began this course on _________________________ .
Date to Actual Feedback
Submit Date Sent Received Grade
Lesson 1 _________ _________ _________ _________
Lesson 2 _________ _________ _________ _________
Lesson 3 _________ _________ _________ _________
Lesson 4 _________ _________ _________ _________
Lesson 5 _________ _________ _________ _________
Lesson 6 _________ _________ _________ _________
Exam 1 _________ _________ _________ _________
Lesson 7 _________ _________ _________ _________
Lesson 8 _________ _________ _________ _________
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Lesson 10 _________ _________ _________ _________
Lesson 11 _________ _________ _________ _________
Exam 2 _________ _________ _________ _________
Lesson 12 _________ _________ _________ _________
Lesson 13 _________ _________ _________ _________
Lesson 14 _________ _________ _________ _________
Lesson 15 _________ _________ _________ _________
Lesson 16 _________ _________ _________ _________
Final Exam _________ _________ _________ _________
DEADLINE: Grade must be received by _________________ .
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Planning Schedule
(Submit this copy with your first lesson.)
STUDENT NAME: _________________________________ .
E-MAIL ADDRSS: _________________________________ .
CONTACT PHONE: _________________________________ .
START: I began this course on _________________________ .
Date to
Submit
Lesson 1 _________
Lesson 2 _________
Lesson 3 _________
Lesson 4 _________
Lesson 5 _________
Lesson 6 _________
Exam 1 _________
Lesson 7 _________
Lesson 8 _________
Lesson 9 _________
Lesson 10 _________
Lesson 11 _________
Exam 2 _________
Lesson 12 _________
Lesson 13 _________
Lesson 14 _________
Lesson 15 _________
Lesson 16 _________
Final Exam _________
DEADLINE: Grade must be received by _________________ .
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Part One
Setting The Stage:
Principles of Marine
Science
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Scope, History
and Practice of
Marine Biology
Marine Biology may be simply defined as the scientific study of living things in saltwater
environments. And here simplicity ends, for in this brief definition we have just introduced
three vast, complex and open-ended subjects.
Objectives of this Lesson
1. Plan a realistic schedule for completing the work required in this course.
2. Gain an historical perspective on human exploration and use of the sea and its
resources.
3. Learn about technologies for studying the marine world.
4. Understand how the scientific method works, and why science is an open-ended
process.
5. Appreciate the difficulties inherent in investigating the marine world, and the extent
to which it remains unexplored.
6. Start thinking about some of the fundamental concepts and questions in Marine
Biology.
steps for completing the lesson
1. Read the information for this lesson in the following pages of this Course Guide.
2. Complete the Reading Assignment (below) in C&H (Castro & Huber, Marine
Biology 9th Ed.).
3. Complete the Written Assignment (last page of the lesson) and submit it to UAF
eLearning & Distance Education.
Reading assignment
C&H, Chapter 1 (pp. 116) and Chapter 19 (pp. 425432)
More information, including interactive quizzes, flashcards, videoclips and links, can be
found at the publisher's Online Learning Center: www.mhhe.com/castrohuber9e
1
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M
Marine Biology may be simply defined as the scien-
tific study of living things in saltwater environments.
And here simplicity ends, for in this brief definition
we have just introduced three vast, complex and open-
ended subjects. Saltwater environments cover more than
70% of the Earth, and fill about 95% of its living space,
extending from shorelines lapped by the tides to trenches
as deep as nearly 11 km (7 miles) below sea level. Along
this continuum are numerous zones, both in the water col-
umn and along the sea floor, which span a huge range of
temperature, light, pressure and chemical conditions. The
varying effects of wind, waves, currents and tides, and the
outflow of rivers, also help to shape a rich diversity of ma-
rine environments. Saltwater habitats include sunlit surface
waters and vast realms of perpetual darkness, turquoise
tropical seas and ice-covered polar oceans, as well as salt
marshes, mangrove swamps, coral reefs, and deep-sea vol-
canoes and hot springs, among others. Yet, while scientists
have discovered a staggering variety of marine environ-
ments, more than 95% of the ocean remains to be explored.
In fact, the sea floor (most of the surface of the Earth) is
one of the more poorly mapped surfaces in our solar sys-
tem! What new ocean worlds are waiting to be discovered?
The variety of living things (biodiversity) residing in this
immense, varied and largely unexplored realm may be
equally vast and unknown. About 240,000 species of marine
organisms (not counting bacteria) have been described, dis-
playing incredibly diverse forms and life-styles. As many as
1.9 million marine and terrestrial species have been identi-
fied worldwide. Does the sea, which includes the vast major-
ity of living space on Earth, really contain only one-eighth of
Earths biodiversity? Or is this disparity due in part to our
greater ignorance of the marine world, compared to our own
terrestrial world? New organisms are being discovered all
the time (much faster, in fact, than they can be described,
or even catalogued) everywhere we look, especially at the
microscopic level. Is most of Earths biodiversity already
known to us, or only a fraction? Can we estimate, even to an
order of magnitude, the total number of species yet to be
discovered? Amazingly, we do not have conclusive answers
to these questions. More surprising still, we may never know
all the life that Earth contains, which brings us to the third
element in our definition of Marine Biology: scientific study.
Marine Biology is a very broad field of study, including many
different disciplines. Learning about life in the sea could in-
volve investigating specific chemical conversions performed
by marine bacteria in the laboratory, testing the communica-
tion skills of a group of captive dolphins, or using satellite
imagery and computerized mathematical models to gauge
the productivity of photosynthetic marine organisms on
a global scale. However, all of the different approaches to
studying marine life (as well as those in all other areas of
science) are conducted by following the same basic proce-
dure, called the scientific method. We use this time-tested
process to ensure that each piece of information contributed
to our store of knowledge is as reliable as possible. Scientific
knowledge is anything but stable, though, for all the quality-
control exerted by the use of the scientific method and by
the careful oversight, at many levels, of scientific research.
New information is continually being added by on-going re-
search, and new vistas are continually being opened by the
development of new techniques and equipment, and by new
ways of synthesizing and viewing information. Good science
often brings up more questions than it answers. In other
words, the more we know, the more we realize that we dont
know - and what we do know may pale in comparison to what
we dont know! So, science is an ongoing, open-ended pro-
cess. Scientific knowledge is constantly growing and is sub-
The Scope, History and Practice of Marine Biology
(Spying on the Sea: Ships and Sonar, Scuba and Submarines, Satellites and Sensors)
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ject to continual refinement, and even occasional revision,
like a huge complex building, perpetually under construc-
tion and continually being remodeled, with the occasional
demolition and reconstruction of some parts. It is a human
endeavor and, as such, is imperfect. This is the first of a
number of "themes" repeated throughout this course, im-
portant ideas to keep in mind as you explore Marine Biology:
1) Science, the key to our understanding of the world, is
an open-ended process, continually adding to, refining and
revising our knowledge.
2) The marine world comprises most of Earth's environ-
ments, is extremely varied, and largely unexplored.
3) The life inhabiting these environments also is extremely
diverse and may be mostly unknown to us.
4) Much of this life is smaller than the eye can see. Wher-
ever they look, scientists using new techniques are finding
microscopic organisms in unimagined numbers and diver-
sity, performing many vital functions in the ocean. In fact,
microorganisms appear to be the most abundant, diverse
and important (but least known) forms of life on the planet.
5) Another recurrent theme in this course is the interdepen-
dence of all organisms with each other and with their physi-
cal environments. Each living thing is an integral part of a
biological community and ecosystem. As a result of these
interactions, energy flows and nutrients cycle from organism
to organism, and between organisms and the environment.
At its most basic level, life is made up of these exchanges of
energy and matter.
6) The most important unifying concept in biology is that
of evolution. This course emphasizes an evolutionary
perspective, in order to help put the staggering diversity of
marine organisms into context, both with respect to the geo-
logic timescale (included as an Appendix to this Course
Guide) and to the Tree of Life, the family tree of relation-
ships among all living things.
7) As you will learn in this lesson, humans have always relied
on the ocean and its resources, in many ways. In the next
two lessons, you will learn that ocean waters and the sea
floor also regulate our atmosphere and climate and drive
weather-related phenomena such as drought, flooding, hur-
ricanes and tornadoes. They are responsible for many geo-
logical processes and events, including mountain-building,
earthquakes, volcanoes, tsunamis and even the position of
the continents on the globe! The ocean and its resources,
in turn, are increasingly subjected to impacts from human
activities. We are destroying marine habitats and marine life
on a huge scale and at an alarming rate, even causing the
extinction of species, exterminating unique forms of life,
in some cases even before they have been discovered. This
is perhaps the most important "take-home message" from
this course.
As you learn about Marine Biology, and come to a more
complete understanding of these essential themes,
also consider some of the questions that they raise:
What kinds of marine environments remain to be ex-
plored? How much of Earth's biodiversity is still un-
known to us, and what types of marine life are yet to be
discovered? How have marine organisms been shaped by
interactions with their biological communities and physi-
cal environments, and how will they continue to be shaped
by future interactions? How will current and future sci-
entific research change our understanding of the marine
world? Finally, what are the impacts of our activities on
the world ocean, and how will those impacts change ma-
rine environments and marine life, and affect conditions
globally? Our own future will be strongly influenced by these
changes, and will depend in part on how well we understand
the marine world, and how wisely we use that knowledge.
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Study the information presented for this lesson until you feel that you have a good understanding. Write your answers in your
own words, not those of the textbook or Course Guide. Be brief and clear, but also complete. For example, if asked to
"describe" or "explain" something, do so - don't just list. (To be clear and show your understanding, imagine that you are de-
scribing/explaining to someone who has no knowledge of the subject.)
Include a cover sheet and submit the written assignment to UAF eLearning & Distance Education according to directions found
online at eLearning.uaf.edu and in the course materials you received.
Complete the Planning Schedule on p. ix of this Course Guide. Return one copy of your schedule with this assignment (re-
quired), and keep a copy for yourselfcheck this schedule periodically to make sure you are on track to finish the course
on time!
1. Describe four ways that humans have traditionally used the ocean and its resources.
2. The voyage of the HMS Challenger (1872-1876) was the first worldwide expedition dedicated to the scientific
study of the ocean, and modern oceanography is based on its research. Describe three ways in which this voyage
fundamentally advanced marine science.
3. What do the following acronyms stand for: sonar, scuba, AUV, ROV, COOL, UNCLOS, EEZ?
4. Scientists can directly observe marine organisms at the shore, in the laboratory, on ships and manned submarines,
or by scuba diving. Describe four technologies that allow scientists to study marine organisms at a distance, without
being there in person and observing them directly.
5. If you turn in two assignments this week and two assignments next week, I might use inductive reasoning to con-
clude that you will turn in two assignments every week until you finish the course. If I have the general impression
that students complete this course on time, then I might deduce that you will finish the course on time. From your
own experience, give one example each of inductive and deductive reasoning (concerning any subject).
6. Define "testable hypothesis." Preferably from your own experience, give an example of a testable hypothesis and
briefly tell how it might be tested. Define "scientific theory."
7. The temperature of its surroundings is one variable that may influence an organism's functioning. What are three
examples of other variables that might affect it? When conducting an experiment to test whether increasing light
intensity increases the growth rate of seaweed, how would you make sure that other variables do not "confuse" the
results? (In other words, briefly tell how you would conduct a controlled experiment.)
8. The 1982 United Nations Convention on the Law of the Sea established a framework for regulating the use of the
world's oceans, providing for the conservation and equitable usage of the marine environment and its resources,
and addressing matters of sovereignty, rights of usage, and the resolution of disputes. As of 2012, 163 countries
and the European Union have signed and ratified the Law of the Sea treaty. Is the United States among them?
9. Explain why the study of life in the oceans is important. (give three reasons).
Written Assignment for Lesson 1
(Please make your answers complete and concise)
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The Benthic
environment
The benthic environment is vast, complex and largely unexplored. Submarine topography is
not only dramatic, it is also extremely dynamic.
1. Understand the concept of a world ocean and be able to identify its principal geo-
graphical regions: the major ocean basins, seas, bays and gulfs.
2. Have a basic knowledge of the internal structure of the Earth and the characteris-
tics of the continental and oceanic crusts.
3. Gain an historical perspective of the exploration of the sea floor and appreciate the
difficulties inherent in studying these environments, and the extent to which they
remain unexplored.
4. Be familiar with the "anatomy" of the sea floorits profiles, habitat zones and major
features.
5. Understand the process of plate tectonics and how it transforms the Earth's surface
and shapes the ocean environment.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 2 (pp. 18-38)
More information, including interactive quizzes, flashcards, videoclips and web-links, can
be found at the publisher's Online Learning Center: www.mhhe.com/castrohuber9e
2
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The Benthic Environment

T
To prepare for meeting the organisms that make the
sea their home, in Part 2 of this course, we need
some background knowledge. In the next two les-
sons, we will study the physical characteristics of the ocean
and the processes that created and continue to shape this
dynamic environment. The ocean includes two general habi-
tats, the environments where organisms live: benthic and
pelagic. The benthic ("depth") habitat is that on or in the sea
floor, and the pelagic ("sea") habitat comprises the water
column above the bottom. Each of these two general envi-
ronments is divided into several more specific habitat zones.
Here, we explore the physical features and processes of the
benthic environment, and in Lesson 3 we will study physical
and chemical conditions in the pelagic environment. Lesson
4 completes our preparation, by reviewing basic character-
istics of living organisms, and general principles governing
their interactions with each other and with their environments.
The world ocean, the interconnected expanse of seawater
covering 71% of the Earth, is classified into 4 major basins.
The Pacific, Atlantic, Indian and Arctic Oceans. Within the
major basins, smaller basins are specified (e.g., the Northwest
Pacific Basin and South Indian Basin), and many seas, bays
and gulfs are identified as arms or geographically specific
regions of the ocean. At the back of your textbook, there is
a fold-out map of the world ocean. Keep this map open
for reference, as you read about the benthic environment.
The ocean basins are not simply hollows in the Earth that
have collected water, but are chemically and geologically
distinct parts of the Earth's crust, and are subject to ex-
tremely dynamic processes. The crust is the outermost layer
of our multi-layered planet, which might be compared to a
peach. The Earth's inner core, outer core and mantle
correspond to the peach seed, pit and flesh, respectively.
The thinnest layer by far is the crust, corresponding to the
fuzzy peach skin. It holds the oceans, continents and all living
things. Oceanic crust, underlying the ocean basins, is rela-
tively thin. It is mainly composed of a dark-colored rock called
basalt, which is rich in iron and therefore quite dense. The
thicker continental crust consists largely of lighter-colored
granite, which is rich in silicon and aluminum and therefore
less dense, so the continents float higher on the underlying
mantle while the oceanic crust is submerged. Density differ-
ences between the two types of crust also figure significantly
in plate tectonics, the dynamic processes involving the entire
surface of the planet, which we will learn about in this lesson.
History of Seafloor Exploration and
Mapping
The benthic environment is vast, complex and still largely un-
known simply because it is so difficult to observe. We have
learned about the sea floor using bathymetry, the mea-
surement of depth in the ocean. Early measurements were
made by sounding (not related to the word "sound" but
from the Old English "sund" meaning sea/water/swimming),
lowering a weighted hemp rope from a ship until the weight
hit bottom. Depth was measured in fathoms, 6-ft incre-
ments of rope equivalent to the distance between a man's
outstretched arms. It could take hours to make one sounding,
the boat and line being subjected to movement by wind and
current during the process. Imagine how difficult it was, try-
ing to plumb the depths of the ocean and decipher the shape
of this immense, unseen surface, sounding by sounding!
The Survey of the Coast, the oldest scientific agency in
the U.S., was established by President Thomas Jefferson and
Congress in 1807 to conduct hydrographic (bathymetric)
surveys and chart the nation's coasts. The first hydrographic
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maps showed only the largest benthic features in shallow
coastal waters, detected by painstakingly looking for patterns
among many soundings. They served primarily to identify haz-
ards to near-shore shipping. During its early years, however,
the Coast Survey also was responsible for the discovery of the
continental shelf break and continental slope (from sound-
ings in the western North Atlantic in 1849), the first known
seafloor canyon (in Monterey Bay off the coast of California
in 1857) and the first indications of seamounts and trenches.
The deep-sea floor wasn't sounded until the mid-1800s.
Until then, a bottomless ocean (as some believed) was a
testable hypothesis that hadn't yet been proven false! The
first successful deep-water soundings were taken dur-
ing the 1839-1843 voyage of HMS Erebus and Terror
as they sailed toward Antarctica, under the command of
James Clark Ross, in search of the South Magnetic Pole. A
line was constructed on board that was 3600 fathoms long,
fitted at intervals with swivels and strong enough to sup-
port a weight of 76 lbs. On January 3, 1840, they reached
latitude 27
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26' S and longitude 17
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29' W, the middle of
the South Atlantic Ocean between Rio de Janeiro, Brazil and
Namibia. Here, in dead calm weather, they sounded the sea
floor at 2,425 fathoms (14,550 ft). Ross's crew went on to
make more deep-water soundings, including some deeper
than the first. It was the 1872-1876 HMS Challenger ex-
pedition that first conducted and documented deep-sea
soundings on a large enough scale to begin to give some
characterization to the floor of the world ocean, using 144
miles of rope and 12 miles of piano wire as they sailed
nearly 115,000 km through all the oceans except the Arctic.
The presence of life on the deep-sea floor also was unknown
until the mid-1800s. In 1843, the British naturalist Edward
Forbes declared that life didn't exist below 300 fathoms.
Sometimes referred to as the "Azoic Theory," we know it to
be a hypothesis, waiting to be disproved. Forbes' announce-
ment started a debate that lasted for 20 years, despite the
fact that Ross and crew had hauled up baskets full of corals
and other life from dredgings as deep as 400 fathoms during
their 1840-1843 expedition, which prompted Ross to write,
"although contrary to the general belief of naturalists, I have
no doubt that from however great a depth we may be able
to bring up the mud and stones of the bed of the ocean,
we shall find them teeming with animal life." The debate
was officially ended by dredging operations conducted from
1867-1869 (after Forbes' death). In 1867, the Coast Sur-
vey found prolific life as deep as 517 fathoms off the coast
of Florida, and in 1868, the Norwegian scientist Michael Sars
documented 427 species brought up from 450 fathoms off
the coast of Norway. Inspired by these results, the Scottish
naturalist Charles Wyville Thomson (a student of Forbes')
petitioned the British Royal Society and Admiralty to fund
dredging operations off the coasts of Scotland and Ireland,
in 1868 and 1869. Many productive hauls were made well
below 300 fathoms, including one at 4,289 m (2,346 fath-
oms), the deepest ever dredge haul at that time, emphatically
disproving Forbes' azoic hypothesis. It was these successes
that led to Thomson's organization of the HMS Challeng-
er expedition, which might not otherwise have happened.
Around the time of Challenger's voyage, hydrographic
surveying was greatly advanced by the replacement of hemp
rope with wire lines, the invention of the wireline sounding
machine, and deep-sea anchoring. It was truly revolution-
ized, however, by the development of sonar during World
Wars I and II. We saw in the previous lesson (C&H Fig. 1.7)
how sonar using a single beam of active sonar is used to
measure the depth of the sea floor (echo sounding).
Today, hydrographic mapping is carried out primarily using
multibeam sonar and side scan sonar, which produce
a broad swath of echo soundings (Fig. 1). The GPS (Global
Positioning System), the global satellite navigation system,
accurately determines the geographic position of the sur-
vey vessel and the location of each sounding on the Earth.
The huge amount of data generated is processed and im-
aged digitally with specialized computer systems. In addi-
tion to sonar, LIDAR (Light Detection And Ranging) systems
use pulses of laser light, emitted from aircraft and reflected
off objects on shore and in water down to 50 m, to pro-
vide seamless coverage of elevation and depth between
land and sea. LIDAR also allows bathymetric measurements
to be made safely along complex and rugged shorelines.
Side scan sonar and precise measurements of the relative
strength of reflected signals also can be used to determine
the texture and composition of the sea floor (rocky, sandy,
muddy, etc.) as well as depth. Seismic reflection and
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seismic refraction, echo sounding using explosive en-
ergy pulses, detect subsurface features down to hundreds
of meters within the sea floor (Fig. 1). Deep-sea drilling
has pulled up core samples of the earth from as deep as
nearly 7,000 ft beneath the ocean floor. DSDP, the Deep Sea
Drilling Project carried out with the vessel Glomar Chal-
lenger from 1968-1983, provided key evidence in support
of seafloor spreading and plate tectonics. It was succeeded
in 1983 by OCP, the Ocean Drilling Program involving a U.S.-
led consortium of 25 countries. OCP was followed in 2003
by IOCP, the Integrated Ocean Drilling Program, a U.S.- and
Japan-led effort. Its mission is to examine life deep under
the sea floor, seek information on climate change, and in-
vestigate mass and energy transfers between the crust and
mantle. In addition to drilling vessels and a fleet of sur-
vey vessels, submersibles like Alvin (C&H Fig. 1.10), and
AUVs and ROVs have continued exploring the ocean floor.
The Coast Survey, together with the Weather Bureau (formed
in 1870, the first agency dedicated to atmospheric science)
and the Bureau of Commercial Fisheries (formed in 1871,
the first conservation agency), is now a part of NOAA, the
National Oceanic and Atmospheric Administration, formed in
1970 as one of 12 agencies under the U.S. Department of
Commerce. NOAA conducts extensive and exciting ocean ex-
peditions and research around the globe, and you can find
a wealth of information on their ocean explorer website. We
have made immense strides in understanding the features
and processes of the sea floor, which probably could not
have been imagined by the explorers and scientists of the
1800s and early 1900s. Still, only about 10% of the sea floor
has been comprehensively mapped. (Half of that, however,
is available for you to explore at earth.google.com/ocean!)
Epilogue: The rest of the story of HMS Erebus and Ter-
ror is legendary and, in a strange way, comes full circle with
respect to seafloor exploration. After the Ross expedition
to the Antarctic, these sailing ships (which began their long
careers as bomb vessels in the Royal Navy) were outfitted
with steam engines and iron-plated hulls. In May of 1845,
they turned north toward the Arctic in an attempt to com-
plete the Northwest Passage, under the command of Sir John
Franklin. After a brief siting in Baffin Bay a few months later,
Erebus and Terror were never seen again. Over the next
150 years, many searches (including one headed by Ross)
and investigations have pieced together the broad outlines
Figure 1. Hydrographic mapping techniques.
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of the demise of the Franklin expedition, but the search for
the vessels continues. In 1992, Canada designated HMS
Erebus (fittingly, Greek for "deep darkness") and Terror as
a National Historic Site (their only undiscovered such site).
Since 2008, the Canadian Hydrographic Service, in collabo-
ration with Parks Canada, has been using side scan and mul-
tibeam sonar and an ROV to chart the sea floor in search of
the ships, and in an effort to strengthen Canada's claim of
sovereignty over large portions of a rapidly warming Arctic.
Dramatic Features
The sea floor can be described in superlatives. The most
extensive geographic feature on Earth, covering 60% of its
surface, is the abyssal plain. It lies at depths of 3,000-
5,000 m (10,000-16,500 ft) and is the flattest place
on the planet, covered with mud that is hundreds, even
thousands, of feet deep. The vast expanse of the abys-
sal plain is broken by many features. The most dramatic is
the mid-ocean ridge system, a colossal chain of volca-
nic mountains that encircles the globe, running through all
the ocean basins (C&H Fig. 2.5). Occupying nearly a third
of the ocean floor, this is the largest geologic formation on
Earth. Running along the top of the mid-ocean ridge is a
great depression, the central rift valley, the longest val-
ley on Earth. Individual volcanoes that grow to 1,000 m
or more above the sea floor, called seamounts, also can
be seen. When seamounts rise high enough to break the
surface, they become islands. One such volcano, Mauna
Kea, lying mostly submerged on the island of Hawai'i, is
the world's tallest mountain. It rises 9,632 m (31,601 ft)
from base to summit, more than 2,500 ft taller than Everest.
The lowest places on Earth also are found on the sea floor,
at trenches. The Mariana Trench, near Guam in the west-
ern Pacific, holds the deepest place known, the Challenger
Deep, plunging to about 11 km (7 mi) below sea level. If
Mt. Everest were tipped into the Challenger Deep, it would
sink out of sight and it's base would be covered by more
than a mile of water. In 1960, a real-life "Voyage to the Bot-
tom of the Sea" took place when Jacques Piccard, a Swiss
undersea explorer, and U.S. Navy Lieutenant Don Walsh, an
engineer, descended into the Challenger Deep in the bathy-
scaphe ("deep boat") Trieste. Her descent took 4 hours
and 48 minutes, the ascent, 3 hours and 15 minutes. Pic-
card, the son of Trieste's designer, Auguste Piccard, de-
scribed the touchdown thus: "the Trieste balanced herself
delicately on the few pounds of guide rope that lay on the
bottom, making token claim, in the name of science and hu-
manity, to the ultimate depths in all our oceans - the Chal-
lenger Deep." Their feat was not repeated for more than 50
years, until James Cameron (director of Titanic and The
Abyss, among other films!) also descended into the Chal-
lenger Deep, in a submersible he designed, on Mar. 26,
2012. His descent took about 2 and a half hours, the ascent
only 70 minutes ("heckuva ride!"), and live updates of the
event were posted on Twitter from a yacht accompanying the
expedition. The Japanese-built, remote-controlled submers-
ible Kaiko also paid a visit to the Challenger Deep in 1995.
Many other dramatic features of the benthic environ-
ment are known, including submarine canyons.
The Hudson Submarine Canyon off the coast of New
York is deeper, wider and longer than the Grand Can-
yon. With most of the sea floor still unexplored, what
other amazing formations remain to be discovered?
Dynamic Processes
The sea floor not only displays dramatic features, it also un-
dergoes extremely dynamic processes. Sediment continually
rains down on the abyssal plains, and avalanches of rock and
sand sweep through submarine canyons. Lava erupts from
cracks in the sea floor, forming volcanoes, which grow into
seamounts. Seamounts rise above the surface to become is-
lands, and islands erode away and sink below the surface,
again becoming seamounts. Guyots (or tablemounts) are
flat-topped seamounts, formerly islands, whose tops became
eroded by wind and waves as they subsided. Chains and
other groupings of islands also form over "hot spots" in the
ocean's crust where columns of hot rock from deep in the
earth (mantle plumes) burn through (although there is
debate about this: C&H pp. 36-37). About 50 hot spots have
been identified. Hot spots under parts of the mid-ocean ridge
system are thought to have formed islands like the Azores
and Iceland (C&H Fig. 2.7) on the Mid-Atlantic Ridge,
and the Galapagos Islands at the Galapagos Spreading
Center, part of the mid-ocean ridge system in the east-
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ern Pacific off the coast of Ecuador. Since 2005, scientists
have been anticipating the birth of an island where another
hotspot volcano, the Vailulu'u seamount near the Samoan Is-
lands, is rising from the sea floor. Using bathymetric mapping
techniques and deploying a submersible and ROV, they found
that a new cone, 300 m tall, had appeared within the crater
of this seamount since the last bathymetric measurements
in 2001. Named Nafanua after the Samoan goddess of war,
this new cone could breach the surface within decades, at
its present rate of growth. If that happens, a new island
would be born, a highly revered event in Polynesian culture.
The sea floor also is home to dynamic biological activity. The
summit of Nafanua already supports a thriving community of
marine organisms. Many tropical volcanic islands are fringed
by coral reefs, the richest and most complex marine eco-
systems. Even the deep-sea floor holds astonishing oases
of life around hot springs called hydrothermal vents,
discovered in the late 1970's with the aid of Alvin. The dis-
covery of these vents changed our understanding of the
chemistry, physics and biology of the ocean. They occur in
association with the mid-ocean ridge system and trenches,
when seawater seeps down into countless cracks in the sea
floor, becomes superheated, and is vented back up to the
surface. Often the water emitted from these vents is laden
with minerals, producing dense clouds which solidify and
build into chimney-like structures called "black smokers"
(C&H Figs. 2.26 and 2.27). It is thought that all the ocean's
water passes through the earth's crust in this way, recy-
cling itself completely about once every 6-8 million years.
In addition to all the volcanic activity (volcanism) that pro-
duces hydrothermal vents, seamounts and the mid-ocean
ridge system, the sea floor also is subject to much seismic
("shaking") activity. Frequent earthquakes characterize the
areas near trenches, the mid-ocean ridges and other volca-
noes. Vailulu'u, for example, experiences an average of four
earthquakes a day. As a result of all this seismic activity, the sea
floor is littered with faults, or rifts, where the brittle oceanic
crust has fractured. Normal faults, where the blocks on ei-
ther side of the fracture have been displaced vertically from
each other, are prominent along the mid-ocean ridge system,
running parallel with its axis. The blocks at the crest of the
ridge have dropped, creating a depression running along the
crest line, the central rift valley (Fig. 2 and C&H Fig. 2.25).
The mid-ocean ridge system also displays many transform
faults, where the two sides of the fracture slip past each
other horizontally. Transform faults occur perpendicular to
the axis of the ridge, offsetting it in many places so that it
zigzags through the ocean basins (Fig. 2 and C&H Fig. 2.5).
In fact, the entire sea floor is constantly in motion. Molten
magma is continually rising up from the mantle at the crest
of the mid-ocean ridges, giving birth to new sea floor that
spreads outward, as if on a conveyor belt (moving at about
the same rate as your fingernails growing). This process is
called seafloor spreading (C&H Fig. 2.10). At the same
time, subduction is occurring at the trenches, where old
sea floor is being pushed back down and recycled into the
mantle. Spreading zones at mid-ocean ridges, and sub-
duction zones at trenches, mark the boundaries of great
lithospheric plates, which partition the Earth's surface
like giant puzzle pieces arranged over the globe (C&H Fig.
2.11). As these plates grow away from the spreading zones,
they collide with neighboring plates. When a collision involves
oceanic crust on one plate edge and continental crust on the
other, the lighter continental crust overrides the dense oce-
anic crust, pushing it into the mantle and creating a trench
(C&H Fig. 2.12). When the two converging plates both carry
oceanic crust, one overrides and pushes the other down,
also creating a trench (C&H Fig. 2.13). Earthquakes and
volcanoes are common near these subduction zones (C&H
Fig. 2.8), as the subducted plate edge begins to break up
and melt on its way down into the mantle. When two plates
bearing continental crust collide, neither is subducted and
great mountain ranges are created as the plates buckle
against each other (e.g. the Himalaya, where the Indian Plate
is being pushed into the Eurasian Plate). In addition to the
spreading zones at mid-ocean ridges and the subduction
zones at trenches, plates interact in a third way, along a third
type of plate boundary called a shear boundary. Exempli-
fied by the San Andreas Fault in California (C&H Fig. 2.15),
a shear boundary involves two plates scraping past each
other (i.e., transform faulting), catching and building tension
which is eventually released explosively, causing more earth-
quakes. As a result of these plate interactions, oceanic crust
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is relatively thin and short-lived, since it is continually being
recycled, while continental crust is being built up and mostly
endures. The oldest known continental rocks are nearly 4 bil-
lion years old, whereas the oldest sea floor was "just" formed
some 200 million years ago. Continents are built up by a
process called accretion, which occurs during subduction,
as sediments on the subducted plate are scraped off and
plastered onto the overiding plate. Lava erupting from volca-
noes is another way that material is added to the continents.
Plate tectonics (tectonics means "to build") is the all-en-
compassing term describing the creation, movement, defor-
mation and destruction of the lithospheric plates, through
the cyclic creation, spreading and subduction of the sea floor.
The story of how the theory of plate tectonics developed, re-
counted in your textbook, is a great example of science as an
ongoing processgradually expanding and refining, and oc-
casionally making huge leaps or sweeping alterations, in our
understanding of the world. The latter typically face strong
opposition at first, gaining acceptance only after the accumu-
lation of overwhelming evidence. Often this is achieved with
the aid of new technologies, or the synthesis of information
from different disciplines. All of these factors were involved
in the development of plate tectonics theory, which has rev-
olutionized our understanding of this dynamic planet. The
unifying theory of geology, it draws together evidence from
disparate branches of science (e.g., seismology and paleon-
tologythe study of fossils) to explain where and why earth-
quakes, volcanoes and mountain-building occur, how ocean
basins open and close, and how continents drift, collide and
split apart, continually transforming the face of our planet.
Originally, almost all the crust was probably sea floor, and
cycles of spreading and subduction were rapid. By about
3 billion years ago, granitic rock had built up at the sur-
face to form numerous microcontinents, which eventually
clumped together to form the core of our modern conti-
nents. Around 220 million years ago, when dinosaurs ap-
peared, the Earth held one supercontinent, Pangaea ("all
earth"), and one gigantic ocean, Panthalassa ("all sea").
Two bays off Panthalassa enclosed enormous seas, the Te-
thys Sea and Sinus Borealis. As rifts formed and flooded
with seawater, Pangaea began to break apart into smaller
continents, separating smaller oceans and seas. The rift
dividing Pangaea into Laurasia, to the north, and Gond-
wana, to the south, filled with water from the Tethys Sea,
creating the North Atlantic Ocean. As Gondwana broke apart,
India split off and moved northward, establishing the Indian
Ocean, and South America moved away from Africa, creating
the South Atlantic Ocean; the Sinus Borealis shifted north
Figure 2. Hydrologic cycle.
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to become the Arctic Ocean. Fig. 2.17 in your textbook il-
lustrates these changes, from about 200 million years ago.
Today, the Atlantic, Indian and Arctic Oceans are "mature"
basins, still expanding by seafloor spreading. The Pacific
Ocean (the remnant of Panthalassa) is a "declining" ba-
sin, continuing to shrink as its borders are subducted.
The Red Sea is a "juvenile" basin, on its way to becom-
ing a mature broad basin, while an "embryonic" ocean
basin is forming in the rift valleys of East Africa. Far into
the future, the configuration of oceans and continents will
again be very different from the arrangement we take for
granted today, as the process of plate tectonics continues.
Other factors also change the relationship between the
ocean basins and continents, through geologic time. Land-
masses sink or rise as a result of weight being added or
removed. Loading happens when ice accumulates in glaciers,
lava is extruded, or sediments accumulate. Unloading occurs
through the melting of glaciers, weathering and erosion of
rocks, or extraction of groundwater or oil. Subsidence or
uplift of the Earth's surface on a local scale due to such
processes, as well as to earthquakes and fault motions, are
referred to as isostatic changes. Other processes result
in global adjustments to sea level, called eustatic chang-
es. Change in the volume of ocean basins due to seafloor
spreading is an example of a eustatic change. Climatic fluc-
tuations also produce eustatic changes. During glacial pe-
riods, when more of the Earth's water accumulates on land
as ice and snow, sea level falls. During interglacial peri-
ods, sea level rises again as glaciers melt. The volume of
the ocean also expands when its temperature increases, and
contracts when it cools. We are now in an interglacial period,
and sea level today is about 130 m (425 ft) higher than at
the peak of the last ice age, about 18,000 years ago. Most
of the glaciers from that ice age had melted by about 10,000
years ago, and sea level had risen very little in the last few
thousand years. In recent decades, however, atmospheric
and ocean temperatures have been increasing. Warming
waters are expanding and glaciers are melting at an accel-
erating rate. The impacts of these eustatic changes in sea
level are already being felt, especially in coastal areas. Global
changes like these, due in large part to an enhanced green-
house effect, will be discussed more in the next lesson.
The Sea Floor Profile
Because these processes work to shape the sea floor in
similar ways around the globe, all the ocean basins share
the same general features (C&H Fig. 2.20). They all have
two main regions: the continental margin, and the deep-
sea floor. The continental margin generally exhibits a gently
sloping continental shelf, where the edge of the conti-
nental crust is submerged, a shelf break that more or less
marks the end of the continental crust, and a more steeply
sloping continental slope. When sediment accumulates at
the base of this slope, a continental rise will also form.
The abyssal plain stretches away from the continental mar-
gin, to form the deep-sea floor. The profile of the continental
margin in a particular area is shaped by the type of plate
tectonic processes operating there (C&H Fig. 2.23). The
continental margin is covered by a thick wedge of sediment
eroded off the continent. When the edge of a continent oc-
curs near an advancing plate edge, where two plates are
colliding and subduction is occurring, sediments are carried
down into the trench, and a sharp and dynamic profile forms,
called an active margin. When a continent does not oc-
cur at a plate edge, or is on a receding edge, then it has
a passive margin with the wedge of sediment forming a
long shelf (more than 1,000 km, or 620 mi, in the case of
the Arctic Ocean), a gentle slope and a pronounced rise.
Benthic Habitat Zones
Corresponding to this profile of the ocean basins, there are
five divisions of the benthic environment: the intertidal, sub-
tidal, bathyal, abyssal, and hadal zones (Fig. 3). Conditions
differ in each zone, and each supports different communities
of organisms. The intertidal (or littoral) zone is the area
between the low and high tide marks, exposed to the sun and
waves, and to the air when the tide is out. The subtidal (or
sublittoral) zone runs from the low tide mark to the shelf
break, including the continental shelf below the intertidal. It
remains submerged at low tide, but is exposed to sunlight,
waves and currents. The bathyal (from the Greek bathys,
"deep") zone extends from the shelf break (usually 120-200
m deep) to about 4000 m, including the continental slope and
rise, and much of the mid-ocean ridge system. It is subject to
increasing darkness, cold and pressure. The abyssal (from
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the Greek abyss, "bottomless") zone comprises the sea floor
from 4000 to 6000 m, the abyssal plains and deep ocean
basins. The hadal (from Hades, the classical Greek under-
world) zone refers to the bottom below 6000 m, the trench-
es. Both the abyssal and hadal zones are completely without
sunlight and subject to extreme pressure and cold, except at
hydrothermal vents, which are the hottest habitats on Earth.
The bathyal, abyssal and hadal zones together make up the
"deep-sea floor." In the next lesson, we will fill in this basin
of benthic zones with the pelagic habitat zones, when we
study the physical characteristics of the pelagic environment.
Shelf Break
Baythal
Hadal
Abyssal
Subtidal
High Tide Mark
Low Tide Mark
Intertidal
Shelf Break
Baythal
Hadal
Abyssal
Subtidal
Benthic Habitat Zones
High Tide Mark
Low Tide Mark
Intertidal
Figure 3. Benthic habitat zones..
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glossary terms
Listed below are some key terms for this lesson. Do a self-check for understanding. If you're unclear about the meaning of
any of these terms, re-read the material in this course guide and in your textbook, and check the glossary at the back of
your textbook.
benthic and pelagic
water column
world ocean
Pacific, Atlantic, Indian, Arctic, and Southern
Oceans
inner core, outer core, mantle, and crust
oceanic crust and continental crust
basalt and granite
asthenosphere and lithosphere
lithospheric plates and plate tectonics theory
continental drift
bathymetry/hydrography
sonar/echo sounding and LIDAR
lithogenous sediment and biogenous sedi-
ment
calcareous ooze and siliceous ooze
abyssal plain
mid-ocean ridge system, Mid-Atlantic Ridge
and East Pacific Rise
central rift valley
seamount and guyot
submarine canyon
hot spot/mantle plume
trench, Mariana Trench, and the Challenger
Deep
hydrothermal vent and black smoker
volcanism and seismic activity
fault/rift, normal fault and transform fault
seafloor spreading and spreading zone
subduction and subduction zone
shear boundary
magnetic anomalies
accretion
Pangaea and Panthalassa
Gondwana and Laurasia
isostatic and eustatic
continental shelf, shelf break, continental
slope and continental rise
passive margin and active margin
intertidal (littoral), subtidal (sublittoral),
bathyal, abyssal and hadal
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"describe" something, do so - don't just list. (To be clear and show your understanding, imagine that you are explaining to
someone who has no knowledge of the subject.)
1. Define the following pairs of terms. Be brief, but also include all the essential information. Comparing/contrasting the
two terms will often help you to do this. For example:
normal faults and transform faults: normal faults are cracks in the earth's crust where the two sides are
displaced vertically, while in transform faults the sides are displaced horizontally. Along the mid-ocean ridge, normal
faults are parallel to the axis while transform faults are perpendicular to the axis of the ridge.
a. benthic and pelagic
b. asthenosphere and lithosphere
c. Gondwana and Laurasia
d. lithogenous sediments and biogenous sediments
e. sonar and LIDAR
f. Trieste and HMS Terror
g. Glomar Challenger and the Challenger Deep
h. Mariana Trench and Mauna Kea
i. guyots and Galapagos Islands
j. active margins and passive margins
2. Name the four major basins of the world ocean. Which continents separate which oceans from each other?
3. Identify the two principal types of rock that distinguish the continental and oceanic crusts. Describe three other
important differences between continental and oceanic crust.
4. Observation is the currency of science. When scientists towed a magnetometer above the mid-ocean ridge and plot-
ted the data on a map, they observed a surprising pattern of magnetic "stripes," a discovery that played a key role
in the development of the theory of plate tectonics. Describe the pattern they observed and explain how it supports
the hypothesis of seafloor spreading. Describe two other kinds of observations of the sea floor around the mid-
ocean ridge that scientists have made, and explain why they also support the hypothesis of seafloor spreading.

Written assignment continued on next page
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5. The three major features of the Earth's crust that form the boundaries of the lithospheric plates are the central rift
valley of the mid-ocean ridge system, trenches, and shear boundaries. Describe the processes occurring at each of
these boundaries.
6. Put on your scuba gear and take a walk across the ocean floor, along a direct route starting at Cape Cod, Massa-
chushetts, and ending at the west coast of Portugal. Describe or diagram (with labels) 11 specific seafloor features
you have learned about in this lesson, which you might see along this path, including features on the continental
margins and deep-sea floor. Describe these features in chronological order, as you would encounter them along
this route. (Use the fold-out map at the back of the textbook, other maps in C&H Chapt. 2, and the illustrations of
continental margins. You may also use your imagination to come up with some geologic features you might reason-
ably expect to find on your trek.)
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3
The Pelagic
environment
Water is a simple molecule with complex and extraordinary properties. It is the
primary requirement for life and the principal constituent of living things. Close to
98% of the water that cycles on Earth is held in the ocean. Water also is continually
being circulated within the ocean, in complex patterns that strongly influence both
marine environments and global climate, thus affecting all of Earth's inhabitants.
1. Appreciate the extraordinary chemical and physical properties of water, and be
familiar with the hydrologic cycle.
2. Be familiar with the chemical composition of seawater, and the physical characteris-
tics of the pelagic environment, including sea ice.
3. Learn how the ocean is structured vertically, according to temperature, salinity and
density.
4. Understand the concept of thermohaline circulation and its impact on global climate
patterns.
5. Recognize the major surface currents of the ocean, how they are generated and
how they regulate climate, and have a basic understanding of waves and tides.
6. Be able to classify the pelagic environment into habitat zones.
7. Understand the significance of the ocean in the heat balance of the planet, and as a
moderator of the greenhouse effect and global warming.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 3 (pp. 4062), and "Special Report" pp. 231237 ("Rolling The Dice:
Climate Change" and "Ocean Acidification: The Other CO2 Problem") and p. 239 ("Will
There Be a Last Straw?")
http://highered.mcgraw-hill.com/sites/0073028193/student_view0
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The Pelagic environment

C
Contrary to its name, "Earth" is really the water
planet. With 74% of its surface covered by water,
71% by seawater at an average depth of 3.7 km
(2.3 mi), our world is a predominantly blue sphere when
viewed from space. This water originated as steam emitted
from the molten rock on the early surface of the planet, as
it began to cool and harden. Comets and meteorites striking
the Earth have also contributed water, and volcanic erup-
tions continue to bring steam to the surface. Our distance
from the sun, and the complex processes that created and
maintain our atmosphere, are in large part responsible for
retaining this water and keeping it in a liquid state, the pri-
mary requirement for life. Processes occurring in the ocean,
in turn, strongly influence Earth's atmosphere and climate.
Venus, our neighbor closer to the sun, has an atmosphere
100 times denser than Earth's, with a runaway green-
house effect producing a surface temperature of about
450C (800F). Any liquid water it once held would have
boiled away eons ago. On the other side is Mars, with a much
thinner atmosphere than ours and an average temperature
of 60C. The Martian landscape appears to have been
eroded by flowing water in the distant past, but is now too
cold for liquid water to persist at the surface, although ice
caps occur at the poles and permafrost exists in the ground.
Water is a simple molecule with complex and extraordinary
properties. It is the primary requirement for life and the
principal constituent of living things. The availability of water
largely determines the distribution of life on land. About 95%
of the water on Earth's surface is bound into rocks. The re-
maining 5% participates in the hydrologic cycle (Fig. 4),
as it evaporates from the oceans, lakes, rivers and land, and
is transpired from the leaves of plants, then forms clouds
which are blown by the wind over the land or sea, eventually
precipitating in the form of rain or snow. On land, this pre-
cipitation refills lakes and rivers, builds glaciers, percolates
into the earth to become groundwater, and is again taken
up by the roots of plants. Glaciers melt, surface waters drain
into rivers which flow to the sea, and most groundwater finds
its way to the ocean as well, as the cycle continues. Close to
98% of this cycling water is held in the ocean. Water also
is continually being circulated within the ocean, in complex
patterns that strongly influence both marine environments
and global climate, thus affecting all of Earth's inhabitants.
Chemical Composition
Seawater is composed, on average, of 96.5% water and
3.5% dissolved materials, or solutes. Water can dissolve
more kinds of solutes (i.e., is a better solvent) than any
other natural substance, one of its many remarkable prop-
erties. Most of the solutes in seawater are salts, predomi-
nantly sodium chloride (NaCl - table salt), which dissociate
in water into their component electrically-charged ions: so-
dium (Na+) and chloride (Cl-), in the case of NaCl. The main
types of ions present in seawater are listed in C&H Table
3.1. Salinity, the amount of salt dissolved in the water, may
be expressed as a percentage, or as psu (practical salinity
units) or ppt (parts per thousand), all of which are numeri-
cally equivalent. The average salinity of seawater is 35 psu
(or ppt or %). Salinity in some places may be higher, usually
due to the removal of water (through evaporation or the
formation of ice), or lower due to the addition of water
(through precipitation, river runoff or the melting of
ice). However, the proportions of the major constituent
ions of seawater (C&H Table 3.1) remain more or less the
same, regardless of salinity, throughout the world ocean.
This rule of constant proportions is another important
discovery that came out of the HMS Challenger expedition.
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In addition to the major ions, small amounts of virtually all
other chemicals can be found in seawater. Nutrients, the
materials needed for life, include the major ions and other
required substances which may occur in nearly undetect-
able, yet critical, concentrations in seawater. The concentra-
tion of some nutrients can vary greatly in the ocean and
nutrient concentration, as well as salinity, strongly influences
the functioning and distribution of marine life. Ions and
other nutrients enter the ocean in a variety of ways (C&H
Fig. 3.6). Weathering and erosion release materials from
rocks and soil, which are washed into rivers or picked up by
winds, and carried to the sea. Chemicals also are released
from the sea floor, through hydrothermal vents and subma-
rine volcanoes, and from land volcanoes whose emissions
rain down on the ocean. Pollutants, chemicals released
by humans which negatively affect the quality of the envi-
ronment (like fertilizers and exhaust fumes), also find their
way into the ocean and can profoundly affect marine life.
Gases, as well as solids, dissolve in water and are important
constituents of seawater. Gases diffuse across the air-sea
interface, moving from a higher to a lower concentration un-
til equilibrium is reached, a process called gas exchange.
Turbulence due to waves and currents accelerates this
process. As the most abundant gases in the atmosphere,
nitrogen(N
2
)and oxygen (O
2
) readily enter the sea surface
through gas exchange. Carbon dioxide (CO
2
) is present at
very low concentrations in the air, but its concentration is 500
times greater in surface waters, making it the most prevalent
gas in the ocean. The ocean holds 50 times more carbon
than in the atmosphere. This is because CO
2
reacts chemi-
cally with water, becoming incorporated into bicarbonate
and carbonate ions, which allows more CO
2
gas to be ab-
sorbed at the sea surface through gas exchange. Gas con-
centrations vary in different regions and at different depths
in the ocean. Generally, the solubility of gases increases with
decreasing salinity, decreasing temperature, and increasing
pressure. The primary influence on gas concentrations below
the surface of the ocean, however, is the activity of marine
organisms. During respiration, marine organisms take up
oxygen and release carbon dioxide, while during photosyn-
thesis they produce oxygen and absorb carbon dioxide.
Many marine organisms also incorporate carbon dioxide into
skeletons and shells made of calcium carbonate. Over
Figure 4. Hydrologic cycle.
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eons of time, immeasurable quantities of these shells have
accumulated on the sea floor, contributing to biogenous
sediments and forming carbonate rocks like limestone.
Through their photosynthesis, marine organisms are respon-
sible for releasing into the air about half of the oxygen that
we breathe. Also through photosynthesis, and the calcium
carbonate shells and skeletons that they build, marine or-
ganisms are responsible for absorbing huge quantities of
carbon dioxide from the air. Atmospheric carbon dioxide
is a greenhouse gas that prevents the escape of heat
radiated from the surface of the Earth, causing the atmo-
sphere to become warmer. Since the industrial revolution,
this greenhouse effect has been increasing as a result of
increasing levels of carbon dioxide released from the enor-
mous quantities of fossil fuels that we have been burning
(about 3 tons of carbon dioxide are produced for each
ton of carbon burned), and from cement production. Global
warming is the result of this increasing greenhouse ef-
fect. The ocean and its inhabitants have been moderating
the greenhouse effect and global warming, by absorbing
and storing huge quantities of carbon dioxide, about half
of that produced by human activities in the past 200 years.
While the increased uptake of carbon dioxide has moder-
ated warming of the atmosphere, it has also altered sea-
water chemistry. The addition of carbon dioxide shifts the
balance of chemicals in seawater, producing a higher con-
centration of hydrogen ions, or lowering pH, making
it more acidic. The hydrogen ion concentration near the
surface of the ocean has increased about 30% since pre-
industrial times and may increase much further if the pres-
ent trend continues. This acidification of the ocean can
affect both physical processes (e.g., increasing the rate of
weathering of rocks) and biological processes (e.g., limit-
ing the ability of marine organisms like shellfish and cor-
als to form shells and skeletons of calcium carbonate).
Temperature
Water has many unusual physical properties, in addition to
its unsurpassed solvent power. One of the most important
is its unusually high heat capacity (the amount of heat re-
quired to raise the temperature of 1 gram of a substance
by 1C). The ability of water to absorb a large amount of
heat without a large increase in temperature, coupled with
the tremendous volume of water in the ocean, means that
temperatures remain relatively stable and homogeneous
in marine environments, compared to terrestrial environ-
ments. This is of crucial importance to marine life, as all
organisms are greatly affected by changes in temperature.
The ocean's enormous capacity to absorb and store heat,
moreover, makes it the dominant component in the heat bal-
ance of the Earth. While the ocean has been moderating the
increasing greenhouse effect by absorbing huge quantities
of carbon dioxide, it also has been moderating the warming
of the atmosphere by absorbing and storing an immense
amount of heat from the atmosphere. Scientists have cal-
culated a mean temperature increase of 0.037C in the up-
per 3000 m of the world ocean during the latter half of the
twentieth century. Although this increase may seem small, it
reflects an incredible amount of heat energy absorbed by
the world ocean, in fact 84% of all the excess heat add-
ed to the entire global system during this period of global
warming. To put this in perspective, scientists calculate that
a mean temperature increase of 0.1C of the world ocean
would equal a mean temperature increase of 100C of the
global atmosphere, if all the heat required for this increase
in ocean temperature were transferred to the atmosphere!
Just as the absorption of excess carbon dioxide by the ocean
has limits and consequences, so too does the absorption of
excess heat by the ocean. We will see that changing ocean
temperatures directly affects the marine environment and
its inhabitants in many ways. Ocean warming also has many
indirect effects, for example decreasing oxygen solubility.
Changing temperatures produce changes in the circulation
of the ocean, which in turn regulates weather patterns and
climate globally, thus affecting all of Earth's inhabitants.
Temperature varies in complex ways in the ocean. Surface
temperature varies according to location. Because the sun is
more directly overhead at the equator than at higher latitudes,
its rays are focused onto a smaller area, so insolation (the
amount of solar energy striking the Earth's surface) is great-
er, warming the atmosphere and sea surface more at the
equator and tropics, and less toward the poles (Fig. 5). This
effect is intensified by the higher albedo (reflectivity) of the
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ice and snow covering the poles, as they reflect more of the
sun's light (and heat) than do the darker-colored sea and land
surfaces at lower latitudes. Satellite imagery gives us a global
picture of sea-surface temperatures (C&H Fig. 3.10).
Because both insolation and the extent of ice and snow cover
vary seasonally, sea-surface temperatures also vary season-
ally. This picture is made more complex by surface cur-
rents, which carry warm water from the equator to higher
latitudes, and cold water in the opposite direction, in complex
patterns that are driven by global wind systems. These sur-
face-current patterns also vary seasonally, with insolation.
Longer term variations in these patterns occur as well, most
notably due to periodic El Nino-Southern Oscillation
(ENSO) events, which we will learn more about later. Chang-
es in the patterns of sea-surface temperatures (such as with
an ENSO event) have dramatic consequences for marine life
and produce major shifts in weather around the world. Sea-
surface temperatures are now changing, in unpredictable
ways, due to the intensifying greenhouse effect. The conse-
quences of these changes to ocean environments and ocean
life, and to Earth's climate and weather systems, are of great
concern. The major surface currents, the global wind sys-
tems that produce them, and other sea-surface phenomena
(waves and tides), are described in detail in your textbook.
We have seen that temperature varies in complex ways near
the surface of the ocean, but this is just the 2-dimensional
picture. To fully understand the important role that temper-
ature plays in the ocean, we must look below the surface.
Measuring temperature at different depths in the water col-
umn produces a temperature profile (C&H Fig. 3.22). At
temperate and tropical latitudes, a relatively shallow band
of warm surface water floats over a much broader band of
near-freezing deep water. In between the surface and deep
waters, from about 200 m to about 1000 m (660 to 3,300
ft) below the surface, there is an intermediate band of water
where the temperature gradient (rate of change with depth)
is steep, called a thermocline. This thermocline is a persis-
tent feature in these regions and is referred to as the per-
manent thermocline (C&H Fig.3.22a). At polar latitudes,
the surface water is much colder, so the thermocline is much
less pronounced than at lower latitudes (C&H Fig. 3.22b). In
summer months, however, a shallower seasonal thermo-
cline may develop within the surface layer of temperate and
polar waters, as this layer is warmed by increased insolation
Figure 5. Insolation (greater at the equator than the poles).
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(C&H Fig. 3.22c). Diurnal thermoclines also occur at very
shallow depths, as the surface water heats up on a daily ba-
sis. Salinity also may vary with depth, and a gradient called
a halocline is seen in some salinity profiles (C&H Fig.
3.22a), although usually not as steep or consistent as the
permanent thermocline. In addition to their direct effects on
marine organisms, the salinity and temperature of seawater
are of crucial importance because they determine its den-
sity, and density controls the vertical structure of the ocean.
Density: Stratification and Stability
Density (the amount of mass per unit volume) increases
with decreasing temperature or with increasing salinity, so
the presence of a thermocline or halocline produces a steep
density gradient called a pycnocline. Density profiles for
much of the ocean, at low and middle latitudes, tend to look
like mirror images of temperature profiles for these regions,
with a pycnocline of increasing density mirroring the perma-
nent thermocline (C&H Fig. 3.22a). Because denser water
sinks, this results in water stratification, which imposes a
three-layered structure on the ocean: a surface layer, an
intermediate layer (also called the pycnocline layer),
and a deep layer. The thin surface layer (about 100200
m or 330660 ft thick) represents about 2% of the ocean's
volume. It is warmed and lit by the sun, subject to evapora-
tion, precipitation, freezing, the outflow of rivers, and gas
exchange with the atmosphere. This layer also can be mixed
by wind, waves and surface currents, and is affected by daily,
seasonal and annual fluctuations in weather and climate. It is
here that changes in temperature and salinity take place. The
intermediate or pycnocline layer is a transition zone, where
the surface layer grades into the deep layer, and represents
about 18% of the ocean's volume. The deep layer consists
of cold, dense water sitting below about 1500 m (5,000 ft),
comprising 80% of the ocean's volume. It actually includes
two water masses, deep water and the even deeper bot-
tom water, both essentially out of reach of the processes
that influence temperature and salinity. Since their tempera-
ture and salinity are more or less fixed, deep and bottom
water masses from different regions of the world ocean
have unique temperature-salinity "signatures" en-
abling scientists to identify them and track their movements.
Stratification creates a stable water column, with the dif-
ferent layers effectively separated from each other, which
has important consequences. Because matter sinks in the
ocean, essential nutrients accumulate in deep water and
bottom sediments, remaining out of reach of organisms
in the surface layer. Thus, nutrient profiles in the open
ocean often appear similar to density profiles, with low levels
near the surface and high levels in deep water (C&H Fig.
15.28, p. 352). Stratification also results in a distinct pat-
tern of oxygen distribution in the water column at middle
and low latitudes. Oxygen levels are high near the surface
because of gas exchange with the air, and oxygen produc-
tion by photosynthetic marine organisms. In deep water,
oxygen is not produced by photosynthesis, due to the lack
of light, but it also is not used up quickly by respiring organ-
isms, because this vast layer is so sparsely populated. In
the pycnocline layer, on the other hand, while oxygen is not
being generated by photosynthesis, it is being depleted by
respiring organisms, which results in an oxygen minimum
zone at these intermediate depths (C&H Fig. 16.18, p. 372).
In polar regions, the chilling of surface water generally pre-
vents the formation of a strong pycnocline and stratifica-
tion, resulting in a relatively unstable water column. As
surface water cools, it becomes more dense and sinks, a
process called downwelling. Downwelling occurs whenever
the temperature of surface water decreases or its salinity in-
creases (through evaporation or the formation of ice on the
surface), to an extent that it becomes more dense than the
water below, causing it to sink. An opposite movement, called
upwelling, is produced as deeper water is displaced up-
ward by the sinking water mass. Together these movements
are referred to as overturn, which is critical to marine life,
as it brings nutrients which have accumulated in deep water
back up to the surface. This also is extremely important as
a seasonal process in temperate regions, where cold tem-
peratures and stormy weather during the winter can combine
to produce overturn and mixing, replenishing nutrients for
use by surface organisms during the spring. Upwelling also
occurs when water rises to replace surface water driven off-
shore by winds, a process called coastal upwelling (C&H Fig.
15.30, p. 354). Downwelling also occurs when water is driven
against the shore, forcing surface water down. The eastern
sides of the Atlantic and Pacific Oceans, where winds and
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surface currents tend to move surface water away from the
coast, are important areas of coastal upwelling (C&H Fig.
15.31, p. 355). Similarly, upwelling occurs when water rises
to replace surface water drawn apart by two diverging sur-
face currents, and downwelling occurs where two converging
surface currents pile up surface waters, inducing sinking. The
former process occurs primarily at the equator, where it is
called equatorial upwelling (C&H Figs. 15.33, p. 357),
and the latter often occurs at about 30N and 30S latitudes.
Density controls not only the vertical structure of the ocean,
but also the circulation of the vast majority of its volume.
The types of water movement we have discussed so far,
surface currents and the vertical flow of water involved in
upwelling and downwelling, are important but they are not
the dominant form of circulation in the ocean. Using their
temperature-salinity "signatures," scientists have tracked
deep- and bottom-water masses and discovered that these
water masses (80% of the ocean's volume) are flowing, very
slowly, in immense subsurface currents that circulate wa-
ter throughout the world ocean. Because this deep-ocean
circulation is driven by density differences produced by tem-
perature and salinity changes at the surface, it is called ther-
mohaline circulation. It begins in the polar seas south of
Greenland and around Antarctica, in relatively shallow basins.
Here, surface water becomes very cold and very salty, and
therefore very dense, so it sinks (we will see how this hap-
pens when we discuss sea ice, below). This sinking water
becomes trapped at the edge of the shallow basin, which
forms a sill and acts as a dam at the entryway to a deeper
basin. The dense water piles up until it eventually overflows
the sill and streams out and down into the deeper basin. In
this way, Antarctic Bottom Water is formed and streams
north, past the equator. Similarly, North Atlantic Deep
Water streams south, all the way to the Southern Ocean
(C&H Fig. 3.24). This deep circulation is connected with shal-
lower circulation, returning warm surface-water from the
Pacific and Indian Oceans to the North Atlantic Ocean, thus
completing a circuit (C&H Fig. 3.25). Like an immense con-
veyor belt, this circuit is called the great ocean conveyor.
Requiring perhaps thousands of years for the completion of
one cycle, it mixes the ocean on a global scale. The volume
of water involved is calculated to equal 100 Amazon Rivers,
or all the rain falling on the planet. The great ocean con-
veyor is vitally important to marine life, as it carries oxygen to
deep waters and returns nutrients to the surface. It also has
immense influence on global weather patterns and climate,
as it transports heat from the tropics toward the poles and
vice versa. The possibility of disturbances to the thermoha-
line circulation of the ocean, as a result of the increasing
greenhouse effect and global warming, is of great concern.
Pressure, Light and Sound
Depth in the water column also affects several other impor-
tant physical properties of seawater. Pressure, exerted
by the weight of the column pressing down from above, in-
creases steadily with depth. We hardly notice the weight of
the enormous column of air above us, exerting 1 atm (atmo-
sphere = 1 kg per square cm, or 14.7 lb per square in) of
pressure at sea level. Marine organisms are subject to this
1 atm of pressure, plus the pressure of the column of water
above them. The pressure exerted by water adds up very
quickly, about 1 atm for each 10 m (33 ft) of depth, since
water is much heavier than air (the pressure profile in Fig.
6 shows this linear relationship). The pressure in the Chal-
lenger Deep of the Mariana Trench, at 11 km below sea level,
would therefore be about 1100 atm, or 16,170 lb per square
inch! Water is nearly incompressible, but gases are highly
compressible. A plane flying at 10 km with a cabin pressure of
1 atm must be engineered to prevent it from exploding, while
a submarine with the same cabin pressure at depth must be
built so as not to implode. Similarly, marine organisms with
gas-filled structures like lungs and air bladders must con-
tend with extreme pressure from without, when at depth, and
extreme pressure from within, when rising in the water col-
umn. Sperm whales, which can dive down more than 2000 m
and back up to the surface within the space of an hour, may
be subjected to the most extreme pressure changes of all.
How seawater transmits light, and how this varies with
depth, also is vitally important. Sunlight provides the energy
for photosynthesis, and photosynthesis directly or indirectly
supports virtually all life. Light is readily transmitted through
the atmosphere, but the ocean reflects much of the light
striking its surface, and quickly absorbs the light that does
penetrate, converting it to heat (Fig. 7). A light profile
(C&H Fig. 3.11) shows that about 65% of visible light is ab-
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sorbed within 1 m of the surface in clear water, and less than
1% penetrates to 100 m. As a result, photosynthesis takes
place only in the upper reaches of the ocean, and organism
densities generally are highest near the surface. The quality
of light also changes with depth. Longer wavelengths, at the
red end of the spectrum, are absorbed more readily than
blue light. This selective absorption accounts for the blue
and green colors of the sea, and the blueish or greyish ap-
pearance of objects at depth, in natural light (C&H Fig. 3.12).
Unlike light, sound travels faster and farther in water (about
1500 m/s) than in air (about 340 m/s). The sea is alive with
sounds produced by marine creatures and, increasingly, with
human-generated "noise pollution." Marine organisms use
sound for communication and echolocation (nature's sonar),
and humans employ sound waves to explore the ocean, as we
learned in the previous lesson. The speed of sound increases
with increasing salinity, temperature and pressure. Accord-
ingly, sound velocity profiles (Fig. 8) show an initial de-
crease in velocity with depth in the water column, as temper-
ature decreases in the region of the permanent thermocline,
then an increase at greater depths, as pressure continues to
increase. This produces a region of minimum sound velocity,
sandwiched between layers of greater velocity, much like the
oxygen minimum zone. Sound waves traveling in this layer
of minimum sound velocity are "trapped" here because they
are refracted (bent) from above and below, and are directed
toward the center of the layer. Thus focused, sound waves
can travel great distances without losing much energy in
this layer, called the SOFAR channel (for SOund Fixing And
Ranging), as if echoing through a tunnel. Scientists place re-
cording equipment at the center of the SOFAR channel (600
1200 m down, at low and middle latitudes), to monitor whale
vocalizations, earthquakes, and man-made sounds. Sound
transmissions across entire ocean basins, as far as 25,000
km (about 15,500 mi), have been recorded in this zone.
Icebergs, Ice Shelves and Ice Islands
Another important feature of the pelagic environment is the
presence of ice at the surface. Water's ability to coexist on
the planet in all three phasesgas, liquid and solidis
another of its unique properties. Even more remarkably, wa-
ter becomes less dense as a solid than as a liquid, so ice
floats. This is crucial for marine and aquatic life, since ice
forms at the surface, insulating and maintaining the liquid
Figure 6. Water pressure increases steadily with depth. Figure 7. Light is quickly absorbed as it penetrates water.
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environment below, and it melts quickly when temperatures
rise again. Ice is found in marine environments in two gen-
eral forms. Freshwater ice originates on land in glaciers and
ends up in the ocean as icebergs, ice shelves and ice islands.
The second form, sea ice, occurs when seawater freezes.
Icebergs, or just "bergs," calve (break off) from glaciers
which flow to the sea, primarily the huge ice sheets cover-
ing Antarctica and Greenland. They come in many shapes
and sizes, ranging from "growlers" (less than 1 m above the
sea surface by 5 m across the sea surface) and "bergy bits"
(14 m by 514 m) to "very large bergs" (over 75 m by over
213 m). The tallest known iceberg in the North Atlantic tow-
ered 168 m above the sea, the height of a 55-story building.
Since 6080% of a berg's bulk is submerged, however, this
is literally just the tip of the iceberg! Icebergs can weigh hun-
dreds of thousands of tons, and are dangerously unstable,
tending to break or capsize. Subsurface "rams" projecting
far beyond the visible part of the berg are another great
danger to vessels. Large bergs may be surrounded by fog
banks, due to the difference in temperature between the ice
and the surrounding water. They may run aground, and can
scour and gouge the sea floor, preventing the installation
of undersea cables and pipelines in some areas. Afloat in
open water, icebergs are transported primarily by currents,
due to their large underwater extension and relatively small
"sail" area. They become locked into the sea ice during win-
ter, and are released again in spring. Since they continually
melt and erode away, only a relatively few, large icebergs
last long enough to be carried into shipping lanes, which
may take 23 years. It was one of these persistent bergs
that sank the Titanic in 1912, off the coast of Newfound-
land. Ever since that disaster, the North Atlantic has been
patrolled to locate icebergs. Today, icebergs are monitored
worldwide, using remote sensors on polar orbiting satellites.
Ice shelves are found attached to 44% of the Antarctic
coastline, and the coasts of Greenland and northern Elles-
mere Island, Canada. They are created by the seaward ex-
pansion of glaciers, augmented by annual snow accumula-
tion, until a tongue of ice extends out over the sea surface.
Their ice covers immense areas, is 1001000 m thick, and
thousands of years old. The largest is the Ross Ice Shelf, at
472,000 km
2
(182,000 mi
2
), about the size of France. It was
discovered in 1841, during the Erebus and Terror Ant-
arctic expedition, and later named in honor of its commander.
(Ross' response on seeing it was, "there's no more chance
of sailing through that than through the Cliffs of Dover. ") Ice
Figure 8. The SOFAR channel.
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shelves also calve icebergs. One of the largest icebergs ever
recorded, Iceberg B15, broke off the Ross Ice Shelf in March
of 2000. It was 11,000 km
2,
about the size of Connecticut,
before it began breaking apart within months of calving.
The pieces of this huge berg formed dams that blocked the
normal flow of sea ice out of the Ross Sea (also discov-
ered in 1841 by the Ross expedition), upsetting the eco-
system and resulting in a loss of productivity in this region.
Huge, flat icebergs that calve from ice shelves are called ice
islands. Some have even been inhabited. T-3, an ice island
calved off of Ellesmere Island, carried a manned research
station complete with power plant and runway, from 1952-
1978. The entire Ayles Ice Shelf fractured off of Ellesmere
Island on August 13, 2005, becoming an ice island the size
of Manhatten. Caught by satellite imagery, this amazing event
took less than an hour. The Markham Ice Shelf, another huge
one, also calved entirely off of Ellesmere Island in 2008. In
all, Ellesmere Island ice shelves were reduced by 90% during
the twentieth century, the most dramatic example of a gener-
al decrease in ice shelf extent. The disintegration of shelves,
a phenomenon different from normal calving events, also is
becoming more common. And ice shelves are become thin-
ner, as they are being melted from beneath (basal melting),
due to the gradual warming of Arctic and Antarctic waters,
and to changing winds and currents bringing warmer water
to the poles. All of these changes are linked to the accelerat-
ing greenhouse effect, global warming and climate change.
Thinning and disintegration of ice shelves may have global
consequences, in addition to disrupting marine life locally,
and hazarding ships and oil platforms. Ice shelves support
unique communities of organisms. They regulate the tem-
perature and salinity of the water beneath them, thus in-
fluencing thermohaline circulation. They also contribute
to the high albedo of polar areas. Decreasing ice cover-
age results in less light being reflected, and more heat be-
ing absorbed, by the darker exposed water. This produces
a positive feedback effect, accelerating warming at the
poles. As warming surface waters expand, and meltwater
from land glaciers runs to the sea, sea level also is in-
creasing. Melting of icebergs and ice shelves does not di-
rectly raise sea level, because floating ice already displaces
a volume of water equal to the volume of ice above the
water. However, it can indirectly increase sea level, since
ice shelves act like dams on glaciers. When they break off,
the glacial ice that was held back will move faster toward
the ocean, another case of a feedback loop that will ac-
celerate changes due to the runaway greenhouse effect.
Sea Ice
To the untrained eye, sea ice may appear indistinguish-
able from glacial ice, but it actually is very different. Sea ice
forms from seawater, and is far more complex and extensive,
accounting for 95% of ice found in the sea, and far more
important both locally and globally. When fresh water cools,
its density increases to the point of maximum density, at
4C, and then density decreases and freezing occurs at 0C.
In this manner, a thin, glass-like sheet of ice will form on the
surface of a calm lake. The presence of salt in water lowers
the temperature both of maximum density and of freezing
(the freezing point). In seawater of average salinity, the
temperature of maximum density and the freezing point co-
incide at about 2C, so seawater continues to get denser
as it cools, until it freezes at 2C. As a result, water at the
surface sinks as it cools, with ice crystals floating in a com-
paratively thick layer of congealing slush, which gives the sea
surface a cloudy or oily appearance ("grease ice" is one form
that occurs during this process). Further freezing creates a
flexible crust of new ice 510 cm thick, called "nilas," which
bends in response to waves and wind, and is able to support
the weight of a gull but not of a man, unlike freshwater ice
of similar thickness. This crust thickens by the addition of ice
to the bottom (opposite to how glacial ice accumulates), and
becomes more brittle. Wind and wave action break and shape
it into circular pieces, called "pancake" or "pan" ice, which
unite to form larger ice floes and eventually massive ice fields
as freezing continues. In this way, the seasonal ice pack (or
winter ice pack) is formed, consisting of annual ice that
melts each spring and forms again each winter. At higher lati-
tudes, the permanent polar ice pack occurs, consisting
of multiyear ice, which has survived at least two summers'
melt. The summer ice pack also remains but is broken up
to some extent each summer. The ice pack includes landfast
ice, or fast ice, which is attached to the shore, and drift ice.
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New ice attains a thickness of 30 cm to 23 m during its
first winter. Even multiyear ice is usually less than 10 m
thick, much thinner than glaciers and ice shelves, because
it undergoes a continual cycle of loss and replacement. The
surface layer melts each summer, while new ice is added
to the bottom during each winter. Thus, the youngest lay-
ers are at the bottom, in contrast to glaciers, and they mi-
grate upward over time. The migration of each new layer
to the top may take about 12 years, also in contrast to ice
present in glaciers, which may be thousands of years old.
As seawater freezes, not all the salt can be incorporated
into the ice, and extremely salty liquidbrineis expelled
into the water beneath the forming ice. Called brine rejec-
tion, this process results in fresh ice near the top of the
pack and increasingly salty ice in the lower layers. As the
cold water below the forming ice gets saltier it also gets
denser and begins to sink, becoming a major contributor to
the deep-water masses near the poles that drive ther-
mohaline circulation and the great ocean conveyor.
Sea ice is important to regional and global heat balance
in other ways, as well. The ice pack inhibits heat exchange
between the sea and the air. It insulates the sea from the
cold air, like a blanket, and actually makes the air colder by
blocking access to the water's huge heat stores. Because
of its high albedo, especially when covered with snow,
sea ice also reflects heat away from the Earth's surface.
Sea ice is extremely complex and variable, subject to alter-
nate growth and melting, and to movement and deformation
by wind, tides and currents. Ice floes are carried thousands
of kilometers away from the ice pack. Leads, long cracks in
the ice wide enough to allow passage of a ship or a whale,
open and close back up again. Other areas of open water,
called polynyas, occur over upwellings of relatively warm
water, or on the leeward side of islands and coasts, where
prevailing winds keep blowing the ice away from shore. Col-
liding ice floes push up pressure ridges, piles of jumbled
blocks of ice 10 m or more high, with "keels" that may ex-
tend even farther downward from the bottom of the pack.
These keels can gouge and plow the sea floor in shallow
areas. Beaches also are shaped by the actions of sea ice.
One of the most impressive sea-ice phenomena, referred
to as push ice or "ivu" by Inupiat Eskimos, occurs when
blocks of ice are driven by wind and currents, with such
speed and energy that they violently override the shore or
landfast ice. A group of hunters on the fast ice off Barrow,
Alaska, had to abandon their dog teams and gear and run
for their lives, in the path of an ivu in 1957. Hundreds of
years earlier, a family was crushed in their home by an ivu,
their bodies remaining frozen in the Arctic soil until a re-
cent storm unearthed the site, on the coast near Barrow.
This challenging environment is, surprisingly, one of the eco-
logically richest and most important in the world. Sea ice is
home to unique and diverse communities of marine life, on
which many other organisms (including humans) depend, as
we will see later in this course. The ice packits extent,
its movements, and the timing of freeze-up and meltis
critical not only to life and physical conditions regionally, but
also globally, because of its role in ocean circulation and
global heat balance. Alarming changes to the Arctic ice pack
and parts of the Antarctic ice pack, similar to the changes
seen with ice shelves, have been observed in recent de-
cades. Sea ice is thinning, its freezing later and breaking
up earlier, and there has been a dramatic decline in the ex-
tent of ice coverage in the Arctic and parts of the Antarctic.
Multiyear ice in the Arctic shrank an average of 8 percent per
decade, since satellite monitoring began in the late 1970s,
until the beginning of this century, when it began dropping
more precipitously. In September of 2007, a record minimum
extent was seen, 40% less than the previous 28-year aver-
age. (The minimum extent is the smallest area covered by ice
during the summer melt season, March-September.) This re-
cord was broken in September 2012, when the minimum ex-
tent of Arctic sea ice was nearly 50% less than the historical
average. In other words, Arctic sea ice is melting at an accel-
erating rate, faster than most scientific models had predicted,
and likened by some to a "death spiral." Revised models are
predicting that the Arctic may be completely ice-free in sum-
mer by the 2030s, for the first time in about a million years.
We know that the poles are warming faster than the rest of
the planet, but we are still trying to understand why. One
logical reason is that decreasing albedo due to shrinking ice
cover is creating a feedback loop. Sea ice reflects 80% of
the sunlight that hits it, while open water absorbs 90%. Huge
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losses in ice cover in summer (when sunlight is at a maxi-
mum) therefore may contribute both to accelerated warming
and melting at the poles, and to global warming. Recent stud-
ies also suggest that changing weather patterns are trans-
porting energy to the poles, which may be an even more
important factor. Regardless of the mechanisms, the loss of
sea ice will have enormous impacts on conditions and life in
these regions. Moreover, through disturbances to currents
like the Gulf Stream and to thermohaline circulation, it
may also influence weather systems and climate worldwide.
Thus, sea ice conditions serve both as an indicator, and
as a driver, of changing conditions regionally and globally.
Ocean Weather
We have seen that the ocean is far from homogenous, but in-
stead is horizontally and vertically differentiated with respect
to many characteristics, including chemical composition, sa-
linity, temperature, density, pressure and light. We also have
seen that, far from being static, water masses are moving in
this 3-dimensional environment in varied and extremely com-
plex ways. Physical oceanographers who study these move-
ments are coming to view them as examples of "weather" in
the ocean. Water masses are in contact along "ocean fronts"
similar to weather fronts in our atmosphere. An example of an
ocean front is the edge of a mud plume spreading from the
mouth of a river after a flash flood, or the abutment of a sur-
face current like the Gulf Stream against another water mass.
Turbulence is generated along ocean fronts, as it is in the
atmosphere, but operates more slowly and over longer time
periods in the ocean, because of water's greater density. Sea
floor topography also affects the flow of water masses, in the
same ways that air masses are affected by landforms like
mountains. Submarine "waterfalls" are a spectacular example
of this effect. When dense surface water sinks and accumu-
lates behind a barrier in the sea floor, eventually spilling over
onto the other side (as in the formation of Antarctic Bottom
Water and North Atlantic Deep Water, discussed above), the
cataracts produced can dwarf any waterfall found on land.
On its way north in the great ocean conveyor, Antarctic Bot-
tom Water forms numerous submarine waterfalls, e.g. as it
passes over the Rio Grande Rise, off the coast of Brazil.
Pelagic Habitat Zones
Now that we have described the physical and chemical char-
acteristics of seawater, we are ready to fill in our benthic
basin (Fig. 3) with the habitat zones found in the pelagic
environment (Fig. 9). The pelagic realm is divided into two
general regions, the neritic zone, which lies over the con-
tinental shelf, and the oceanic zone, which includes all the
open water beyond the shelf break. These zones are further
divided according to depth. The deepest hadopelagic and
abyssopelagic zones lie over the hadal and abyssal benthic
zones, below 6000 m, and from 40006000 m deep, respec-
tively. Over the bathyal benthic zone, the pelagic environment
is divided into two zones: the bathypelagic, 10004000 m
deep; and the mesopelagic, from about 100 or 200 m, to
1000 m deep. The shallowest zone is the epipelagic, from
the surface down to about 100 or 200 m, including nearly all
the neritic zone plus the surface waters of the open ocean.
The pelagic environment also can be classified according to
light penetration, into photic, dysphotic and aphotic zones.
The photic zone corresponds to the epipelagic, where
sunlight warms the surface waters and supports photosyn-
thesis. Life thrives here, given sufficient nutrients. Below the
photic lies the dysphotic zone, roughly corresponding
to the mesopelagic. This is the "twilight zone" where some
light penetrates but not enough to support photosynthesis,
and organisms are subjected to increasing cold and pres-
sure. Below the dysphotic lies the aphotic zone, includ-
ing the bathypelagic, abyssopelagic and hadopelagic, which
are sometimes elided into a single "deep-sea zone." This
vast realm receives no sunlight at all and is subject to ex-
treme pressure and cold (or extreme heat, around hydro-
thermal vents). We will explore these zones, and the fas-
cinating life forms they support, in Part 3 of this course.
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High Tide Mark
Low Tide Mark
Hadopelagic
Abyssopelagic
Bathypelagic
Mesopelagic
Epipelagic
Oceanic
Neritic
Pelagic
Intertidal
Benthic
Shelf Break
Hadal
Abyssal
Subtidal
4000 m
1000 m
100-200 m
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a
Bathyal
Benthic and Pelagic Habitat Zones
High Tide Mark
Low Tide Mark
Hadopelagic
Abyssopelagic
Bathypelagic
Mesopelagic
Epipelagic
Oceanic
Neritic
Pelagic
Intertidal
Benthic
Shelf Break
Hadal
Abyssal
Subtidal
4000 m
1000 m
100-200 m
D
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Bathyal
Figure 9. Benthic and pelagic habitat zones
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glossary terms
your textbook.
hydrologic cycle
solvent and solute
universal solvent
salt and ion
salinity (psu, ppt)
rule of constant proportions
nutrient
pollutant
calcium carbonate
Niskin bottle
gas exchange
greenhouse gas, greenhouse effect, and
global warming
pH and ocean acidification
heat capacity
insolation and albedo
profile (of temperature, pressure, etc.)
thermocline, permanent thermocline, halo-
cline and pycnocline
stratification
downwelling, upwelling, and overturn
deep water and bottom water masses
thermohaline circulation and great ocean
conveyor
oxygen minimum zone
SOFAR channel
freezing point
iceberg, ice shelf and ice island
winter/seasonal ice pack, summer ice pack
and permanent ice pack
annual ice and multiyear ice
drift ice and fast ice
lead, polynya, pressure ridge and ivu
Coriolis effect and Ekman transport
trade winds, westerlies and polar easterlies
equatorial currents and Gulf Stream current
gyres
fetch
seas, swells and surf
tsunami
diurnal tides and semidiurnal tides
neritic zone and oceanic zone
epipelagic zone, mesopelagic zone, bathype-
lagic zone, abyssopelagic zone and hadope-
lagic zone
photic zone, dysphotic zone and aphotic zone
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1. Complete the crossword puzzle on the following page. All the terms in the puzzle are included in the "Glossary
Terms" for this lesson. (Submit the completed puzzle, or a list of the answers, with this written assignment.)
2. How do the ions found in seawater get there? (Describe 4 ways.)
3. Imagine that you take the bathyscaphe Trieste out of retirement (she's now on display at the National Undersea
Museum in Keyport Washington), for another trip down to the Challenger Deep of the Mariana Trench, in the tropical
waters of the western Pacific. Describe (or diagram) the temperature, pressure, density, salinity and light conditions
that you observe in the water surrounding your submarine as you descend. Take measurements at the surface, and
at 50 m, 500 m, 1000 m, 5000 m and 10,000 m deep.
4. Off the east coast of Greenland, pack ice covers the sea for most of the year, while at about the same latitude and
only a few hundred miles away, Norwegian ports remain ice-free and temperatures are considerably warmer. This is
due in part to relatively warm waters from the Gulf Stream current bathing Norway's coast, while the East Greenland
current brings frigid water from the Arctic Ocean to Greenland. The Gulf Stream begins with air at the equator being
warmed by the sun, rising and being displaced, which creates winds. Taking it from here, continue explaining (step-
by-step) how the Gulf Stream current develops and carries warm water from the equator all the way to Norway.
Include the Coriolis effect, Ekman transport and gyres in your explanation.
5. Most coasts experience semidiurnal tides. Define these tides and explain what causes them to occur.
6. The freezing point of sea ice (-2
o
C) is lower than that of freshwater ice (0
o
C). Describe 5 other important differ-
ences between sea ice and glacial ice.
7. Explain the role of brine rejection, which occurs as sea ice freezes, in the formation of deep-water masses, and how
it helps to drive the great ocean conveyor. Describe some specific changes that have been observed in the Arctic ice
pack in recent decades. Speculate on how these changes could affect the great ocean conveyor, and how altering the
great ocean conveyor could have global consequences. (Try to be very clear, as if explaining to someone who has no
knowledge of the terms you are using, in order to show your understanding of these complex processes.)
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crossword puzzle for Lesson 3
(Tear out and submit for grading)
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37 4
Biological
Principles, Cells,
Organisms &
Ecosystems
Marine organisms are incredibly diverse in form and function, with diverse strategies for
meeting their needs and for coping with the challenges presented by their surroundings.
They also are involved in complex interactions, both with other organisms and with their
physical environments. Underlying all this complexity, however, are basic characteristics,
needs and challenges shared by all organisms, and general principles by which organisms
interact with each other and with their environments. In this lesson, we review these
fundamentals of biology and ecology.
Objectives for this Lesson
1. Know the basic characteristics of living things, the fundamental ways in which they
differ from nonliving matter, and the molecular basis for these attributes.
2. Be familiar with the general structure of ecosystems, and the basic types of interac-
tions among organisms and between organisms and their environment.
Steps for Completing the Lesson
Biology 9th Ed.).
Reading Assignment
C&H: Chapter 4 (pp. 6483) and Chapter 10 (pp. 211229)
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N
Now that we are familiar with the abiotic (nonliv-
ing) features of the ocean, we turn to its biotic
(living) aspect. In the previous lessons, we learned
that the marine environment is extremely varied, dynamic and
challenging. Marine organisms, too, are incredibly diverse in
form and function, with diverse strategies for meeting their
needs and for coping with the challenges presented by their
surroundings. We also learned that the ocean is involved in
complex interactions, not only within the marine environ-
ment, but also with the land and the atmosphere. Marine
organisms, too, are involved in complex interactions, both
with other organisms and with their physical environments;
ecology is the study of these interactions. If all this sounds
complicatedit is! Underlying all this complexity, however,
are basic characteristics, needs and challenges shared by all
organisms, and general principles by which organisms inter-
act with each other and with their environments. In this les-
son, we review these fundamentals of biology and ecology,
in preparation for meeting the ocean's diverse inhabitants,
in Part 2, and its ecological systems, in Part 3 of this course.
Diverse, Complex and Organized
Living things can be broken down into simple, lifeless mol-
eculesthe same molecules found in inanimate matter,
conforming to the same laws of chemistry and physics. So,
what distinguishes these collections of molecules, and what
are the unique attributes of life? At the most basic level, or-
ganisms consist of a unique assortment of chemicals. Of
the 90-plus naturally occurring elements, only about 30 are
essential ingredients of life. Whereas oxygen, silicon, alumi-
num and iron are the most abundant elements in the Earth's
crust, 99% of living matter is made of hydrogen, oxygen,
carbon and nitrogen. These four happen to be the lightest
elements capable of forming strong bonds with each other,
and are supremely adept at combining together in myriad
ways to produce organic compounds. Organic compounds
are distinguished from inorganic compounds by their carbon
"backbone" consisting of two or more carbon atoms bonded
together. Of all the elements, carbon atoms are the most
versatile at bonding with each other and with other atoms,
and are capable of forming by far the greatest variety of
sizes and shapes of molecules, including very large, com-
plex ones called macromolecules. Living things consist
of organic matter, which is composed of water and a vast
assortment of organic molecules. Most of these mole-
cules belong to one of four major classes: carbohydrates,
lipids, and the macromolecular nucleic acids (DNA
and RNA) and proteins. All of life's attributes stem from
the properties of these molecules and their interactions.
The most obvious qualities of life are its extraordinary di-
versity, complexity, and organization. Inanimate matter
has little structural organization, usually consisting of ran-
dom mixtures of simple molecules. In contrast, a microscopic
bacterium contains thousands of different organic molecules,
including several thousand types of proteins. None of these
proteins is identical to any of the approximately 35,000 dif-
ferent proteins found in a human. In fact, each type of living
thing has its own unique complement of proteins and nucleic
acids. Scientists have only begun to identify, and may never
fully know, all the different types of life on Earth, let alone all
the kinds of organic molecules they contain. Yet, the virtually
limitless variety of macromolecules found in living things is
constructed from a very limited variety of smaller, building-
block molecules. All proteins are formed from just 20 kinds
of amino acid molecules, which are identical in all living
things. These 20 amino acids are linked together in chains of
different sequences, to create the endless variety of different
proteins. Similarly, all DNA and RNA molecules are created
Getting organized: Cells, organisms & ecosystems
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from just 5 nucleotide molecules, repeating in chains of
various sequences. Large carbohydrate and lipid molecules
also consist of chains of building blocks, usually simple sug-
ars like glucose, and fatty acids, respectively. In all, there
are around 100-200 kinds of building block molecules and
other relatively small, organic compounds that can be found
in virtually all cells. These universal organic compounds are
capable of producing the limitless variety of more complex
molecules found in organic matter. We can see, therefore,
that the molecular diversity and complexity of living things
is based on an underlying simplicity and economy. Further,
since all forms of life use the same building-block molecules,
we can infer that they share a common origin. Finally, since
all individuals of a certain type of organism, or species,
share similar sets of proteins and nucleic acids, distinct from
those of other species, we can surmise that these character-
istic sets of macromolecules are the basis for the identity of
the different forms of life, and for its astonishing diversity.
The diverse, complex molecules of organic matter are assem-
bled into even more complex, highly organized structures in
living things. This complexity and orderliness, again, is based
on an underlying simplicity and economy. Macromolecules
are combined into supramolecular complexes, such as
membranes and ribosomes. Supramolecular complexes,
in turn, may be organized into more intricate, membane-
bound structures called organelles. Organelles and other
structures are organized within a watery matrixthe cyto-
plasmwhich is surrounded by a cell membrane to form
a cell, the basic unit of life. The vast majority of living things
are unicellular, consisting of a single cell. Most are pro-
karyotes, such as bacteria, whose cellular components are
usually not organized into organelles (C&H Fig. 4.7). More
complex unicellular, and all multicellular, forms of life con-
sist of more complex eukaryotic cells (C&H Fig. 4.8). Mul-
ticellular forms contain different types of cells. Similar cells
group together into tissues, various tissues assemble into
organs, and organs are combined into organ systems,
which together make up the individual organism (Fig. 10).
Every structural component of an organism also has a specific
function. Biomolecules serve basically the same functions in
all cells. Carbohydrates and lipids store fuel and form struc-
tural frameworks. The nucleic acids, DNA and RNA, store and
transmit the instructions for the construction and functioning
of an organism, and proteins are the instruments for carry-
ing out these instructions. Each cellular component also has
a specific role to play. For example, the mitochondria are
the cell's power plants, and ribosomes manufacture proteins.
Each type of cell and tissue (muscle or nerve) serves a spe-
cific purpose, as does each organ (heart or kidney), organ
system (respiratory or digestive) and gross structural com-
ponent (a branch, a wing, or an arm) within the organism.
Self-Assembling
Complex and highly organized, both structurally and
functionally, living things are further distinguished
by their capacity to assemble themselves. This
unique ability also originates in the unique proper-
ties of biomolecules, and the ways they interact.
Nucleic acids and proteins are informational macromol-
ecules. The sequences of nucleotides in the DNA molecules
of a cell encode the instructions for constructing all the oth-
er cellular components. DNA molecules serve as templates
for the transcription of their nucleotide sequences into
complementary nucleotide sequences in strands of RNA.
Some of these RNA molecules, called messenger RNAs or
mRNAs, in turn serve as templates for the construction of
proteins, during translation. Translation takes place on the
ribosomes, where amino acids are linked together in spe-
cific sequences, corresponding to the code of nucleotide
sequences in the mRNA strands being translated. Amino
acids are carried to the ribosomes by other RNA molecules,
called transfer RNAs or tRNAs, different types of tRNA for
each type of amino acid. In this way, the instructions encoded
in DNA molecules are used to construct protein molecules.
Once assembled, protein molecules also become informa-
tion-rich. The particular amino acid sequence of a protein
determines its specific 3-dimensional structure, which in
turn determines its specific biological activity. In the watery
environment of the cell, the long strand of amino acids pro-
duced by the translation of mRNA, called a polypeptide
chain, is unstable in its extended form. Each polypeptide
spontaneously curls and folds into a specific 3-dimensional
shape, or conformation, which is determined by its par-
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41
ticular amino acid sequence. The specific conformation of a
protein, and the specific positioning of its functional groups,
allows it to recognize and interact with other molecules in
very specific ways, like pieces fitting together in a 3-dimen-
sional jigsaw puzzle. Through these interactions, the instruc-
tions encoded in DNA are carried out. The interactions of
biomolecules, according to the complementary fit of
their 3-dimensional structures and the positions of
their functional groups, is a fundamental principle of life. It
applies to the transcription of DNA into RNA molecules, the
translation of mRNA into proteins, and the specific interac-
tions of proteins and other molecules, which ultimately ac-
count for all aspects of structure and function in living things.
Proteins initiate the assembly of cell structures by recogniz-
ing and spontaneously associating with other molecules, to
form supramolecular complexes. Ultimately, organelles, cells
and organisms assemble in essentially the same way, accord-
ing to the specific interactions of proteins with other mole-
cules. Through these interactions, proteins are able to carry
out the instructions contained in DNA because they are key
components in all cell structures, and key participants in all
cell functions. Fibrous proteins help to shape, support and
protect cells and organisms. They include the keratin proteins
of skin, scales, hair, nails and feathers, the collagen proteins
of bones, cartilage, and tendons, and cytoskeleton proteins,
which give structure and organization to the cytoplasm of
many cells. Most proteins are globular, folded into a great
variety of conformations, which enable them to recognize
and bind to many different molecules and mediate many dif-
ferent biological functions. Motor proteins, for example, pro-
duce movement in many structures including cilia, flagella and
muscles. Other globular proteins transport substances (e.g.,
hemoglobin, the oxygen-carrier in blood), defend against
foreign substances (antibodies), or serve as messengers
(hormones such as insulin and growth hormone), among
many other functions. The most abundant and diverse group
of globular proteins, the enzymes, function as catalysts.
Catalysts are substances that speed up chemical reactions
without being consumed in the process. All the chemical re-
actions of a cell are catalyzed by enzymes, a different one for
each reaction, which requires the presence of thousands of
different enzymes in each cell. Each type of enzyme cataly-
ses the conversion of one specific substrate molecule into
one specific product molecule, again according to the prin-
ciple of structural complementarity: Enzyme and substrate
interact with an exclusive, lock-and-key fit. The thousands of
different enzyme-catalyzed reactions in a cell are organized
into series of reactions called pathways. Together, these
interrelated pathways constitute the cell's metabolism,
another fundamental characteristic of living things. Central
pathways acting on the main constituents of cellsnucleic
acids, proteins, carbohydrates and lipidsare essentially
the same in all living things, another indication of their com-
mon origin. Enzymes are far more specific and efficient than
man-made catalysts. The great specificity, efficiency and
diversity of enzymes allows cells to simultaneously con-
struct thousands of different molecules from relatively few
precursor molecules. Enzyme-catalyzed pathways exist for
the synthesis of precursors into all the different building
blocks, and for the synthesis of building blocks into all the
more complex biomolecules, including DNA itself. Thus, by
encoding the structure of proteins, DNA ultimately encodes
the structure of all the components of cells and organisms.
Chemical Engines
We have seen that living things are complex and highly or-
ganized. They synthesize their own components, assemble
themselves and perform all other biological functions, ac-
cording to the instructions encoded in their DNA. Synthesiz-
ing biomolecules, and assembling and maintaining cells and
organisms, are forms of work. Work requires the input of
energy. There are many forms of energy: radiant, electri-
cal, mechanical and chemical energy, which is the energy
stored in the bonds holding atoms together in molecules.
Energy can be converted from one form into another, and
it can be transferred by doing work or by heat flow (e.g.,
when a pot of water on the stove heats up, or your cup of
coffee cools down). All processes, biotic or abiotic, are the
result of conversions or transfers of energy, and they al-
ways proceed according to the laws of thermodynamics.
The first law of thermodynamics states that energy cannot
be created or destroyed, only transferred or converted from
one form to another, so the total amount of energy in the
universe always stays the same. The second law of thermo-
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dynamics states that all processes proceed with an increase
in the randomness or disorder (entropy) of the universe.
Living things have the unique ability to extract energy from
the environment, convert it into a useable form, and channel
it into biological workall in accordance with the first law.
They also obey the second law, by increasing the entropy
of their surroundings (by breaking down complex molecules
into simpler ones and by giving off heat), while they are cre-
ating and maintaining order within themselves. Inanimate
matter, on the other hand, cannot use external energy to
maintain its organization and usually becomes more disor-
dered when it absorbs light or heat from the environment.
Living things are chemical engines. Their ultimate source of
energy is the radiant energy of sunlight, which is converted
into the chemical energy stored in organic molecules. This
chemical energy is released by the breakdown of complex
molecules into simpler ones during metabolism, and can be
used to perform work. Anabolic metabolism synthesizes
simpler molecules into more complex ones, and catabolic
metabolism breaks down complex molecules into simpler
components. Anabolism is an energy-requiring (endergonic)
process, while catabolism is an energy-yielding (exergonic)
process (C&H Fig. 4.1). In living cells, energy is transferred
from exergonic reactions to endergonic reactions, through
a shared intermediateusually ATP (adenosine triphos-
phate). Some of the energy released during an exergonic
reaction goes into synthesizing ATP, from ADP (adenosine di-
phosphate) and phosphate. ATP can then be used to drive an
endergonic reaction, by breaking it back down into ADP and
phosphate and capturing the released energy to do work.
ATP is the common carrier of energy, used in synthesis and
all other types of biological work, in all cells and organisms.
Catabolic pathways converge into a common pathway, called
respiration, which breaks down glucose to supply most of
the ATP for biological work. Anaerobic respiration (or
fermentation), breaks down glucose incompletely, in the
absence of oxygen. This less efficient type of respiration
was probably used by ancient forms of life, at a time when
Earth's atmosphere lacked molecular oxygen. Some organ-
isms still obtain their energy from fermentation, for example,
primitive fishes called coelacanths, living deep in the ocean
where the oxygen concentration is near zero. More efficient
aerobic respiration, in which glucose is completely bro-
ken down into carbon dioxide and water, is used by most
cells and organisms. Most higher organisms, however, still
retain the capacity for anaerobic respiration, and can use
it to supply energy for short periods when there is insuf-
ficient oxygen for aerobic respiration. For example, normally
sluggish crocodiles use anaerobic respiration during bursts
of activity, as do whales and seals when performing long
dives. Respiration is summarized in the following equation:
C
6
H
12
O
6
(glucose) + 6O
2
6CO
2
+ 6H
2
O
i
chemical energy
Glucose is central to the metabolism of virtually all living
things. It is rich in energy and therefore a good fuel and it
can be converted into a huge variety of other molecules. In
many bacteria, it provides the carbon skeleton for every oth-
er organic molecule needed by the cell. It also can be ware-
housed in the form of larger carbohydrate molecules, like
starch (in plants) and glycogen (in animals), and saved
for future use. Although almost all organisms use glucose
as a fuel and as raw material for building other molecules,
only some organisms are capable of synthesizing organic
molecules like glucose from inorganic ingredients. Only pho-
tosynthetic organisms are able to harness the radiant en-
ergy of sunlight and transfer it into the work of synthesizing
glucose from carbon dioxide and water. In the first phase
of photosynthesis, called the light phase, pigment mol-
ecules absorb light energy, some of which is converted into
the chemical energy of ATP. In the second phase, the dark
phase, ATP's energy is transferred into the synthesis of glu-
cose and other organic compounds. The overall process of
photosynthesis can be summarized in the following equation
light energy
i
6CO
2
+ 6H
2
O C
6
H
12
O
6
(glucose) + 6O
2
All other organisms depend on organic matter from pho-
tosynthesizers, directly or indirectly, to provide raw mate-
rials for their growth and to fuel their processes, through
respiration. Photosynthetic organisms also use respiration
to derive energy from stored glucose, during dark pe-
riods when ATP is not being generated by the light phase
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of photosynthesis. In the case of humans, photosynthetic
organisms are the ultimate source of energy not only for
our bodies, but also for our machines, since coal, petro-
leum and natural gas were all formed from the accumula-
tion of countless dead organisms over millions of years.
As the equations for photosynthesis and respiration show,
these are essentially opposite processes (C&H Fig. 4.5). In
photosynthesis, sunlight, carbon dioxide and water are ab-
sorbed from the environment and used to create organic
matter, glucose. In respiration, glucose is broken down into
carbon dioxide and water, and energy is released. These are
also reciprocal processes in the sense that photosynthesis
uses carbon dioxide and generates oxygen as a by-product,
while respiration uses oxygen and generates carbon diox-
ide. Thus, photosynthesizers and other organisms are in-
terdependent. Photosynthesis provides the fuel as well as
the oxygen required for aerobic respiration. Aerobic respira-
tion, in turn, generates carbon dioxide which can be used
again in photosynthesis. Sunlight is the driving force behind
this continuous cycling of oxygen and carbon dioxide among
organisms, and between organisms and their environment.
We will explore these relationships further, when we discuss
some of the basic concepts of ecology, later in this lesson.
The photosynthetic organisms we are most familiar with are
the flowering land plants. However, at least half of the photo-
synthesis on Earth is conducted in the sea, and the vast major-
ity of marine photosynthetic organisms are microscopic, uni-
cellular prokaryotes and eukaryotes. Some marine microbes,
such as some bacteria living at great depth around hydro-
thermal vents, synthesize organic matter in the absence of
light. Instead of solar energy, they use inorganic compounds
such as hydrogen sulfide as an energy source for the pro-
cess of chemosynthesis. We will learn more about photo-
synthetic and chemosynthetic marine organisms in Lesson 6.
Self-Regulating
In its composition, complexity and organization, life exists in
sharp contrast to its surroundings. Instead of existing in isola-
tion, however, living things are absolutely dependent on their
environment. Living cells and organisms continually release
and absorb energy, converting it and transferring it into bio-
logical work. They continually absorb raw materials, synthe-
size and break down molecules, and release wastes. Because
they are constantly exchanging energy and materials with
their surroundings, living cells and organisms are described
as open systems. Although they do not visibly change from
moment to moment, their individual atoms and molecules are
constantly being exchanged. The types of molecules they
contain, and the concentration of each type, however, remain
relatively constant. The concentration of glucose in cells, for
example, will remain nearly constant because the rate of its
intake or synthesis is just balanced by the rate of its break-
down or conversion into another product. This is called a dy-
namic steady-state. When changes occur in their environ-
ment and this steady-state is disturbed, cells and organisms
make adjustments in order to return to a new steady-state.
This ability to maintain a dynamic steady-state is called ho-
meostasis, which is another fundamental attribute of life.
Homeostasis is achieved by regulating metabolism. All of the
cell's interconnected metabolic pathways are precisely regu-
lated into a superbly coordinated and exquisitely respon-
sive system. Each of the thousands of different metabolic
pathways supplies its product at a rate that exactly matches
the cell's demand for that product, continually adjusting for
the changing needs of the cell, and for changing external
circumstances. Through regulation, metabolism achieves
maximum efficiency and economy, and homeostasis is main-
tained. Living things are self-regulating at every level, and
with respect to every aspect of their functioning. The instruc-
tions for regulation are included in DNA and are carried out
primarily by regulatory proteins. A large portion of DNA is
devoted to encoding these proteins, and much of the work
of life goes into self-regulation and maintaining homeostasis.
In order to regulate themselves, cells and organisms must
continually obtain vital information about their surround-
ings, and respond appropriately. The ability to receive and
respond to signals from the environment is another fun-
damental property of living things. Organisms may sample
their surroundings for the availability of light, oxygen or
glucose, or the presence of toxic chemicals, or other or-
ganisms that may pose a threat, compete for food, or be
potential mates. Similarly, cells within a multicellular organ-
ism must exchange information. Cells and organisms must
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respond appropriately, for example moving toward light
or food, or away from a threat, or by coordinating fuel al-
location among different tissues in a complex organism.
The temperature, pH and salinity of the environment are
particularly important signals for cells and organisms. The
most critical components of living thingsthe informational
macromolecules, DNA and proteinsare fragile. Their abil-
ity to function depends on their 3-dimensional structure,
which in turn depends on the temperature, pH and salinity
of their surroundings. When these conditions exceed cer-
tain limits, proteins and DNA undergo denaturation, losing
their 3-dimensional shape and thus their biological activity.
As is true of the other attributes of life we have discussed,
the ability of living things to receive and act on information
stems from processes at the molecular level and is de-
pendent on the specific actions of proteins. All sensation,
including sight, smell, taste and touch, are mediated by
mechanisms at the molecular level. The most complicated
responses, for example a whale locating a mate across hun-
dreds of miles of ocean, or the perfectly synchronized move-
ments of schooling fish, can be traced to the molecular ma-
chinery by which living cells receive and respond to signals.
Self-Replicating
Living things are unique and incredibly complex, self-assem-
bling and self-regulating chemical machines. They continually
extract raw materials from the environment, transform them
through metabolism and convert energy into work, like syn-
thesis, transport and movement. They receive and act on
signals from their internal and external environments, and
maintain homeostasis. The most remarkable characteristic
of living things, however, and a major investment of their en-
ergy and materials, is their ability to grow and reproduce.
All organisms can produce offspring, new individuals like
themselves. Multicellular organisms also grow and develop,
and continually replace worn-out cells and repair damaged
tissues, by creating new cells from pre-existing cells.These
processes, again, are ultimately molecular in nature, in-
structed by DNA and implemented by proteins. A cell repro-
duces by first duplicating its DNA molecules, a process called
replication. It then divides into two daughter cells, each
containing a copy of the parental DNA. The elucidation of the
structure of DNA and how it replicates were among the most
important advances in biology during the twentieth century,
made possible by contributions from different scientific fields
(chemistry, physics, genetics), and by new technologies.
The characteristics, or genetic traits, which identify an or-
ganism as an individual and as a species are encoded in the
nucleotide sequences of its DNA, inherited from the previous
generation. Transmitting this information to the next genera-
tion maintains the continuity of these individual and species
characteristics, and perpetuates life. An organism's unique
set of DNA molecules is called its genome, which is dupli-
cated in each of its cells. Each molecule of DNA is organized
into a supramolecular complex called a chromosome, one
chromosome for the single DNA molecule contained in most
prokaryotic cells, and numerous chromosomes in the cells
of eukaryotes. A gene is a segment of DNA which contains
the information for synthesizing one functional product, ei-
ther a protein or RNA molecule. Gene expression occurs
when that segment of DNA is transcribedand in the case
of proteins, translatedto yield its product. Regulation of
gene expression, through the regulation of transcription
and translation, is critical for the regulation of all processes
in cells and organisms. In multicellular organisms, different
genes are turned on or turned off in different types of cells,
giving them their different structures and functions. Changing
gene expression over time is responsible for the changes that
occur in a cell or organism at different developmental stages.
DNA molecules are by far the largest molecules. A bacte-
rial DNA sequence may be several million nucleotides
long and include several thousand genes. The 46 chromo-
somes in the human genome contain a combined sequence
of about 3 billion nucleotides, comprising about 25,000
genes. The Human Genome Project, begun in 1990, com-
pleted the first human whole genome sequence in 2003,
which is available to the public and already being used by
researchers in biology, medicine and biotechnology. En-
tire DNA sequences have been determined for numer-
ous other organisms, with more being added all the time.
The ability of living things to preserve these immense in-
formational sequences is truly incredible. DNA is auto-
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matically repaired by specific enzymes, and continues to
replicate itself, in generation after generation of cells,
almost without error. Some forms of bacteria have been
reproducing themselves for countless generations with
nearly perfect fidelity, remaining essentially unchanged for
almost 4 billion years! When an unrepaired error does in-
frequently occur, a portion of the nucleotide sequence is
changed, and a genetic mutation is produced. Through
such changes, the structure of DNA also allows for the fi-
nal distinguishing characteristic of lifeits evolution.
Evolving
Species are altered over vast spans of time, as changes
due to the accumulation of mutations in DNA, and to the
recombination of genetic material that occurs during
sexual reproduction, are passed from generation to gen-
eration. Most mutations are harmful, resulting in defective
organisms unable to reproduce themselves. Occasionally,
however, changes in DNA are beneficial. This often occurs
under challenging or changing environmental conditions.
For example, if a previously available fuel or nutrient mol-
ecule became unavailable in the environment, a mutation
which enabled an organism to use a different fuel or nutri-
ent would confer an advantage on that organism. It would
be more likely to survive and reproduce, passing its al-
tered DNA, and the advantage it confers, onto its offspring.
In the mid-1800s, before DNA and the nature of genetic in-
heritance was known, Charles Darwin first hypothesized "the
survival of the fittest under selective pressure" as an ex-
planation for the changing appearance of species over time
and the origin of new species. Since then, an overwhelm-
ing body of evidence has accumulated in support of Dar-
win's hypothesis, demonstrating that natural selection
for advantageous genetic traits results in the evolution-
ary adaptation of species over time. The modern theory
of evolution has become the unifying concept of biology,
much as plate tectonics is the unifying theory of geology. The
evolution of life since its first appearance in the geological
record almost 4 billion years ago, through changes in DNA
which were transmitted to subsequent generations, accounts
for both the immense diversity of living things today, and
the fundamental characteristics shared by all forms of life.
Interacting
We have outlined the unique attributes of living things and
seen that living things continuously interact with their envi-
ronments, extracting and releasing energy and matter, and
responding to a myriad of signals. Organisms interact with
both the abiotic (physical) and biotic (living) aspects of
their environment, or habitat. They interact intraspecifi-
cally, with members of the same species living in that partic-
ular area, that constitute a population, and interspecifi-
cally with other species. All the interacting populations in an
area form a community. The system of interactions within
a biotic community, and between the organisms and their
physical environment, constitute an ecosystem. The full
range of ecological characteristics of an organism or spe-
cies (including its specific habitat, feeding habits, reproduc-
tive strategy, etc.) is called its ecological niche. Ecology
is the study of all these interactions. All these interactions
result in changes, including the physiological adapta-
tion of individual organisms, and the growth, decline and
evolutionary adaptation of populations, and physical
changes in the environment. As a result, communities are
constantly changing. The continual replacement of one com-
munity by another is called ecological succession. Eco-
systems are complex and dynamic, with many interdepen-
dent parts whose interactions continually reshape the whole.
At the most fundamental level, ecosystems can be viewed
in terms of the flow of energy and matter, from organism
to organism and between organisms and the environment.
Energy and materials flow from organism to organism ac-
cording to their feeding, or trophic, relationships. It begins
with photosynthetic and chemosynthetic organisms, which
are autotrophs (self-feeders). All other organisms are
heterotrophs (feeding on others), directly or indirectly de-
pendent on autotrophs for their energy and raw materials.
An autotroph is fed upon by a heterotroph, which may itself
become food for another heterotrophic organism, and so on,
forming links in a food chain (C&H Fig. 10.13). In a com-
munity of many interacting organisms, many food chains are
interwoven into a food web (C&H Figs. 10.14 and 10.15).
Feeding relationships also can be shown 3-dimensionally as
a trophic pyramid, with a large volume of autotrophs as
the base and increasingly fewer heterotrophs at higher tro-
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phic levels (C&H Fig. 10.16). Autotrophs are a community's
producers, and heterotrophs are its consumers. Consum-
ers that feed directly on producers are primary consum-
ers (or herbivores or grazers). Secondary consumers
feed on primary consumers, and so on up the food chain.
Secondary and higher trophic-level consumers are carni-
vores (predators) and those at the top of the chain are
top carnivores (top predators). Organisms with both
herbivorous and carnivorous feeding habits are omnivores.
Decomposers (or saprotrophs) are consumers that feed
on dead organic matter, from organisms at all trophic levels.
The flow of energy through the living system, or biosphere,
is unidirectional and irreversible. Solar energy enters the
system through producers, is converted into chemical energy
and transferred from one link of the food chain to the next.
Only about 2% of the light energy absorbed by photosyn-
thetic organisms is available to primary consumers, and only
about 10% of that is available to secondary consumers, and
so on up the food chain. The other 90% of available energy,
at each successive link in the chain, is dissipated as heat
into the environment. So, in a food chain with five links and
500,000 units of solar energy absorbed by the producers,
only one unit of energy would be available to the organisms
at the end of the chain. Vast numbers, or a huge biomass,
of producers are required to support a decreasing biomass
of consumers at each higher level of the trophic pyramid.
The energy that is lost to the environment, each time it is
converted and passed up the food chain, can never be re-
captured and used by living things. Eventually, with the death
and decomposition of all the organisms in the chain, all the
energy that originally entered it is dissipated into the envi-
ronment. The continual input of new solar energy, therefore,
is necessary for life to continue. Matter, on the other hand,
can be used over and over, passed from organism to organ-
ism, and between organisms and the environment, in organic
and inorganic forms. The recyling of different types of matter
necessary for life - nutrients - occurs in biogeochemical
(nutrient) cycles. We are already familiar with the car-
bon cycle, in which carbon dioxide in the atmosphere or
dissolved in the ocean is absorbed by photosynthetic or-
ganisms, incorporated into organic matter, passed along
the food chain, and eventually returned to the environment
as carbon dioxide (through respiration), where it can again
be used by photosynthetic organisms (C&H Fig. 10.20).
The role of photosynthetic organisms in the carbon cycle is
critical because they are responsible for carbon fixation
the conversion of inorganic carbon (CO
2
) into organic forms
that can be used by all other organisms for energy and raw
materials. The role of decomposers also is critical for the
continuation of all nutrient cycles, and all life, since they
break down dead organic matter, releasing nutrients which
can be used again by producers to make more organic mat-
ter. Marine photosynthetic organisms are responsible for
about half the carbon fixation (and oxygen production) on
Earth. In addition to fixing carbon into organic matter, marine
organisms incorporate a huge amount of carbon into calcium
carbonate shells and skeletons. In contrast to the terrestrial
environment, much of the dead organic matter in the ocean
sinks out of reach of decomposers, ending up in sediments
on the bottom where it is sequestered, an enormous res-
ervoir of unavailable carbon and other nutrients. This down-
ward movement of vast amounts of carbon, fixed by marine
photosynthesizers and incorporated into calcium carbonate
shells and skeletons, is sometimes referred to as the carbon
pump or carbon sink. These processes have been critically
important for moderating the greenhouse effect and global
warming caused by increasing carbon dioxide emissions.
Nitrogen, an essential component of nucleic acids and pro-
teins, is another nutrient commonly occurring in the envi-
ronment in forms that must be converted in order to be
used by producers. Nitrogen-fixing organisms (certain
types of bacteria and other microorganisms) perform this
vital role in the nitrogen cycle (C&H Fig. 10.21). Phos-
phorus, another crucial ingredient of nucleic acids, is gen-
erally available in the ocean as phosphate, which can be
absorbed by producers, incorporated into organic matter
and passed up the food chain in the phosphorus cycle
(C&H Fig.10.22). The other nutrients required by organisms,
such as iron and silica, participate in similar biogeochemical
cycles. When a bottleneck occurs, at any point in the cycling
of any of the essential nutrients, then the availability of that
nutrient may become a limiting resource, restricting the
growth of individuals and populations, and affecting whole
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communities. Other essentials, such as light, warmth or
space, also become limiting resources when in short supply.
Organizational Schemes
We have seen that organisms and ecosystems, while in-
credibly complex, are governed by basic principles and are
highly organized. Let's review some of the ways we have
learned (here and in C&H Chapts. 4 and 10) to represent
this organization and make sense of all this complexity.
Organisms are assembled from atoms which bind to-
gether into molecules, building-block molecules and
macromolecules. Macromolecules assemble into su-
pramolecular complexes, organelles (in eukaryotes)
and cells. Cells (in complex eukaryotes) form tissues,
which build organs, which cooperate in organ sys-
tems, which make up the individual organism (Fig. 10).
Each type of organism can be identified by its genus
and species (Homo sapiens for us), according to the
worldwide system of binomial nomenclature. Simi-
lar genera (plural of genus) are grouped into a fam-
ily, then into progressively larger groups: order, class,
phylum, kingdom and domain (C&H Table 4.3).
Organisms of the same species living in a particu-
lar area constitute a population, and all the popula-
tions living and interacting together in an area form a
community. Communities interacting with their physi-
cal environment form an ecosystem (C&H Table 4.1).
Organisms participate in communities at different trophic
(feeding) levels. There are autotrophs (producers) and
heterotrophs (consumers). We can further order them
into primary producers, primary consumers, second-
ary consumers, etc. (C&H Fig. 10.15). Or we can call them
producers, herbivores (grazers), omnivores, carni-
vores (predators), top carnivores, and decomposers.
We can show these relationships in a food chain, food web
or trophic pyramid (C&H Figs. 10.13, 10.14 and 10.15).
Organisms interact in communities in many ways. They com-
pete with their own kind, intraspecifically, or with other
species, interspecifically. The most intimate species inter-
actions occur in symbiosis: commensalism, parasitism
and mutualism. Mutualism may be facultative or obligate.
All interactions in ecosystems can be reduced to the flow
of energy and cycling of matter. A food web shows
the flow of energy and matter through a community. The
"big picture" of how matter is continually cycled between
organisms and the environment can be shown in nu-
trient cycle diagrams, like those for the carbon cycle
(C&H Fig. 10.20) and nitrogen cycle (C&H Fig. 10.21).
The marine environment contains two divisions, ben-
thic and pelagic. In Lesson 2, we organized the ben-
thic habitat according to depth, into intertidal (litto-
ral), subtidal (sublittoral), bathyal, abyssal and
hadal zones (Fig. 2). In Lesson 3, we divided the pelagic
habitat into neritic and oceanic zones and, according to
depth, into epipelagic, mesopelagic, bathypelagic,
abyssopelagic and hadopelagic zones (Fig. 9). We
also organized the pelagic environment according to il-
lumination, into photic, dysphotic and aphotic zones.
Similarly, we can organize marine organisms according
to habitat and lifestyle. Benthos (bottom-dwellers) live
in benthic habitats, while nekton (active swimmers) and
plankton (drifters and weak swimmers) live in the epipe-
lagic. Later in this course, we will further classify benthos
as epifauna (animals living on top of the sea floor) or in-
fauna (those within the sea floor), among others, and di-
vide plankton into size categories (C&H Fig. 15.2 p. 334).
We can categorize organisms in many other ways. For ex-
ample, they may be osmoregulators or osmoconform-
ers depending on how they control internal salt concentra-
tions. Or ectotherms, endotherms, poikilotherms or
homeotherms, depending on temperature regulation.
They may be distinguished according to reproductive strat-
egy, for example, asexual reproduction, sexual repro-
duction, or hermaphrodism. In the next lesson, we will
learn more about some of these relationships, as we move
on to Part Two, The Players: A Survey of Marine Organisms.
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glossary terms
Listed below and on the following page are key terms for this lesson. Do a self-check for understanding. If you're unclear
about the meaning of any of these terms, re-read the material in this course guide and in your textbook, and check the glos-
sary at the back of your textbook.
abiotic and biotic
organic compound and inorganic compound
building-block molecule and macromolecule
glucose, fatty acid, amino acid, and nucleotide
carbohydrate, lipid, protein, and nucleic acid
(DNA, RNA)
supramolecular complex, membrane, organ-
elle and cell
prokaryote and eukaryote
unicellular and multicellular
tissue, organ, organ system and organism
transcription, translation and replication of
DNA
conformation and denaturation
catalyst, enzyme and substrate
anabolic metabolism and catabolic metabo-
lism
endergonic and exergonic
ATP
homeostasis
aerobic respiration and anaerobic respiration
(fermentation)
photosynthesis and chemosynthesis
sexual reproduction and asexual reproduction
gene, chromosome, genome, gene expression
and genetic trait
mutation, recombination and natural selection
evolutionary adaptation and physiological
adaptation
habitat and ecological niche
population, community and ecosystem
ecological succession
intraspecific and interspecific competition
symbiosis: commensalism, mutualism and
parasitism
symbiont and host
facultative and obligate symbiosis
autotroph (producer) and heterotroph (con-
sumer)
primary consumer (herbivore), secondary
consumer (carnivore/predator), top predator/
carnivore, omnivore and decomposer (sapro-
troph)
food chain, food web and trophic pyramid
biomass and pyramid of biomass
biogeochemical (nutrient) cycles
carbon cyle and nitrogen cycle
carbon fixation and nitrogen fixation
carbon sequestration/pump/sink
greenhouse effect and global warming
limiting resource
benthos, plankton and nekton
osmoregulator and osmoconformer
ectotherm and endotherm
poikilotherm and homeotherm
binomial nomenclature
domain, kingdom, phylum, class, order, fam-
ily, genus and species
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organic and inorganic compounds - organic compounds consist of a "backbone" of 2 or more carbon atoms,
bonded to other atoms such as hydrogen and oxygen. Inoganic compounds lack a carbon "backbone."
a. amino acids and nucleotides
b. conformation and denaturation
c. anabolic and catabolic pathways
d. photosynthesis and respiration
e. prokaryote and eukaryote
f. osmoconformer and osmoregulator
g. osmosis and active transport
h. ectotherm and endotherm
i. intraspecific competition and interspecific competition
j. population and communitiy
k. habitat and ecological niche
l. autotroph and heterotroph
m. benthos and nekton
n. gene and genome
2. The ability to self-assemble is a fundamental attribute of living things. Briefly explain how they do this, and briefly
describe three other unique attributes of living things.
3. Describe three modes or strategies used by organisms to accomplish reproduction.
4. Symbiotic relationships are the most intimate type of relationship between species. Describe the three types of
symbiosis and, from your own experience, give an example of each type.
5. Without photosynthesizers, nitrogen-fixers and decomposers, life on Earth would grind to a halt. Explain the vital role
that each of these groups plays in the biosphere.
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Part two
The Players: A
Survey of Marine
Organisms
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53
Microorganisms
And The Tree of Life
In this lesson, we begin a survey of the forms of marine life, starting with the smallest, but
by no means least significant: microorganisms.
1. Recognize the great abundance, ubiquity and diversity of microorganisms.
2. Understand the importance of microbes to every aspect of life on Earth, and recog-
nize our lack of knowledge of them.
3. Become acquainted with some of the marine microbes, representing the three major
divisions of life.
4. Be familiar with the Tree of Life, and how it is a work in progress.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 5 (pp. 85100) and pp. 349-350 ("The Microbial Loop") and review p. 73
("Evolutionary Perspective"), pp. 79-83 ("Classifying Living Things"), p. 216 ("Eye on
Science: The Census of Marine Life") and pp. 218-219 ("Symbiosis")
5
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Microorganisms and the tree of life
(A Huge Cast of Tiny Characters)
I
In this lesson, we begin a survey of the forms of marine life,
starting with the smallest, but by no means least signifi-
cant: microorganisms. Microorganisms, or microbes,
are not all related. They include all members of Bacteria and
Archaea, two of the three major divisions of life. They also
include single-celled algae and protozoans, the small-
est members of the third division of life, Eukarya. Combining
these diverse organisms into one category, and attempting to
describe them in one lesson, is simply a measure of how little
we know about them. Recent research is showing us, how-
ever, that microorganisms are the most abundant, diverse
and, in many ways, the most important organisms on Earth..
The Tree of Life
Because of the difficulty of studying them, our knowledge
of microorganisms has lagged far behind that of more vis-
ible and accessible life. As a result, the universal Tree of
Life (the branching diagram depicting how living things are
related to each other) has been very lopsided, until quite
recently. The redrawing of the tree of life in recent biology
texts is a prime example of science as an ongoing pro-
cesssubject to change as new information becomes avail-
able, often with the emergence of new technologies. How
we draw the tree of life in the future will continue to evolve.
Several factors make studying and classifying microorgan-
isms so difficult. Taxonomy, the practice of classifying
living things, has traditionally been done by looking at the
complex morphology (structural characteristics) of plants
and animals, and their detailed fossil record. Microbial struc-
tures, even when made visible under the microscope, are
too simple to be very useful for classifying these organisms,
and they don't leave much of a fossil record. The tradi-
tional way to study microorganisms has been to cultivate
(grow) samples taken from nature, isolate all the different
types of microbe that grow out of the sample, and then cul-
tivate each of those types in a "pure culture." The success
of this laborious process depends on providing the right
combination of nutrients in the growth medium, in the right
concentrations, and at the right conditions of temperature,
pressure, light, acidity, etc., for each unique organism to sur-
vive and proliferateabout as likely as winning the lottery!
Only a tiny fraction of microbes found in nature have been
cultivated. In marine environments, even the process of col-
lecting samples is a problem, since microbes can be sparsely
and unevenly distributed in the vast ocean. Plankton nets
that scientists used for more than a century are too coarse
to capture many marine microbes. Contamination of samples
is another problem, since microbes are everywhere, living
in and on every living and nonliving thing on the planet,
especially where there is moisture. At the same time, the
practice of isolating microbes may prevent us from studying
their true nature, since they usually exist in intimate and com-
plex associations, with each other and with other organisms.
Revolutionary new technologies have changed all this. In the
late 1960s, molecular sequencing was developed, which
allows scientists to determine the order of amino acids in
protein molecules, and the order of nucleotides in RNA and
DNA molecules. At the same time, powerful computers be-
came available, which were needed to analyze such huge
databases (remember, an the organism's DNA sequence is
millions to billions of nucleotides in length). These technolo-
gies made it possible to detect and characterize microbes,
without culturing or isolating them. Molecular sequencing
also allows scientists to directly analyze and compare the
genetic information of organisms, so their relationships can
be determined in a much more accurate and meaningful
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way. Applying these techniques to microorganisms has not
only revolutionized microbiology, but has also radically
altered our view of the relationships among all living things.
We identify organisms by their genus and species name,
then group them into families, families into orders, orders
into classes, classes into phyla (or divisions), and phyla
into kingdoms. When Carl Linn (a.k.a. Carolus Linneaus)
introduced this taxonomic system in the mid-1770s, only
two kingdoms were recognized: Plantae and Animalia.
A century later, a third kingdom, Protista, was proposed
for the diverse group of unicellular organisms that did not
seem to fit either the plant or animal category, and Charles
Darwin's The Origin of Species was published. His hy-
pothesis that species evolve over time through the pro-
cess of natural selection eventually became the unify-
ing concept of biology. It directed taxonomy to become a
search for the phylogeny (evolutionary history, or relat-
edness) of species, their place in the natural order of liv-
ing things. Phylogenetics was still based on what could be
observed at the time: morphology and the fossil record.
Another century passed before two more kingdoms were split
out: Monera for the bacteria, and Fungi for the mushrooms,
molds and yeasts. The two basic cell types also were recog-
nized: the smaller, simpler prokaryote cell seen in bacteria,
and the more complex eukaryote cell found in plants, ani-
mals, fungi, and protists. This 5-kingdom phylogenetic
system, in use since the 1950s and still found in many
textbooks, produced a tree of life with five main branches:
Animalsmulticellular eukaryote heterotrophs
Plantsmulticellular eukaryote autotrophs
Protistsmostly unicellular eukaryotes, with diverse
structures and lifestyles
Fungimostly multicellular eukaryote osmotrophs
(heterotrophs that absorb organic material)
Monera/Bacteriaunicellular prokaryote autotrophs
and heterotrophs
With 4 out of 5 branches devoted to eukaryotes, this tree
suggests that most living things, by far, are eukaryotes.
Similar proportions are evident when we look at the total
numbers of species that have been identified. The Glob-
al Biodiversity Assessment, commissioned by the
United Nations Environment Program in 1995, reported:
1,320,000 animals
270,000 plants
80,000 protists
72,000 fungi
4,000 bacteria
More species, identified in the years since, may be added to
these totals but the proportions remain roughly the same. Do
these proportions reflect the actual proportions of living things
in existence? Do these five groups all deserve to be ranked
at the highest taxonomic level, and do the organisms within
each group share a common ancestry? Have we found most
of the forms of life on Earth? Discoveries in recent decades
have provided some surprising answers to these questions.
In the 1970s, Carl Woese and coworkers used molecular se-
quencing to compare RNA nucleotide sequences in bacteria,
and made an astonishing discovery. Several species were as
different from the other bacteria as they were from eukary-
otes. These unique organisms, methane producers found in
swamp muck and animal intestines, were named Archaea.
As more of these organisms were discovered and analyzed,
it became clear that this previously unrecognized group
is so unique that it requires its own branch on the tree of
life. In 1990, Woese proposed an additional taxonomic rank
above Kingdom, to distinguish the three fundamentally dif-
ferent Domains of life: Bacteria, Archaea, and Eukarya.
This 3-domain phylogenetic system (Fig. 11 and C&H
Fig. 4.24) has become widely adopted. The number of king-
doms within each domain is still being determined. At least
three major groups of archaea, and several dozen major
groups of bacteria, have been identified. Within Doman Eu-
karya, animals, plants and fungi remain as kingdoms, but it
is still unclear how to classify the diverse unicellular eukary-
otes, the protists. A tree of life with three main branches,
two of which are occupied by prokaryotes, represents a
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radically different view of life on Earth and relationships
among living things. It makes sense, however, that pro-
karyotes should make up a significant portion of life. After
all, they arose 3.8 billion years ago, about 2 billion years
earlier than the unicellular eukaryotes and about 3 billion
years earlier than multicellular plants and animals! They
have been evolving and diversifiying all that time, for most
of Earths existence. Prokaryotes also replicate at a much
faster rate, and exchange genetic material more freely, so
they evolve new forms more rapidly than do eukaryotes.
As to the question of how many species there are, and wheth-
er we have found most of them, the answer is: We don't know.
New species are being discovered all the time, much faster
than they can be described or catalogued. Global biodiversity
estimated by taxonomic experts ranges wildly from 3 million
to 100 million species. One recent study estimates global
biodiversity to be 8.7 million species, 2.2 million of which are
marine. If this is accurate, then we have yet to discover 86%
of the total species on Earth, and 91% of marine species.
At the same time, we know that species are disappearing
at an alarming rate, and global biodiversity is decreasing.
It appears that human activities are causing a mass ex-
tinction of proportions unprecedented in the history of life
(C&H p. 215). If known species are disappearing, then how
many more are being lost before they can be discovered?
This all suggests that microbial life in the sea is pro-
lific and diverse, but mostly unknown to us, and that
many unique microbes may be lost before we can dis-
cover them. What little we do know bears this out.
Abundant, Ubiquitous and Diverse
Microbial Abundance
Microbes, we are discovering, are everywhere, in great
abundance. They fill the oceans and freshwater bodies, and
saturate every substrate on the planet. They can even be
found airborne, floating at altitudes as high as 50 miles. The
abundance of microbes in the ocean is astounding. The up-
per 200 m of the ocean holds an estimated 5x10
5
/ml, or
500,000 prokaryote cells in every thimbleful of water. Below
200 m, the concentration of cells drops to about 5x10
4
/ml.
When the immense volume of the ocean is taken into account,
this adds up to 10
29
(1 followed by 29 zeroes) prokaryote
cells in the pelagic environment, 99.8% of the total pro-
karyote abundance in both saltwater and freshwater bodies.
Figure 11. A simplified version of the Tree of Life, following the 3-domain system.
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Many of these marine microbes are so small that they es-
caped detection until the development of the epifluores-
cence microscope in the late 1970s. When treated with
fluorescent dyes, microbial cells glow under ultraviolet light
and can be easily seen and counted with this instrument.
In 1979, a new type of microorganism was reported using
epifluorescence microscopy, a genus of miniature cyano-
bacteria called Synechococcus. These picoplankton
(the size category for the smallest living plankton) were
so numerous and widely distributed in the ocean that they
appeared to be the most abundant organisms on Earth,
until a related genus, Prochlorococcus, was discovered
in 1988, that was even more abundant. Finding new types
of bacteria and archaea, in great abundance in pelagic wa-
ters, is now a common occurrence (C&H p.92, Tiny Cells,
Big Surprises). It appears that prokaryote members of
the plankton far outnumber eukaryotes, the microscopic
algae and protozoans previously thought to dominate the
plankton. Together, bacteria, archaea and microbial eukary-
otes account for more than 90% of the oceans biomass.
Microbes are even more abundant on the ocean floor than in
the waters above. Shallow marine sediments and soils con-
tain about 1000 times more prokaryote cells than in water.
Since the sea floor accounts for 70% of the Earths surface,
this represents almost unimaginably large numbers of mi-
croorganisms in marine sediments. The greatest numbers
of microbes, however, may be not in the water or on the
ocean floor, but deep beneath it. The existence of subsurface
microorganisms has been known for many years, from for-
mation fluids of oil, gas and water wells. Recent studies have
shown that prokaryotes are ubiquitous and often abundant,
in groundwater, aquifers, petroleum deposits and sediments.
Sediments underlying most of the ocean floor and the coastal
plains extend as deep as 4 km (2.5 mi). The volume this rep-
resents is so enormous that, although the concentrations of
microbes here may be less, the total number of subsurface
microbes is greater than in shallow sediment or in the water.
There are an estimated 10
30
subsurface prokaryote organ-
isms. Most occur beneath the oceans, constituting perhaps
one-third of Earth's biomass. It is becoming clear that mi-
croorganisms in and beneath the sea are the most abundant
organisms on Earth, and they are largely unknown to us.
Extremophiles
Microbes are being found not only miles beneath the Earth's
surface, but in every extreme environment imaginable, so
we call such microbes extremophiles (extreme-loving).
Extreme only from our point of view, these environments
are perfectly normal for the organisms living there. In fact,
most couldn't survive anywhere else. Extremophiles include:
endolithsthose living deep beneath the Earths surface,
without light or oxygen, at extreme temperatures and pres-
sures, feeding on hydrocarbons and inorganic compounds,
some reproducing only once every 100 years.
thermophileslike it hot, found in hot springs and around
hydrothermal vents.
hyperthermophileslike it really hot, also found in hot
springs and hydrothermal vents, and on active seamounts,
where lava is emitted directly onto the sea floor.
psychrophilesfound in extremely cold environments, like
Arctic and Antarctic seawater, sea ice and sediments, the cold
deep sea, and lake ice and glaciers.
barophiles (or piezophiles)live under extreme pressure,
such as deep-sea and subsurface environments.
halophileslive in extremely salty environments, like deep-
sea hypersaline basins, hypersaline lakes like the Dead Sea
and the Great Salt Lake, and solar salt-evaporation ponds.
acidophilesgrow best under extremely low pH condi-
tions, found at some hydrothermal vents, hot springs and
mines.
alkaliphilesprefer extremely high pH conditions, found
in carbonate springs, soda lakes and alkaline soils.
Most extremophiles found so far are bacteria. Many ar-
chaea also are extremophiles, including the record-holding
hyperthermophile, called Strain 121. This archaean, col-
lected from hydrothermal vents at the Juan de Fuca Ridge,
nearly 1.5 miles down in the Pacific Ocean, survived and
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even doubled its numbers when cultured at 121C (250F).
The conventional sterilization regimen is 15 minutes in a
pressurized autoclave at 121C, previously thought suffi-
cient to kill all microorganisms and spores. Some eukary-
ote microbes also are extremophiles, including some soft-
shelled foraminifera pulled from the Challenger Deep
in the Mariana Trench, at pressures of 8 tons per square
inch. Multicellular extremophiles also exist, like the animals
living around hydrothermal vents. Many extremophiles are
polyextremophiles, such as psychrobarophilic bacteria
and archaea living in the cold deep oceans, and psychro-
halophiles in the deep hypersaline basins of the Mediter-
ranean Sea. Organisms living at deep hydrothermal vents
not only thrive under extremes of temperature and pres-
sure, but also in the presence or extremely acid, alkaline
or toxic conditions, and in the absence of light or oxygen.
Microbes survive not only in virtually any conditions,
but also for seemingly indefinite periods of time. Bacte-
rial spores have survived in a spacecraft, after orbiting
the Earth unprotected for years, and have been found
and revived after 750,000 years buried 300 ft deep in a
glacier, after 2540 million years in the gut of a bee en-
tombed in amber, and after 250 million years in salt crys-
tals from an ancient sea bed now 2,000 ft underground!
Intimate Relationships
Microbes saturate not only every nonliving substrate, but
also every living substrate on Earth. Throughout the liv-
ing world, microorganisms may be found engaging in every
kind of symbiosis. Ectosymbionts live on, and endo-
symbionts live in, the bodies of larger organisms. Some
form commensal symbiotic relationships, neutral guests
that neither harm nor help their hosts. Others are para-
sites or pathogens, benefiting themselves at the expense
of the host. Still others form mutualistic relationships
that benefit both partners, a biological win-win situation.
Some are merely opportunistic, engaging in facultative
symbiosis. Obligate symbionts, on the other hand, require
the host or partner for their existence. Microbial mutual-
ists may provide their partners with a variety of services
including nutrients, defenses, and even light (C&H p. 90).
In return, they may receive shelter and a stable environ-
ment, nutrients, or a route for reproduction and dispersal.
One such partnership, involves photosynthetic dinofla-
gellates of the genus Symbiodinium, which drives the
formation of coral reefs. These endosymbionts, commonly
called zooxanthellae, provide reef-building corals with
energy and organic nutrients, receiving inorganic nutrients
and a home in return. Other zooxanthellae form similar as-
sociations with relatives of reef-corals, like soft corals, sea
fans and sea anemones, as well as with sponges, mollusks
and even microscopic foraminifera. Other microbial symbi-
onts drive highly productive communities around hydro-
thermal vents and cold seeps. The rich animal life in these
communities, including tube worms, clams and mussels,
harbor chemosynthetic microbes which provide the fuel
for life in these lightless, sulfur- and methane-rich worlds.
Symbiotic relationships develop over vast spans of time,
transforming the participants in remarkable ways. Some
sediment-dwelling oligochaete worms, for example, have
become gutless wonders: they have no mouth, gut, anus
or excretory system, but instead are entirely filled with
chemosynthetic symbiotic bacteria. In tube worms and bi-
valves living around hydrothermal vents and cold seeps, the
digestive tract also is absent or greatly reduced, with en-
dosymbionts embedded in specialized cells to perform this
function. Shipworms (actually a type of wood-boring clam)
also house bacteria in specialized glands within their gills.
These endosymbionts provide cellulase and nitroginase,
enzymes for cellulose digestion and nitrogen-fixation. This
allows the shipworm to survive on a pure wood diet, without
any additional carbon or nitrogen source. Luminous bacte-
ria colonize specialized light organs, or photophores, in
some fishes, octopuses and squids, enabling them to com-
municate, lure prey, attract mates or blend into their sur-
roundings. These are all examples of obligate symbiosis.
We are only beginning to recognize the pervasiveness and
importance of symbiosis. These relationships are not mere
curiosities, but are essential bonds, deeply woven into the
fabric of life. In untold numbers and varieties, mutualis-
tic microorganisms mediate processes essential to life,
while pathogenic microbes mediate disease, in all organ-
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isms. We are largely ignorant of these relationships, and
most of what we do know comes from studying terrestrial
plants and animals, especially humans. We are walking eco-
systems of commensal and mutualistic bacteria, mites and
other tiny creatures, not to mention our pathogens. Each
cm
2
of skin (about the area of a postage stamp) on the
average adult carries 1,000 to 10,000 bacteria, except the
armpit and groin, which are home to about a million bacte-
ria per cm
2
. Others inhabit our eyes, nose, mouth, throat,
and urinary tract. Residing in the human colon are trillions
of bacteria, including hundreds of types, living in symbio-
sis with each other as well as with us. Many of these invis-
ible guests are essential to our normal healthy functioning.
Equally intimate and complex associations, mostly un-
known to us, must extend throughout the biological world.
In fact, eukaryote organisms arose as a result of symbio-
sis, according to the generally accepted Endosymbiont
Theory (C&H, p. 73). Aerobic bacteria in the ancient oceans
were engulfed by anaerobic prokaryotes and transformed
over evolutionary time into mitochondria, the cell's power
plants where respiration takes place. Similarly, cyanobacteria
engulfed by other prokaryote cells eventually became chlo-
roplasts, where photosynthesis occurs. The ingested cells
were gradually transformed from prey into endosymbionts,
and finally into permanent structures of the host cells. The
nucleus and other membrane-bound organelles of eukaryote
cells are thought to have arisen through similar processes.
The Scum of the Earth
At least 3.5 billion years ago, long before the first eukaryotes
appeared, microbes began forming intimate associations as
microbial mats. Primitive microbes congregated together,
forming patches of scum that grew to cover all the available
space in the shallow ancient seas. Organisms with different
capabilities and requirements took up residence in different
layers of these communities. Primitive photosynthetic bacte-
ria grew near the surface and various heterotrophs resided
within. These ancient communities changed over time, much
as a forest is structured and develops over time, through the
process of ecological succession. They grew in thickness
as living microbes multiplied upward and layers of debris
accumulated below, building into calcareous columns called
stromatolites (C&H Fig. 5.4). Fossil stromotolites can be
found, and recent stromatolites continue to form, in the ocean
today. Microbial mats also occur as patches of scum and
slime, floating in water bodies and coating wet rocks and sed-
iments, in the deep ocean or on intertidal flats, at cold seeps
and gas hydrates, in hot springs and lagoons, salt marshes
and mangrove swamps, lakes and ponds, and even puddles.
These unprepossessing patches of scum, called biofilms,
are becoming important subjects of scientific research.
Biofilms are microbial communities adhering to surfaces.
These living veneers coat just about any damp surface, biotic
or abiotic, on the planet. Some biofilms even occur on sun-
baked desert rocks, creating a colorful coating called "desert
varnish." Biofilms are composed of a variety of microorgan-
isms, which secrete a gooey matrix, glueing them together and
fixing them to the substrate. Like microscopic cities, biofilms
are complex, dynamic communities of interacting organisms.
They manufacture, transport, consume, and compete for re-
sources, and communicate via chemical signals. Biofilms
corrode metal pipes and contaminate medical equipment
and the systems that deliver our water, air and heat. They
cling to the shower curtain and the toilet bowl, and cause
us to slip on wet rocks. As plaque on our teeth, they give us
cavities and periodontal disease. Elsewhere, such as in the
lungs or on a heart valve, their presence may be life-threat-
ening. Biofilms are even more ubiquitous in marine environ-
ments, where they colonize every living and nonliving sub-
strate, every sediment grain, algal frond, sea urchins spine,
whales back and foul every ships hull. Marine microbes
congregate in the water column, as well, coating suspended
detritus particles to form marine snow. Microbes are
gregarious by nature, and biofilms are a prevailing lifestyle.
Many chemical signals are used to orchestrate life in bio-
films. Bacteria continuously produce and detect molecules
called autoinducers. When the level of autoinducers in
a particular area is sufficiently high, the bacteria begin to
form a biofilm. This process is called quorum sensing.
Microbes in biofilms produce other molecules that inhibit
the quorum sensing of their competitors. Eukaryotes, for
example some seaweeds, may also produce chemicals that
interfere with quorum sensing and therefore prevent fouling
of their surfaces by biofilms. Biofilms themselves serve as
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substrates for the attachment and growth of other organ-
isms. Plant spores and the larvae of a wide range of marine
invertebrates, including corals, sea urchins, and polychaete
worms, settle onto biofilms to begin the benthic phase of
their life history. Biofilm organisms produce chemical cues
that induce both the settlement and the metamorphosis
of these larvae. Biofilms coating reef corals, for example,
induce the settlement of coral larvae and their metamor-
phosis into juvenile polyps. The ocean may be a cacophony
of these chemical signals, inducing, competing and defend-
ing. Some of these chemicals may provide us with new an-
tibiotics or antifouling agents, allow us to enhance aqua-
culture, or help in the recovery of damaged coral reefs.
Microbial Diversity
We know that prokaryotes, bacteria and archaea, account
for two of the three main branches on the universal tree
of life, as distant from each other as they are from the
eukaryotes. But how many different types of archaea and
bacteria are there, how many twigs on those branches?
Clearly, they are incredibly abundant, amazingly ubiqui-
tous, flexible and tenacious forms of life with a history of
3.8 billion years of evolution. They are present everywhere
we look, able to live in any environment. An entourage of
symbiotic microbes, both helpful and harmful, lives on and
in every bigger organism, perhaps hundreds of types of
symbiont living with each host, many of them unique to a
particular host species. This alone suggests that microbial
diversity is huge, far greater than has so far been described.
Metagenomics, or whole genome shotgun sequenc-
ing, is a recently developed technique that has produced
tremendous insights into microbial diversity. It gives a profile
of diversity by sequencing genetic material in samples taken
directly from nature, without cultivating. Among the first to
apply this technique, Norwegian researchers in the 1990s
sequenced DNA in small samples of forest soil and marine
sediment. In 1 gram of forest soil (about the contents of a
tea bag), they found 4.8 billion prokaryote individuals, repre-
senting about 6,000 different types of organism. In a similar
sample of marine sediment, they found 3.1 billion individuals,
of about 11,400 types, almost none of which matched those
in the forest soil sample. The prokaryote diversity detected
in these two small samples alone is several times greater
than that described by scientists during the past 250 years.
The Global Ocean Sampling Expedition, led by Craig
Venter, is perhaps the most ambitious use of metagenom-
ics. In 2003, these researchers began circumnavigating the
globe and collecting metagenomic samples. In initial samples
from six sites in the Sargasso Sea, where very low nutrient
levels occur and species diversity was assumed to be low, they
detected at least 1,800, and possibly as many as 50,000,
new species. The DNA in these samples included 1,214,207
new genes, a significant addition to the number currently in
public databases, including genes coding for 782 new forms
of photoreceptor genes, nearly four times as many as had
previously been discovered from all species. Some of the
sequences analzyed are so unusual that they may even rep-
resent a fourth domain of life! (C&H p. 93: "Eye on Science")
The diversity of microorganisms lies not in the variety of their
physical forms (their morphology) but in their incredible met-
abolic flexibility. The ability of microbes to live everywhere,
under any conditions, stems from their ability to evolve new
metabolic capabilities. They continually evolve to function un-
der different physical conditions, exploit different chemicals to
obtain nutrients and energy, and synthesize needed compo-
nents, via new metabolic pathways. Prokaryotes are capable
of using every natural compound on Earth and most human-
made compounds as well. Some of the metabolic pathways
evolved by marine prokaryotes are outlined in Table 5.1 of
your textbook. Understanding the metabolic diversity of mi-
croorganisms is one of the greatest scientific challenges.
So, how many microbial species are there? Suffice it to say,
we dont know and may never know, but its a lot! The emi-
nent biologist, Edward O. Wilson, refers to prokaryote diver-
sity as the black hole of taxonomy. The decade-long Cen-
sus of Marine Life (COML) begun in 2000 to assess marine
biodiversity and target conservation efforts (C&H p. 216), in
2003 spawned the International Census of Marine Microbes
(ICOMM). The goals of ICOMM are to catalog the known di-
versity, and explore and discover the unknown diversity, of
marine microbes - unveiling the oceans hidden majority.
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Mighty Microbes: The Importance of
Marine Microbes
The importance of microorganisms in the ocean, though
poorly understood, cannot be overstated. In their over-
whelming abundance and diversity, they support marine eco-
systems at nearly every level. They are the producers of
the sea, autotrophs conducting 98% of marine primary
production, as we will see in the next lesson, and supply-
ing the energy and organic materials for all other marine
life. Microbes also are responsible for nitrogen fixation,
supplying this crucial nutrient in useable form to producers.
Microbes make up the bulk of the primary consumers,
as well, the heterotrophic bacteria, archaea and pro-
tozoans, that graze on tiny phytoplankton, too small to
be eaten by anyone else. These zooplankton are them-
selves eaten by predatory zooplankton - secondary con-
sumers - which in turn fall prey to larger marine animals.
Microorganisms are also the principal decomposers of all
dead organic matter in marine systems, playing the vital role
of returning the nutrients contained in this material to the
environment, where they again become available to produc-
ers. In addition to the particulate detritus of dead organ-
isms, as much as half of the organic matter manufactured by
marine producers becomes DOM (dissolved organic matter),
leaked, spilled and excreted into the ocean by organisms.
Microbes are responsible for consuming DOM and returning
its nutrients, which would otherwise be lost, to the rest of
the marine food web. This critical part of the marine food
web is called the microbial loop (C&H pp. 349350).
Without the microbial loop and decomposition by marine
microbes, life in the ocean would quickly grind to a halt.
Countless undecomposed dead microorganisms, on the
other hand, are responsible for current oil and gas de-
posits. In ancient oceans, over millions of years, dead
microbes became rapidly buried in anoxic deep-sea sedi-
ments, where decomposition is slowed or stopped. They
gradually accumulated and "pressure-cooked," forming the
matured oil and gas deposits on which we are so dependent.
Marine microbes also play vital roles in global biogeo-
chemical cycles, which we learned about in Lessons 2 and
4. The CO
2
taken up by marine microbes during photosyn-
thesis is crucial not only to the marine carbon cycle, but
also for the absorption of excess CO
2
in our atmosphere,
moderating the greenhouse effect and global warm-
ing. Methanogens, on the other hand, are adding an-
other greenhouse gas, methane, to the atmosphere.
Mostly archaea, these methane-producers live in anoxic
marine environments, swamps, the intestinal tracts of ani-
mals, and sewage treatment plants where they are used to
break down waste. By generating methane, methanogens
are contributing to the problem of global warming, but they
might also be harnessed to produce methane for energy.
On a geologic time scale, marine microbes also are respon-
sible for Earths habitability. The activity of photosynthetic
marine organisms over billions of years produced the ox-
ygen in our atmosphere that we breathe, and the ozone
in the stratosphere which protects us from the sun's le-
thal radiation. Marine microbes were, in fact, the source of
all life. The only forms of life for most of its history, they
gave rise to all other living things. Microbial life continues
to hold together the fabric of life, as producers, consum-
ers, nitrogen-fixers and decomposers. They live in and on
all organisms, in intimate and necessary associations, and
as pathogens. They even formed the organelles of our cells.
We study microorganisms in our attempt to understand the
origins of life, and to answer many other basic questions in
biology. As pathogens, they are the subjects of biomedical
research into the cause and treatment of disease. We even
look to them when pondering the possibility of life on other
planets. Some extremophiles today live in environments that
may be similar to those on Earth when life began, or similar
to those on other worlds, like Mars or Jupiters moon, Eu-
ropa. The study of these organisms has helped to launch a
new field of science: astrobiology, the study of the effects
of living in space, and the possible origins, distribution and
evolution of extraterrestrial life. Microbes also are valuable
subjects of study in many areas of applied biology. Their
amazing metabolic diversity provides a virtually unlimited
source of potentially useful products and processes. Already,
they are being used to treat wastewater, degrade pollutants,
diagnose diseases, and to produce pharmaceuticals and
many other industrial products. With most microbial diversity
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yet to be discovered, the potential for exploiting microorgan-
isms through bioprospecting is essentially boundless.
Finally, a note about viruses and fungi: It has long been
debated whether to classify viruses as living things, much
less where on the tree of life they might belong. They do
not have a cell membrane, consisting only of genetic mate-
rial in a protein shell, and cannot metabolize or reproduce
on their own. They can, however, respond to the environ-
ment in a limited fashion and infect the cells of organisms
(their name means poison), replicating themselves by
self-assembly. Whatever their status in the living/nonliving
debate, viruses are without a doubt important in the ocean,
where they outnumber microbial cells 10:1 or 100:1, and
infect bacteria (as bacteriophages) and all other forms of
life. By infecting marine microbes, they may profoundly af-
fect the flow of energy and cycling of nutrients in marine
ecosystems, and may even affect the global carbon budget
and global climate. They also appear to play an important
role in influencing microbial diversity. Microbes are constantly
evolving to evade viral diseases, and viruses can shuffle
the genes of microbes, transferring genetic material between
host cells. Viruses are another world largely unknown to us.
There is no question that fungi, on the other hand, are living
organisms. Surprisingly, they are not at all closely related
to plants, but are the sister group to the animal kingdom
(Fig. 11). Marine fungi are mostly microscopic, little-stud-
ied organisms. They are best known in mangrove forests,
where they are the most important decomposer organisms.
They are probably important as decomposers in other ma-
rine environments, particularly in sediments. Some types
of fungi are parasites, whereas others engage in mutual-
istic associations with cyanobacteria and algae, creating
lichens. Fungi are better-studied in terrestrial communi-
ties, where they are intimately associated with the roots of
most plants. These mutualistic relationships are vital to the
health of terrestrial plants, and were crucial to their initial
colonization of the land, as we will see in the next lesson.
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glossary terms
your textbook.
Bacteria, Archaea and Eukarya
protists, Protista
virus and bacteriophage
fungi and lichen
phytoplankton and zooplankton
picoplankton
photosynthesis and chemosynthesis
cyanobacteria, picoplankton, Synechococcus
and Prochlorococcus
algae: diatom, dinoflagellate, silicoflagellate,
coccolithophorid and cryptophyte
protozoan: foraminiferan (foram), radiolarian
and ciliate
frustule and test
siliceous ooze, diatomaceous ooze, and fora-
miniferan ooze
epiphytes and endophytes
extremophile: endolith, thermophile, hyper-
thermophile, Strain 121, psychrophile, baro-
phile, halophile, acidophile, alkaliphile
symbiosis: commensalism, mutualism and
parasitism
facultative symbiosis and obligate symbiosis
ectosymbiont and endosymbiont
Endosymbiont Theory
mitochondria and chloroplasts
zooxanthellae, Symbiodinium
microbial mat, stromatolite, biofilm, autoin-
ducer and quorum sensing
microbial loop, DOM, detritus and marine
snow
methanogen
taxonomy and phylogeny
morphology
molecular sequencing, whole genome shotgun
sequencing (metagenomics)
epifluorescence microscope
Global Biodiversity Assessment, Global Ocean
Sampling Expedition, COML, ICOMM
mass extinction
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unicellular algae and protozoans - both protists, single-celled eukaryotes. Algae are mostly autotrophs, photosyn-
thetic producers. Protozoans are mostly heterotrophs, consumers.
a. Archaea and Bacteria
b. Pfiesteria and Prochlorococcus
c. morphology and molecular sequencing
d. microbial mats and the microbial loop
e. ectosymbiont and endosymbiont
f. cyanobacteria and bacteriophage
g. Strain 121 and stromatolite
h. DOM and ICOMM
2. Most of the ocean floor is thickly covered with biogenous sediments composed of the skeletal remains of marine
microbes. Compare the "skeletons" (shells) of diatoms, dinoflagellates, foraminiferans, coccolithophorids and silico-
flagellates.
3. More than a million animals have been identified, compared to a few thousand known bacteria. Do you think this is
an accurate reflection of the relative diversity of animals and bacteria in nature? Give two detailed reasons for your
answer.
4. The abundance of microbes in the pelagic environment is enormous. The tiny picoplankton Prochlorococcus and
Synechococcus may be the most numerous organisms on the planet. There are 50,000-500,000 prokaryote cells in
each milliliter of seawater, adding up to a total of 10
29
organisms in the pelagic environment. How does this compare
with microbial abundances on the sea floor, and within the sea floor (subsurface) environments?
5. Name and describe three types of extremophile, including their preferred habitats in the marine environment.
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6. Name two microbial mutualists in marine communities and describe their relationships with their hosts.
7. Explain three critical functions that microorganisms perform in marine ecosystems.
8. How common are biofilms in marine environments? Where might you expect to find them? Describe three ways that
chemical signals operate in biofilms.
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The Producers:
Marine Autotrophs
and Primary
Production
This lesson concerns the organisms at the base of the trophic pyramid, the producers
that convert inorganic materials into organic matter, supplying the energy and materials
necessary for all other forms of life.
1. Understand how the types, diversity and history of marine primary producers differ
significantly from those on land.
2. Be familiar with the major types of marine autotrophs.
3. Understand the role of primary producers in marine ecosystems.
4. Understand the factors controlling marine primary productivity.
5. Recognize the importance of marine primary productivity to Earth's habitability
past, present and future.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 6 (pp. 102113) and pp. 237239 ("Overwhelming the Nitrogen Cycle"
and "The No-Zone"), pp. 334-335 ("The Phytoplankton"), p. 336 (Table 15.1), pp.
338-339 ("Red Tides and Harmful Algal Blooms) and review pp. 221-229 ("The
Flow of Energy and Materials")
6
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The producers
(Marine Autotrophs and Primary Production)
T
his lesson concerns the organisms at the base of the
trophic pyramid, the producers (or primary pro-
ducers) that convert inorganic materials into organic
matter, supplying the energy and materials necessary for all
other forms of life. This process, called primary production,
differs greatly in marine and terrestrial environments. We be-
gin with a brief history of the evolution of producers on land
and in the sea, to help put these differences in perspective.
Seeding the Land
In our terrestrial world, the vital function of primary produc-
tion is carried out by the familiar and obvious green plants.
Green plants include about 300,000 species of land plants
and about 7,000 green algae. Land plants evolved from
green algae around 460 mya (million years ago), and these
two groups still share biochemical characteristics that distin-
guish them from other photosynthetic organisms (C&H Table
6.1). Vascular plants, land plants with water-conducting
structures and other specialized tissues, appeared 2040
million years later. These specialized structures allowed
them to evolve much larger forms, and to spread into many
more environments. Seed-bearing vascular plants ap-
peared about 400 mya. Gymnosperms ("naked seeds"),
like our pines and spruces, dominated during the age of the
dinosaurs. Angiosperms (seeds enclosed in an ovary), or
flowering plants, were the last major group to appear, less
than 200 mya.
Flowering plants diversified and spread explosively, now com-
prising the vast majority of land plants. They live in all but the
most extreme environments and conduct most of the primary
production on land. Crucial to human existence, they sup-
ply us with food, clothing, shelter, fuel, medicines and other
useful chemicals. Their productivity is vital to the balance of
oxygen and carbon dioxide in the atmosphere. The peak of
flowering plant diversity and productivity occurs in tropical
rainforests, which contain more than half of all plant and
animal species so far described by science. We are currently
destroying tropical rainforests at a rate of 1% per year, and
have already destroyed more than 50% of their coverage.
All this productivity and diversity on land was made possible
by life in the sea, which flourished for more than 3 billion years
before the first green algae emerged on land and eventually
gave rise to land plants. Fungi emerged with the algae and
many became mutualistic symbionts, now necessary to the
health of most plants. The first land animals, both inverte-
brates and vertebrates, also came from the sea, where near-
ly all the modern phyla had already evolved. Besides seeding
the plant, fungal and animal life on land, marine organisms
also are responsible for making terrestrial environments
habitable for them. For most of Earths history, the land
was uninhabitable, bombarded by the suns lethal radiation.
Through billions of years of photosynthetic activity, marine
bacteria and archaea added oxygen to the ocean and at-
mosphere, which created a protective layer of ozone in the
stratosphere (also damaged by human activities). This build-
up of oxygen also allowed larger multicellular organisms, with
greater requirements for aerobic respiration, to evolve.
Microbial Producers: The Phytoplank-
ton
Unlike their terrestrial counterparts, the vast majority of pro-
ducers in the ocean are microscopic organisms, many still
undiscovered. For the first two billion years of lifes history,
the ocean swarmed with simple prokaryote cells. Micro-
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bial mat communities developed, including photosynthetic
organisms like cyanobacteria, as well as chemosyn-
thetic forms such as sulfur-oxidizing bacteria, which
derive energy from splitting hydrogen sulfide molecules,
rather than from sunlight. Nearly two billion years later,
eukaryotic cells appeared. According to the Endosymbi-
ont Theory, single-celled algae arose as a result of cells
engulfing smaller photosynthetic cells, like cyanobacteria,
which became chloroplasts. The first multicellular pro-
ducers were tiny leaf-like seaweeds, first brown algae,
followed by multicellular red and green algae. About the
same time, the first tiny multicellular animals appeared.
Today, more than 95% of marine primary production is
still carried out by phytoplankton, microscopic photo-
synthetic organisms drifting in sunlit waters. Phytoplankton
have long been known to include unicellular algae such as
diatoms and dinoflagellates, easily caught in standard
plankton nets, and some photosynthetic bacteria, pri-
marily cyanobacteria (formerly called blue-green algae).
However, as we learned in the previous lesson, new tech-
niques (molecular sequencing, metagenomics, and
epifluorescence microscopy) have revealed the most
abundant phytoplankton: Synechococcus and Prochlo-
rococcus. These minuscule cyanobacteria are members
of the tiny picoplankton, too small to be caught in nets,
and are arguably the most abundant organisms on Earth.
The phytoplankton are a hugely diverse group. Prokary-
otic members, the most abundant, include many types of
cyanobacteria, plus purple and green bacteria, and many
photosynthetic archaea. Eukaryotic algae include diatoms,
dinoflagellates, coccolithophorids, silicoflagellates,
and cryptophytes, as well as unicellular brown, red and
green algae. More minute algal cells, previously unknown,
also are being discovered using metagenomics (C&H p. 92:
"Tiny Cells, Big Surprises"). The major groups of phytoplank-
ton are summarized in your textbook (Table 15.1, p. 336).
These groups have evolved various light-capturing photo-
synthetic pigments and pathways for carrying out photo-
synthesis (C&H Tables 5.1, 5.2 and 6.1). Cyanobacteria are
particularly diverse, with an assortment of photosynthetic
pigments imparting a rainbow of colors to these organisms.
The Red Sea gets its color from red cyanobacteria, pink flamin-
gos are colored by the cyanobacteria they ingest, and the fur
of polar bears is sometimes tinged green by cyanobacteria.
Other Microbial Producers
In addition to the free-living plankton, microbial sym-
bionts also contribute to primary production in the sea.
Photoautotrophic symbionts conduct photosynthesis,
providing energy and organic matter to their hosts, in re-
turn for a home and other benefits. The best known are the
zooxanthellae. Symbiodinium, a genus of dinoflagel-
lates, are zooxanthellae who partner with corals and are
critical to the formation of coral reefs. Other symbionts form
associations with mollusks, sponges and even tiny foramin-
ifera. Photosynthetic bacteria of the genus Prochloron
provide similar services to their hosts, the sea squirts.
Most primary production in the ocean, as on land, is fueled
by solar energy. The dark, cold deep-sea environment was
long thought to be lifeless and unproductive. Organic matter
produced by epipelagic phytoplankton sinks down to lower
levels, but it is meager fare in this vast realm. It is now known
that some bacteria and archaea thrive in these environments,
as they do everywhere else. Innovative chemosynthetic
prokaryotes use the energy contained in chemicals like hy-
drogen sulfide, methane, ammonium and iron to synthe-
size organic matter. In certain habitats, unknown until recent
decades, these chemoautotrophs have been found sup-
porting incredibly rich communities. Hydrothermal vents,
the deep-sea hot springs associated with mid-ocean ridges
and trenches, vare surrounded by flourishing communities of
tubeworms, clams, mussels and other invertebrates, fueled
by chemosynthetic bacteria and archaea. Similar communi-
ties also exist at cold seeps, where hydrogen sulfide and
methane seep out from the sea floor, and even at deep-sea
graves, the decomposing remains of large dead animals
like whales which also produce hydrogen sulfide and meth-
ane. Many chemoautotrophs are free-living, primarily in
sediments or growing into thick mats on the ocean floor. Oth-
ers are endosymbionts, living within the tissues of many of
the animals which inhabit these rich, lightless environments.
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Multicellular Producers
While at least 95% of marine primary production is carried
out by phytoplankton, most of which are recently discov-
ered picoplankton, multicellular organisms are important
producers in some coastal habitats. Multicellular algae,
the seaweeds, are common along rocky shorelines and
other shallow-water habitats. Although they appear similar
to land plants, seaweeds lack true leaves, stems and roots,
and are physiologically distinct from land plants (C&H Table
6.1). In distinct contrast to terrestrial systems, few flower-
ing plants are marine producers. Like the seaweeds, they
are restricted to shallow coastal environments. The habitats
where they do occur, however, are some of the most pro-
ductive on Earth. Marine flowering plants include the trees
and shrubs of mangrove forests, the cordgrasses and
pickleweed found in saltmarshes, and the seagrass-
es (more closely related to lilies than grasses) which are
widespread in temperate and tropical bays and estuaries.
The coastal environments that are home to seaweeds and
marine flowering plants, and the rich animal communities
they support, are threatened by many human activities.
Their importance as habitats for migrating birds, spawning
fish, and many other animals, and as buffers for the preven-
tion of flooding and erosion, often has been poorly under-
stood or ignored. Seagrass beds and other coastal habitats
in many places have declined dramatically as a result of
dredging and filling, diversion of freshwater sources, and
pollution and other problems caused by coastal develop-
ment. Another rapidly increasing threat is the introduction
of invasive species (both plant and animal), often car-
ried on the hulls or in the ballast water of ships traveling
from port to port, over great distances. Non-native species
can upset delicately balanced ecosystems, growing uncon-
trollably, destroying native species and spreading disease.
Two Different Worlds
The stark differences between most marine primary produc-
ers and those on land are the result of adaptations for liv-
ing in two very different environments. The low density of
air, in which light readily penetrates to the ground, and the
presence of water and nutrients in the soil, have shaped the
large land plants. They have roots in the ground to take up
water and nutrients, and trunks and leaves raised to the sky
to absorb sunlight and carbon dioxide. The high density of
seawater, in which water and dissolved nutrients are read-
ily accessible, but light decays rapidly as it penetrates, cre-
ates a very different situation. Microscopic phytoplankton
can absorb water and nutrients directly through their cell
walls. They also can remain suspended in the photic zone,
where there is enough light for photosynthesis to occur.
Their buoyancy is dependent on their low weight and rela-
tively high frictional drag. The tiniest cells have minimized
their weight and maximized their surface-to-volume ra-
tio, giving them enough drag to prevent sinking. Larger
cells like diatoms increase their surface area, and their
drag, with elaborate shells bearing spines and pores. The
larger seaweeds also absorb water and nutrients directly
through their cell walls, but are restricted to shallow habi-
tats where their holdfasts can attach to the substrate. Ma-
rine flowering plants, like their terrestrial relatives, use their
roots to attach to the substrate, and to absorb nutrients.
Primary Production
Primary production is the amount of biomass created
by primary producers, usually expressed as the amount of
carbon fixed, or the amount of carbon dioxide taken up.
The primary productivity of a system is the amount
of primary production per unit of time, usually expressed
as the amount of carbon fixed under one square me-
ter of sea surface in a given amount of time, for example:
grams C/m
2
/year
Scientists are extremely interested in these values, since
marine primary productivity supports all other marine or-
ganisms, many of which are important resources for hu-
mans, and strongly influences the global carbon cycle
and global climate. Measuring the primary productivity
of an ecosystem is like taking its pulse: it is a key vital
sign telling us what is happening in the system, indicat-
ing its overall health and useful for tracking changes in
the system over time. Just as primary producers are much
more difficult to detect in the ocean, so primary produc-
tion also is more difficult to measure in marine systems.
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Primary production is a function of many factors, including
light, temperature, and the availability of various nutrients
and water. Primary productivity on land is usually limited by
water availability, temperature and light, in that order. In ma-
rine environments, light, nutrient availability and temperature
are usually the limiting factors, since water is always avail-
able. In other words, lightless and nutrient-deficient environ-
ments are the deserts of the sea. The vast majority of
marine producers therefore live in the epipelagic (photic)
zone, light-filled and relatively warm. The intensity of light
falls off rapidly as it penetrates water, diminishing to a point
where the rate of photosynthesis is only enough to supply the
maintenance needs of the organism, and there is no growth,
or net productivity. At lower light levels, in the dysphotic
zone, phytoplankton are not able to conduct photosynthesis
at all. Deeper still, in the aphotic zone, no light penetrates.
Organisms differ in their sensitivity to light, and light inten-
sity varies widely, depending on latitude, daily and seasonal
changes in solar radiation, cloud cover, and the clarity of
the water. Much of the incoming solar radiation goes into
heating the water. Since the rate of photosynthesis also is
dependent on temperature, it would seem that productiv-
ity would decrease steadily from the equator to the poles,
but often the opposite is true. This is because nutrient
availability also strongly influences production. Availabil-
ity of macronutrients, those required by producers in
large doses, often limits productivity on a regional scale.
Carbon, the most important macronutrient, is always read-
ily available as dissolved carbon dioxide, but nitrogen,
phosphorus and silicon may become limiting nutrients.
Micronutrients, also required but in smaller doses (in-
cluding iron, copper, manganese and zinc), may limit
production on a local scale. Organisms vary in their
need for essential nutrients, and nutrient availabil-
ity is extremely variable, influenced by many other factors.
In the case of some algal species, like diatoms and silico-
flagellates, silicon may be the limiting macronutrient, as it is
needed to build their shells. Most often, however, nitrogen is
the limiting nutrient, in environments with adequate light and
warmth. As N
2
, nitrogen makes up 78% of our atmosphere,
but it must be fixed, or converted into reactive nitrogen,
for producers to use. The presence of nitrogen-fixing
cyanobacteria, and other nitrogen-fixing bacteria and ar-
chaea, strongly influences primary production in marine en-
vironments. Other natural sources of fixed nitrogen in marine
environments include input from rivers and electrification by
lightning. However, humans are the source of most of the re-
active nitrogen in the ocean today, from sewage release, run-
off of agricultural fertilizers into rivers, and burning of fossil
fuels. This degree of nutrient enrichment, actually nutrient
pollution, causes eutrophication, a massive increase in the
growth of algae. Massive growth is followed by massive de-
composition, depriving the environment of oxygen (hypoxia).
Hypoxic zones, or dead zones, created by eutrophication
exist in many places today, such as at the mouth of the Mis-
sissippi River, extending westward along the Gulf of Mexico.
Sudden increases in the growth of phytoplankton are called
blooms. They are a periodic natural occurrence, triggered
by favorable environmental conditions. In temperate and
polar waters, annual spring blooms occur, when light
and warmth return and phytoplankton can take advantage
of nutrients accumulated during the darker winter months.
These seasonal blooms may provide most of the primary
production for the entire year in these regions. Red tides
(which aren't always red and don't involve tides), are an-
other type of bloom. They are caused by a single species
of phytoplankton and occur unpredictably, for unknown rea-
sons. Red tides involving some species of dinoflagellates,
diatoms and cyanobacteria periodically foul coastal waters
around the world, sometimes producing deadly toxins.
They seem to be increasingly frequent, possibly as a result
of increased human activities and pollution along coasts.
Primary productivity in large areas of open ocean are limited
by the micronutrient, iron. Iron was plentiful in the ancient sea,
but today is in short supply in the subarctic Pacific, equatorial
Pacific, and Southern Oceans. When scientists added iron to
these environments, dramatic blooms resulted. This prompt-
ed controversial proposals to fertilize these areas with iron,
both to increase productivity and to slow global warming, by
increasing carbon dioxide uptake. Massive dust storms are an
important source of iron in large areas of the ocean. About
500 million metric tons is blown yearly from the Sahara des-
ert, settling into the North Atlantic, where it boosts primary
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production and sometimes causes harmful blooms. Massive
dust storms originating in the Gobi deserts of northern China
and Mongolia contribute iron to the North Pacific, as well.
The most important source of nutrients in the ocean, how-
ever, is the deep-sea waters and bottom sediments. Gravity
rules in the sea, as on land, causing material to sink. The
constant pull of gravity on dead organic matter results in a
net transport of nutrients toward the sea floor - the carbon
pump we learned about in Lesson 4. Sinking dead organic
matter is a crucial source of energy and materials for deep-
sea consumers. But continued primary production in the epi-
pelagic depends on its decomposition and the return of its
component nutrients to the surface. This is accomplished by
mixing, facilitated by wind, waves, currents and tides, and
especially by upwelling, the slow persistent rising of water
toward the surface. Upwelling occurs along coasts and at
the equator, and in the Southern Ocean around Antarctica,
one reason that these cold waters are highly productive.
Mixing occurs in polar and temperate waters as a result of
overturn. Surface water, made denser during cold winter
months, sinks and displaces nutrient-rich deeper water to
the surface, ready to fuel the spring bloom. The lack of sea-
sonality in the tropics tends to leave tropical waters more
stratified, resulting in lower but relatively constant primary
productivity in the tropical ocean. Many complex processes
cause movements of water, on a small scale or globally.
Their effects on primary production are powerful, and ex-
tremely variable with respect both to time and location.
Since productivity is so variable in this vast and complex en-
vironment, it's difficult to calculate by the traditional meth-
od of measuring the amount of carbon dioxide consumed
(or oxygen given off), or the amount of light absorbed by
chlorophyll, in seawater sampled from ships. Newer tech-
nologies using remote sensors carried by satellites, and
computerized mathematical models, provide scientists with
real-time, global maps of primary productivity. A 2-minute
satellite scan might contain 2 million pixels, covering 2 mil-
lion square kilometers of ocean, whereas for a ship travel-
ing at 20 km/h, it would take more than 11 years to cover
just one pixel of area. The remote sensors detect levels of
chlorophyll-a near the ocean surface, producing images
of phytoplankton concentrations. These images are referred
to by NASA (National Aeronautics and Space Administra-
tion) as ocean color (C&H Figs. 1.13 and 10.19). Ocean
color images can be used to detect blooms and track their
patterns by compositing images over a week, a month or
a year. This technology is revolutionizing our knowledge of
ocean (and global) systems, at a time when understand-
ing ocean dynamics has become particularly important.
Many ocean environments are less productive than the most
productive terrestrial environments, but the vast area cov-
ered by the ocean adds up, so that marine and terrestrial
primary producers contribute about equally to global primary
production (C&H Table 10.1). However, as we learned in Les-
son 3, CO
2
is highly soluble in water, and the ocean holds 50
times as much carbon as in the air. The fate of fixed carbon
differs greatly between terrestrial and marine ecosystems.
When dead organic matter decomposes on land, carbon di-
oxide promptly returns to the atmosphere. But much of the
dead organic matter in the ocean sinks out of reach of decom-
posers and accumulates in ocean sediments. Sediments con-
tain 25 times as much carbon as in ocean water. This carbon
sequestration has been hugely important for moderating
the greenhouse effect and global warming. A study by
NASA scientists calculated that marine primary productivity
has declined by more than 6% globally, since about 1980.
This corresponds with increased atmospheric and sea sur-
face temperatures, which creates more stratification of ocean
waters and inhibits the overturn necessary for spring blooms
in polar and temperate seas. If these estimates are correct,
they are very concerning, considering the significant role that
marine primary production plays in the global carbon cycle.
Efforts to calculate productivity in marine environments, and
to decipher changes that have been occurring and predict
long-term trends, are extremely difficult but are critically im-
portant to our understanding of our rapidly changing world.
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glossary terms
your textbook.
phytoplankton, picoplankton, plankton net
cyanobacteria, purple bacteria, green bacteria
and sulfur-oxidizing bacteria
Synechococcus and Prochlorococcus
single-celled algae (diatoms, dinoflagellates,
silicoflagellates, coccolithophorids, crypto-
phytes, and brown, red and green single-
celled algae)
seaweeds (green, brown and red multicellular
algae)
sea lettuce, calcareous green algae, rock-
weeds, kelps, coralline algae
thallus, stipe, blade, holdfast and pneumato-
cyst
sporophyte and gametophyte
alternation of generations
angiosperms/flowering plants (halophytes:
seagrasses, cordgrasses, mangroves)
free-living, endosymbiont, ectosymbiont,
Endosymbiont Theory
zooxanthellae, Symbiodinium, Prochloron
chemosynthesis and photosynthetsis (chemo-
autotroph and photoautotroph)
chloroplast
photosynthetic pigments (chlorophyll a, c,
carotenoids, phycobilins, etc.)
surface-to-volume ratio, drag
primary producer and primary production
productivity and net productivity
photic, dysphotic and photic zones
upwelling, overturn, stratification
limiting factor/resource
macronutrient and micronutrient
nitrogen-fixation, reactive nitrogen
carbon-fixation, carbon pump/sink/sequestra-
tion
greenhouse gas, greenhouse effect
global warming, acidification
bloom, spring bloom and red tide
eutrophication, hypoxia and dead zone
invasive species
ocean color
ozone layer
After completing the written assignment for this lesson you will be
ready for the rst exam. Locate a proctor in your area and complete
the Exam Request Form on the following pages. (If you live in the Fair-
banks area there is no need to complete the form. Come to our ofce
during business hours to take your exam.) Allow 4 hours for complet-
ing the exam.
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1. Define, by comparing and contrasting (you know the drill), the following terms:
a. photic zone and dysphotic zone
b. phytoplankton and zooplankton
c. seagrass and seaweed
d. silicoflagellate and sea lettuce
e. Synechococcus and Symbiodinium
f. mangal and kelp forest
g. photoautotroph and chemoautotroph
h. chlorophyll a and carotenoids
i. sporophyte and gametophyte
j. macronutrient and micronutrient
k. dead zones and ocean color
2. By far most of the primary production on land is conducted by the hugely diverse flowering plants. Are flowering
plants also responsible for most of the production in the ocean? Who are the most important marine producers?
(include 5 specific examples of marine producers in your answer)
3. Marine primary productivity is strongly influenced by light. When light intensity diminishes below a certain level, pho-
tosynthesis cannot occur. Most primary production takes place at the surface, where light levels are highest. Light
intensity also varies with latitude, daily and seasonal fluctuations, cloud cover and water clarity, all of which affect
primary production regionally and locally. Describe how one other factor influences primary productivity in the ocean.
4. Describe and compare spring blooms, red tides and eutrophication.
5. Marine primary producers are responsible for about half of Earth's primary productivity, but they play a much larger
role in the global carbon cycle. Follow a molecule of carbon dioxide from the atmosphere into the ocean, and de-
scribe what happens to it there. There are various possibilities, but be sure to include the following in your descrip-
tion: photosynthesis, carbon-fixation, decomposition, calcium carbonate, and carbon sequestration.
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INFORMATION LETTER TO BE GIVEN TO YOUR PROCTOR
You have been designated as a proctor for Independent Learning examinations.
The integrity of the examination process is fundamental to our program because it provides the only supervised
check of the students knowledge and capability. We therefore are concerned that prospective proctors under-
stand their responsibilities and agree to ensure that integrity. When an individual is accepted as a proctor, s/he
represents the University of Alaska Fairbanks and is accountable for the examination process.
Proctors must be education officials at a university, community college or an administrator at a public school site
or library, other governmental or community officials, or if, such persons are unavailable, other people approved in
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Proctor responsibilities include:
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Please sign the students Examination Request Form before s/he returns it to our office and keep this Information
Letter for reference when administering the examination.
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EXAMINATION REQUEST FORM
INSTRUCTIONS
1. If you live in the Fairbanks area there is no need to complete this form. Come to our office during regular business
hours to take your exam.
2. Select an approved testing site and arrange an appointment with an eligible individual to proctor the examination.
Ensure that the proctor has read the attached Letter of Information and has signed this Request Form. Schedule
the exam far enough in advance to allow for receipt of all lessons preceding the exam and the mailing of the test
itself.
3. Send this request form in a pre-paid envelope provided with course materials or to:
UAF eLearning & Distance Education
2175 University Avenue South, Suite 200 , PO Box 756700
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PLEASE PRINT OR TYPE
Student Name:
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BIOL F150 UY4 Exam 1
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Marine
Invertebrates
The lower
phyla
Moving up the trophic pyramid, we turn to the consumers, the heterotrophs that rely
directly or indirectly on producers for their energy and organic materials. We are already
familiar with the smallest consumers, the heterotrophic archaea and bacteria, and the
protozoans. Now we begin to study the more visible and familiar consumers, the members
of the animal kingdom.
1. Be aware of the basic trends in the evolution of animals, from sponges to chor-
dates.
2. Be familiar with the "lower" invertebrate phyla, their body plans and life histories.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 7 (pp. 115149) and p. 295 ("Life in Mud and Sand")
7
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Fig. 12 Animal Classification
Subkingdom Parazoa
Phylum Porifera: sponges
Subkingdom Eumetazoa
Radiata
Phylum Cnidaria
Class Hydrozoa: bushy/feathery colonies
of polyps, release minute medusae
Class Scyphozoa: larger jellyfishes
Class Anthozoa: sea anemones, corals, sea
fans, etc.
Class Cubozoa: cubical jellyfishes
Phylum Ctenophora: comb jellies
Bilateria
Superphylum Platyzoa
Phylum Platyhelminthes (flatworms)
Class Turbellaria: freeliving flatworms
Class Trematoda: flukes
Class Cestoda: tapeworms
Phylum Gastrotricha
Phylum Rotifera
Phylum Gnathostomulida
Phylum Cycliophora
Superphylum Lophotrochozoa
Trochozoans
Phylum Mollusca
Class Gastropoda: snails, limpets, abalo-
nes, nudibranchs, etc.
Class Bivalvia: clams, mussels, oysters, etc.
Class Cephalopoda: octopuses, squids,
cuttlefishes, etc.
Class Polyplacophora: chitons
Class Scaphopoda: tusk shells
Phylum Annelida
Class Polychaeta: marine segmented
worms
Class Oligochaeta: earthworms, etc.
Class Hirudinea: leeches
Phylum Nemertea (ribbon worms)
Phylum Sipuncula (peanut worms)
Phylum Echiuria: (spoon worms)
Lophophorates
Phylum Brachiopoda (lamp shells)
Phylum Phoronida(horseshoe worms)
Phylum Bryozoa/Ectoprocta(bryozoans/moss animals)
Phylum Entoprocta: (goblet worms)
Domain Eukarya
Kingdom Metazoa
Superphylum Ecdysozoa
Phylum Arthropoda
Subphylum Chelicerata
Class Merostomata: horseshoe crabs
Class Pycnogonida: sea spiders
Class Arachnida: spiders, ticks, mites
Subphylum Crustacea
Class Branchiopoda: fairy shrimp, water
fleas, etc.
Class Maxillopoda: barnacles, copepods,
ostracods
Class Malacostraca: crabs, shrimp, lob-
sters, krill, amphipods, isopods
Subphylum Uniramia
Class Insecta: insects
Class Diplopoda: millipedes
Class Chilopoda: centipedes
Phylum Nematoda (roundworms)
Phylum Kinorhyncha
Phylum Loricifera
Superphylum Deuterostomia
Phylum Chaetognatha (arrow worms)
Phylum Echinodermata
Class Asteroidea: sea stars
Class Ophiuroidea: brittle stars
Class Echinoidea: sea urchins
Class Holothuroidea: sea cucumbers
Class Crinoidea: sea lillies, feather stars
Phylum Hemichordata (acorn worms)
Phylum Chordata
Subphylum Urochordata (tunicates)
Class Ascidiacea: sea squirts
Class Larvacea: larvaceans
Class Thaliacea: salps
Subphylum Cephalochordata (lanceletsam-
phioxus)
Subphylum Vertebrata (fishes, amphibians,
reptiles, birds, mammals)
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Marine Invertebratesthe lower phyla

I
n the previous lessons, we learned about the least
conspicuous, but by no means least significant marine
organisms - the diverse marine microbes and produc-
ers (most of whom are microbial). Moving up the food
chain from producers, we meet the consumers, the het-
erotrophs who rely directly or indirectly on the produc-
ers for all their energy and organic materials. We have
already met the smallest consumers, the heterotrophic
archaea and bacteria, and the protozoans. Now we are
ready to meet the more visible and familiar consum-
ers, the members of Kingdom Animalia, or Metazoa.
Animals (metazoans) are multicellular, eukaryotic hetero-
trophs that obtain food by ingestion. Of the 3 kingdoms of
multicellular eukaryotes (plants, animals, fungi), animals are
the most diverse. This may reflect the diverse ways they
have evolved to obtain food and to avoid becoming another
animals meal, being unable to manufacture their own food.
Figure 12 shows a taxonomic classification of the meta-
zoans. As you study the different animal groups, keep refer-
ring back to this figure to get the "big picture" of animal clas-
sification. Also refer to Figure 7.54 in your textbook, which
shows the phylogenetic relationships among the major
animal phyla. There is also a table at the end of each text-
book chapter, listing the animals covered and summarizing
their characteristics. These are useful tools for organizing
and reviewing the great diversity of animals. Keep in mind
the position that animals occupy on the Tree of Life shown
in Figure 11, and pictured on the first page of each chapter
in your textbook. Despite their great diversity, animals repre-
sent just one twig on the eukaryote branch, and eukaryotes
account for just one of the three main branches of life. For
a geologic perspective on the appearance of the different
animal groups (and life in general) on Earth, look at the
Geologic Time Scale in the Appendix of this Course Guide.
Traditionally, animals are divided into two groups: verte-
brates (with vertebrae, or a backbone) and inverte-
brates (without a backbone). This is quite a biased view,
considering that at least 97% of animal species are inverte-
brates. Vertebrates constitute just one subphylum, a group
within a larger phylum. Subphylum Vertebrata is one
of three subphyla within Phylum Chordata (chordates),
which is just one of the 3540 animal phyla currently identi-
fied. The other 2 chordate subphyla, plus all the other animal
phyla, are invertebrates. We will devote two lessons to the in-
vertebrates, surveying about half of the 3540 phyla (a cur-
sory introduction to this huge and diverse group). This lesson
describes some of the more primitive invertebrate phyla. In
the next lesson, we survey the more advanced invertebrate
phyla and, in Lessons 911, the vertebrates. Use the mate-
rial in this Course Guide as an introduction to these groups,
which are described in greater detail in your textbook. Re-
calling what you learned in Lesson 4, consider how each of
these highly successful groups has evolved unique adapta-
tions to fulfill the basic requirements of life - obtaining energy
and raw materials, responding to the external environment
and maintaining homeostasis, growing and reproducing.
All the invertebrate phyla have marine representatives, and
some are exclusively marine. In fact, most of the current ani-
mal phyla (plus some others, now extinct) were present in the
sea more than 500 mya, before any life appeared on land.
About 550 mya to 500 mya, during the Cambrian Period,
a huge burst of evolutionary innovation took place in the an-
cient ocean. Known as the Cambrian Explosion, this great
proliferation of life produced most of the present-day animal
phyla. An explosion of eukaryotic life was made possible in
part by the activity of photosynthetic marine prokaryotes,
during hundreds of millions of years, adding enough oxygen
to the environment to support the great metabolic needs of
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large multicellular organisms. New forms have continued to
evolve within these phyla, but they maintain the basic body
plans laid down during the Cambrian Period. It is primarily
on the basis of these body plans, and how they unfold dur-
ing embryonic development, that zoologists (scientists
who study animals) traditionally have classified the Metazoa.
Parazoa
Phylum Porifera (sponges): Sponges are among the
most primitive and unusual animals. They are members
of Subkingdom Parazoa (near animals), distinct from
all other metazoans who belong to Subkingdom Eumeta-
zoa (true animals). Sponges are basically porous sacs
(hence the phylum name "Porifera") with a network of canals
through which water is pumped and filtered. Various spe-
cialized cells cover the surface of the sac, form the pores,
create the water currents, transport, store and digest the
filtered food particles, and secrete materials for structural
support. Parazoa have a cellular level of organization,
in which individual cells function without being organized
into tissues and organs. There is no mouth or digestive
tract, no sense organs, nerve cells or muscles. Digestion
is entirely intracellular (takes place inside the cell) and
the exchange of respiratory gases and wastes takes place
through the cell membrane. Reproduction may be asexual,
by budding, or sexual, in which case the embryo begins de-
velopment inside the parent and is released as a larva which
drifts and settles to the bottom, where it continues to grow.
Sponges are motionless and live a sessile life, permanently
attached to the sea floor or other substrates. In fact, they
were long believed to be plants and were not formally rec-
ognized as animals until 1825. They are almost exclusively
marine, and are classified according to the type of mate-
rial used for structural support, including calcium carbonate
(calcareous sponges), silicon dioxide (glass sponges),
fibers of the protein spongin (demosponges) or a combi-
nation of these materials (sclerosponges).
Radiata
The most primitive members of the Subkingdom Eumetazoa
are radially symmetrical animals, the Radiata. More ad-
vanced, bilaterally symmetrical animals are Bilateria.
Radial symmetry and asymmetry (as in sponges) are char-
acteristic of relatively inactive, sessile or floating animals,
whereas bilateral symmetry allows animals to be more ac-
tive. Unlike the sponges, Radiata display a tissue level of
organization, where cells are organized into tissues, layers
of cells that perform as functional units. In the embryos of
Radiata, there are two tissue layers, the endoderm and ec-
toderm, and a middle layer of gelatinous material with few
or no cells, called the mesoglea. During development, more
specialized tissues grow from the endoderm and ectoderm
cells. Another advancement made by the Radiata is the pres-
ence of a gut cavity, called a coelenteron, with a mouth
but no anus. The body plan of these animals is more complex
than in sponges and less complex than in bilaterians. Radiata
are represented by two phyla, Cnidaria and Ctenophora.
Phylum Cnidaria (sea anemones, jellyfishes and cor-
als): These are some of the most beautiful and important
of all the animals. They range from tiny translucent forms, to
the enormous jellyfish, Cyanea arctica, which weighs almost
a ton and may have tentacles 100 feet long. They include
flower-like sea anemones and huge aggregations of tiny cor-
al animals, which form the coral reefs. Two body forms occur,
the free-swimming medusa or jellyfish form, and the sessile
polyp or hydroid form. Both have whorls of tentacles sur-
rounding the mouth, which is oriented upward in the polyp
and downward in the medusa. Cnidaria are carnivores, but
without well-developed muscles, nerves and sense organs,
they cannot pursue their prey. Instead, they trap food that
comes to them, using their sticky tentacles, equipped with
stinging cells called nematocysts. They can capture and
immobilize prey as large as themselves, and much faster. The
coelenteron functions as a gastrovascular cavity, where
both digestion and circulation take place. Partial extracel-
lular digestion by enzymes takes place in the gut, and final
digestion and absorption take place intracellularly. Cnidar-
ians also have a primitive nervous system called a nerve
net, with protoneurons (primitive nerve cells) and various
sensory cells, sometimes assembled into sense organs.
They reproduce asexually and sexually, and many have com-
plex life histories, including both polyp and medusa stages
and colonial forms. There are about 10,000 known spe-
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cies, almost all marine, which are classified as hydrozoans,
scyphozoans, cubozoans and anthozoans.
Phylum Ctenophora (comb jellies): Comb jellies appear
similar to Cnidarian medusae (they have no polyp stage), but
tentacles are lacking or are not in whorls around the mouth.
They are distinguished by eight radially-arranged comb
rows composed of ciliated plates (combs or ctenes)
which are used for swimming. They lack nematocysts, but
their tentacles, when present, are equipped with adhesive
cells called colloblasts. Food is caught by surface mucus
and carried to the mouth by cilia, or by the sticky tentacles
being wiped across the mouth. The gastrovascular cavity is
somewhat more advanced than in Cnidaria, and a unique
aboral sense organ is used for balance. There are only
around 100 known ctenophore species, but they are vora-
cious predators and can occur in numbers large enough to
be ecologically important. They are exclusively marine, and
are classified as Tentaculata, with tentacles, or Nuda, with-
out tentacles.
BilateriaAcoelomates and Pseudocoe-
lomates
The advent of bilateral symmetry was crucial in the course
of animal evolution. This arrangement allowed animals to
have not only top (dorsal) and bottom (ventral) surfaces
(like the oral and aboral surfaces in Radiata) but also front
(anterior) and rear (posterior) ends, as well as left and
right sides. Movement could be directed forward (anteriorly)
and became more efficient, suitable to more active lifestyles.
Increased activity involved the development of muscles, more
specialized sense organs, and a more complex nervous sys-
tem. At the same time, a head developed (cephalization),
with the mouth, sense organs and nerve center at the forward
end. As locomotor, sensory and nervous systems developed,
more efficient respiration, circulation, digestion and excretion
were required. Bilateria demonstrate the most complex level
of organization, where cells are organized into tissues, and
tissues are organized into organs and organ systems.
Two other important advances occurred in Bilateria. The first
was the appearance of the mesoderm, a third layer of cells
between the ectoderm and endoderm in the developing em-
bryo, instead of the gelatinous mesoglea seen in Radiata.
The mesodermal cells form muscles, the circulatory system,
most organs outside of the digestive tract, and (in verte-
brates) the skeletal system. The endodermal cells form the
digestive tract, while the ectodermal cells become the skin
and nervous system. Having a third embryonic layer of cells
also made possible the second great innovation of bilaterian
evolution: a coelom. This is a second body cavity, located
between the gut cavity and the body wall. The coelom forms
when the mesoderm splits into two layers and creates a fluid-
filled cavity. Organs are suspended in the cavity, cushioned
and protected by the fluid. In soft-bodied animals, the coe-
lomic fluid also acts as a hydrostatic skeleton, providing
support and aiding in locomotion. The development of the
coelom was essential to the evolution of large, complex ani-
mals.
Traditionally, the Bilateria have been divided into three
groups: acoelomates (without a coelom), pseudocoe-
lomates (with a false cavity) and coelomates (with a
true coelom). Zoologists used these morphological char-
acteristics to help classify animals and understand their phy-
logenetic relationships. Once again, molecular sequenc-
ing techniques are shedding more light on the subject, and
it appears that these divisions are not quite so clear-cut. We
will use them here, to divide the many invertebrate groups,
somewhat arbitrarily, into "lower" phyla (acoelomates and
pseudocoelomates), covered in this lesson, and "higher"
phyla (coelomates), covered in the next.
Phylum Platyhelminthes (flatworms): Platyhelminthes
(flat parasitic worms) have long been considered the sim-
plest animals having bilateral symmetry and an embryo with
three cell layers. The mesoderm does not split to form a coe-
lom, but does form muscle and some organs. Cephalization
is present in flatworms, and they have a brain, eye-spots,
chemoreceptive organs, and some have organs for
grasping prey. Their metabolic rate is about 10 times higher
than in Radiata and they are more active. Mostly parasites,
their digestive, respiratory, and excretory systems are primi-
tive, while their reproductive organs are quite complex. Their
hosts supply all their materials, and their small size and flat-
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tened shape increases their surface-to-volume ratio, so re-
spiratory gases, nutrients and wastes are easily exchanged.
About 20,000 species of Platyhelminthes are known, in-
cluding cestodes (segmented tapeworms parasitizing the
intestines of most vertebrates), trematodes (leaf-shaped
flukes, also common parasites of vertebrates), and turbel-
larians (predominantly marine, free-living carnivores, some
found in or on other animals). Many zoologists now place the
flatworms in the superphylum Platyzoa (Fig. 12), along with
some small phyla described at the end of this lesson.
Phylum Nemertea (ribbon worms): There are only
about 900 known species of ribbon worms, mostly marine,
but they are abundant in shallow temperate waters, living
among the rocks and algae or in mucus tubes in the mud
and sand. They range in size from just a few millimeters to
30 m (nearly 100 ft) in the case of Lineus longissimus,
the longest invertebrate. They have more advanced features
than the flatworms, including an enclosed circulatory sys-
tem, sometimes containing the respiratory pigment hemo-
globin, a complete digestive system with both a mouth and
anus, and more developed muscular and nervous systems.
They are carnivores and have a unique muscular tube called
a proboscis, used to capture prey. This organ is explosively
everted from its cavity, turning it inside out and exposing a
sticky surface that ensnares the prey. In some types, the pro-
boscis is also armed with a "stylet" that wounds the prey and
secretes a toxin. The proboscis then wraps around the prey
and retracts, pulling it into the mouth. The cavity housing the
proboscis, the rhynchocoel, is similar to a coelom. Recent
studies place Nemertea with the Trochozoa, a subgroup of
the superphylum Lophotrochozoa (Fig. 12), whose other
members are discussed in the next lesson.
Phylum Nematoda (roundworms): These mostly small,
seldom seen animals occur in incredible numbers, and by
some estimates are the most widely distributed and abun-
dant animals on Earth, accounting for perhaps 4 out of every
5 animals. They are found everywhere, mostly in soils and
sediments where they can occur in densities of up to sev-
eral billion individuals per acre, feeding on bacteria and dead
organic matter. Many roundworms parasitize plants and
animals, and most groups of marine organisms are hosts to
these parasites, including a 13-m long nematode that para-
sitizes sperm whales. Between 10,000 and 25,000 species
are known, but estimates of their diversity, from molecular
evidence, range from 40,000 to 100 million species! If this
were true, their diversity, as well as their numbers, would be
almost unparalleled - second only to (or even ahead of) the
arthropods, among animals.
This enormously successful and diverse group has a decep-
tively uniform and simple body plan. Nematodes (thread-
like") are slender and circular in cross section, tapering at
both ends. Sense organs and a brain are concentrated ante-
riorly, near the mouth which opens into a muscular pharynx,
where food is pulled in and crushed. They have digestive,
excretory, nervous and reproductive organ systems, and
an incompletely lined pseudocoelom. The muscles all run
longitudinally, so nematodes move by thrashing. Their epi-
dermis (skin) is covered by a tough, elastic cuticle which
is periodically shed as the animal grows, a process called
molting. Based on these features, and molecular evidence,
nematodes have recently been placed in the superphylum
Ecdysozoa (molting animals). Ecdysozoa also includes
the hugely diverse arthropods (insects, crabs, etc.), whom
we will meet in the next lesson. Notice that the three phyla
of worm-like animals we have just described (Platyhelmin-
thes, Nemertea and Nematoda), although superficially simi-
lar, are now placed in three different superphyla (Platyzoa,
Lophotrochozoa and Ecdysozoa), based on current morpho-
logical and molecular evidence (Fig. 12).
Phyla Gnathostomulida, Cycliophora, Gastrotricha,
and Rotifera: These tiny, bizarre-looking creatures are
referred to as meiofauna (diminutive animals ), or in-
terstitial animals, which live in the equally tiny spaces be-
tween sediment particles (except Cycliophora, which inhabits
the hairs around a lobsters mouth). Gastrotricha are acoe-
lomate and the others have a pseudocoel. They may be in-
cluded (along with Platyhelminthes) in Platyzoa. Meiofauna
are described in your textbook (pp. 128 and 295).
Phyla Kinorhyncha and Loricifera: Also members of
the meiofauna, these little-known pseudocoelomates have
recently been classified with nematodes and arthropods, in
Ecdysozoa.
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your textbook.
Metazoa, Parazoa and Eumetazoa
vertebrate and invertebrate
Radiata and Bilateria
acoelomate, pseudocoelomate and coelomate
cellular level of organization and tissue level
of organization
radial symmetry and bilateral symmetry
oral and aboral, ventral and dorsal, anterior
and posterior
cephalization
endoderm, ectoderm, mesoderm and meso-
glea
gut/digestive cavity and gastrovascular cavity
coelenteron, coelom and rhynchocoel
hydrostatic skeleton
epidermis, gastrodermis and cuticle
intracellular digestion and extracellular diges-
tion
deposit feeder, suspension feeder, filter
feeder
molting and metamorphosis
asexual and sexual reproduction, hermaphro-
dite and broadcast spawning
sessile
osculum, ostia, collar cells and spicules
hydroid (polyp) and medusa
cnidocyte, nematocyst, statocyst, aboral
sense organ
planula
comb rows/ciliary combs, ctenes and collo-
blasts
protoneuron and nerve net, neuron and brain,
central nervous system
eye-spot
hemoglobin
proboscis
pharynx
meiofauna/interstitial animals
Cambrian Period, Cambrian Explosion
glossary terms
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1. Define, by comparing and contrasting, the following terms:
a. vertebrate and invertebrate
b. parazoa and eumetazoa
c. cellular and tissue levels of organization
d. intracellular and extracellular digestion
e. radial and bilateral symmetry
f. mesoglea and mesoderm
g. coelenteron and coelom
h. nerve net and brain
i. hydroid and medusa
j. nematocyst and statocyst
k. filter feeder and deposit feeder
2. Phylum Cnidaria is represented by four classes. Name these classes and, for each class, describe some highlights of
their life histories and some of the animals included.
3. Describe several advances that accompanied the evolution of bilateral symmetry in animals.
4. Many "lower" invertebrates are worm-like creatures. Describe the basic characteristics of one of the worm-like phyla
included in this lesson.
5. Who are the meiofauna and where do they live? Include several phyla that occur as meiofauna, in your description.
6. Nematodes (roundworms) may be the most abundant animals on Earth. Which of the following animals are their
closest relative: flatworms, ribbon worms, insects, you?
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Marine
Invertebrates
The Higher
phyla
In this lesson, we continue our survey of the invertebrates, covering the "higher"
invertebrates - those with a true coelom.
1. Be familiar with the various groups of "higher" invertebrates, and their characteristic
features.
2. Be aware of some of the major trends in the evolution of these groups, and their
phylogenetic relationships.
2. Complete the Reading Assignment (below) in C&H (Castro & Huber, Marine Biol-
ogy 9th Ed.).
Reading assignment
C&H: Chapter 7 (pp. 115-149)
8
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Marine Invertebratesthe higher phyla
W
e continue our survey of marine invertebrates,
covering the generally more advanced phyla,
those with a true coelom. They fall into three
superphyla, based on morphological and molecular evidence:
Lophotrochozoa, Ecdysozoa and Deuterostomia (Fig.
12). We have already met the acoelomate and pseudocoe-
lomate members of Lophotrochozoa and Ecdysozoa, in Les-
son 7. The third superphylum, Deuterostomia (which includes
us), is named for the particular way the body begins to take
shape in the embryo. Traditionally, zoologist have recognized
two evolutionary paths: protostomia (first mouth) and
deuterostomia (second mouth). The embryo starts as a
ball of cells. As the cells divide and multiply, a dent appears
in the ball, into which cells begin migrating, forming a second
layer of cells inside the ball. Most animals are protostomes,
with the mouth eventually forming from this initial indenta-
tion. In deuterostomes, the first indentation becomes the
anus and the mouth forms later, from a second indentation.
Again, refer back to the animal classification scheme in Fig.
12 of this Course Guide for the big picture (and the Geologic
Timescale in the Appendix for the really big picture). Use Ta-
ble 7.1 in your textbook to review the characteristics of the
major animal phyla. Also look at Fig. 7.54, a family tree show-
ing the phylogenetic relationships of major phyla. As we
will see, our understanding of animal taxonomy and phylo-
genetic relationships is still very much a work in progress.
LophotrochozoaLophophorates
The superphylum Lophotrochozoa includes groups of phy-
la, the lophophorates and the trochozoans. We will con-
sider the lophophorates first. This group is named for a unique
structure called a lophophore, a crown of ciliated tentacles
used for respiration and to capture suspended food particles.
All the phyla in this group share this feature. Lophophorates
also have a coelom divided into two or three compartments,
some form of protective sheathing, and most have a com-
plete U-shaped digestive tract, with the mouth and anus
opening near the lophophore. Most other organ systems are
simple. They are represented by four phyla: Bryozoa (or Ec-
toprocta), Entoprocta, Phoronida, and Brachiopoda.
Phylum Ectoprocta/Bryozoa (moss animals): With
about 5,000 living species, and many more fossil forms
identified, bryozoans have a long record of success. They
are abundant in marine habitats throughout the world. Also
called "moss animals," many look somewhat like cnidarian sea
fans, with branching colonies composed of tiny individuals
called zooids. A sheath of tissue encloses each zooid and
secretes a gelatinous or rigid calcium carbonate structure,
forming a compartment from which the lophophore can be
extended. Different types of zooids within the colony per-
form different functions, including food gathering (autozo-
oids), egg production (heterozooids), and strengthening
or cleaning the structure. The zooids are usually connected
to each other by a thin strand of tissue. Bryozoan colonies
may be encrusting, or upright with solid, branching or cork-
screw shapes. A few are free-floating, or creeping. Their
U-shaped gut is arranged such that the mouth lies inside,
and the anus just outside, the lophophores ring of tenta-
cleshence the phylum name, Ectoprocta (outside anus).
Phylum Entoprocta (goblet worms): These tiny animals
have been classified as Bryozoa, but now have their own phy-
lum. As their name ("inside anus") suggests, they differ from
bryozoans in that their U-shaped digestive tract is arranged
with the anus is inside the ring of tentacles. There are only
about 150 species, mostly marine. Nearly all are sessile, at-
taching to the substrate by a stalk-like continuation of the body
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wall, topped by a cup-shaped ("goblet") body. Some species
are colonial, with multiple individuals on branching stalks.
Phylum Phoronida (horseshoe worms): Only about 20
living species are known, but they occur throughout the world
except polar areas, and may have been more prevalent in the
past. Phoronids are slender and worm-like, from a few mil-
limeters to about a foot long. They burrow into the sediment
or dissolve holes into rocks, shells and even cement piers,
and then line the hole with a tube they secrete. They tend
to aggregate, the tubes twisting around each other. The
name "horseshoe worm" comes from the coiling horseshoe
shape in which the lophophore tentacles are often arranged.
Phylum Brachiopoda (lamp shells): The approximately
350 living species of brachiopods appear to be the vestiges
of a once widespread and abundant marine group, with more
than 30,000 fossil species identified. One genus, Lingula,
includes the oldest (most evolutionarily unchanged) animals
known. Brachiopods are small, up to about 3 in, and are pro-
tected by a shell with two halves, or valves, like the unrelated
clams. Articulate brachiopods have a hinge-like articulation
(joint) between the valves, whereas in inarticulate forms
the valves are held together only by muscles. The shells of
some forms resemble a type of ancient oil lamp, hence the
common name, lamp shells. Most are attached to a sub-
strate by means of a fleshy stalk, or pedicle. They prefer
cold water environments, near the poles or in the deep sea.
LophotrochozoaTrochozoans
Trochozoa, the other lophotrochozoan group, includes two
large phyla, Annelida and Mollusca, plus several smaller
phyla, including Nemertea (ribbon worms), whom we met
in the previous lesson. As adults, trochozoans may appear
completely unrelated (e.g. earthworms and octopuses) but
their larvae are nearly identical. Trochophore larvae are
biconical (like two cones placed base to base), with bands
of cilia around the middle, a topknot of flagellae and a U-
shaped gut. Trochozoans are a good illustration of the Re-
capitulation Principle, or ontogeny recapitulates phylog-
eny. In other words, embryonic development (ontogeny) is
a condensed version of the evolutionary history (phylog-
eny) of an organism, and species with similar ontogenies
probably have a common ancestry. Until recently, annelids
were considered more closely related to arthropods, partly
because their adult forms are both segmented. But anne-
lids have trochophore larvae, and arthropods don't. Using
the recapitulation principle, and other morphological and
molecular sequencing evidence, most zoologists now clas-
sify annelids and molluscs as trochozoans, in Superphylum
Lophotrochozoa, and place arthropods in Superphylum
Ecdysozoa. Similar evidence places the ribbon worms (Ne-
mertea), from the previous lesson, in the trochozoan group.
Phylum Annelida (segmented worms): This familiar
phylum, including earthworms and leeches, is important for
several reasons. A large group, with perhaps 20,000 known
species, annelids are ecologically important members of
soil and marine sediment communities. They are also one
of the most studied groups, as models of developing com-
plexity in animal evolution. The body plan of annelids has a
metameric (segmented) organization, with a distinct and
complex head, and a body divided along its length into many
compartments. Each segment contains a fluid-filled coe-
lom. The digestive tract and organ systems are relatively
well-developed, including a closed circulatory system
with blood vessels, hearts (one per segment), blood and
respiratory pigment, usually hemoglobin. The body wall
is covered by the epidermis, and usually by an additional
outer covering, the cuticle. These protective layers enclose
two muscle layers, an outer circular layer and inner longi-
tudinal layer in each segment. This arrangement of mus-
cles, and a fluid-filled coelom to exert hydrostatic pres-
sure, make annelids powerful swimmers and burrowers.
Traditionally, three major classes of annnelids have been
recognized: Oligochaeta (earthworms and their kin),
Hirudinia (leeches) and Polychaeta. Polychaetes are
the largest group and almost all marine, whereas there
are few marine oligochaetes and leeches. Paired arm-like
muscular extensions of the body wall on each segment,
called parapodia, are characteristic of polychaetes. The
parapodia have skeletal supports, bristles called se-
tae, tactile sense organs and gills, and are used for
locomotion. The setae give the group their common
name, bristleworms (polychaeta means many hairs").
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Polychaetes are extremely abundant and diverse, living in
many different marine habitats, with many different lifestyles.
Two unique groups are the bizzare and beautiful tube-dwell-
ing worms (tubeworms), the pogonophorans (beard
worms - for their long tentacles) and vestimentiferans
("garment" worms). Completely unknown before 1900, they
live in the deep sea. Pogonophorans were not recognized
as annelids until 1964, when the first intact specimens were
recovered. They can be as long as 7 ft and the only seg-
mented part of their body is the posterior part (called the
opisthosoma), which is buried in the mud and had been
torn off of previously collected specimens. We still have a
lot to learn about these tubeworms, which appear to be
the dominant macrofauna of the deep-sea, living in sedi-
ments, around hydrothermal vents, at methane seeps and
even at whale carcasses on the sea floor. Riftia, a genus
of giant tubeworms living around hydrothermal vents, are a
spectacular example (C&H Fig. 16.28, p. 380). Beard worms
are the only free-living animals with no mouth or digestive
tract in the adult. Some nutrients are absorbed through
the tentacles, but most appear to be provided by chemo-
synthetic bacteria living inside the worm in a special-
ized organ called a trophosome, which develops from the
embryonic gut - a great example of obligate symbiosis.
Pogonophorans and vestimentiferans are often placed in
their own phyla, but recent studies confirm them as anne-
lids, in the family Siboglinidae. In fact, current morpho-
logical and molecular research suggests that the phyla
Echiura and Sipuncula (described below) should also be
classified as annelids, and that the other annelid classes,
oligochaetes and leeches, may be derived from poly-
chaetes. Taxonomy is certainly a scientific work in progress!
Phylum Echiura (spoon worms): This small group (about
150 known species) are all marine organisms. They have a
plump, unsegmented sausage-shaped body, with a grooved
proboscis extending beyond the mouth, and small hooks at
the posterior end of the body, giving the phylum its name,
which means spine-tails. The proboscis may be spoon-
shaped (hence their common name) or forked, and can
be greatly extended to be used in sensation and feeding.
Echiurans have relatively simple organ systems, and most
are sedentary, living in burrows and crevices, or the shells of
other animals, and feeding on detritus. One species common
in mudflats of the Pacific coast of California, Urechis caupo,
lives in a U-shaped burrow and uses the proboscis to se-
crete a mucus net for trapping plankton. Commonly called the
innkeeper worm, its burrow may be shared by commen-
sal organisms, such as small crabs, polychaetes and fish.
Phylum Sipuncula (peanut worms): Like Echiura,
this is a small group (320 known species, all marine)
of small, relatively simple, unsegmented worms living
in burrows, crevices or other animals shells, and feed-
ing on detritus. The body is divided into an anterior part,
the introvert, with a mouth and tentacles, and a plump
trunk. When the introvert is retracted into the trunk
by retractor muscles, the animal resembles a peanut.
Phylum Mollusca (snails, clams, octopuses, etc.): With
as many as 200,000 known species, this phylum is second
only to arthropods (and perhaps nematodes) in worldwide
diversity, and is the most diverse in marine environments
(again, with the possible exception of nematodes). The
major groups of molluscs were established even before the
beginning of the Cambrian Period, and they have pursued
individual evolutionary paths ever since. However, all mol-
luscs still have basic features in common. We may have a
more complete picture of this group than of any other in-
vertebrate phylum. They are often conspicuous creatures,
whose shells have long been admired, collected and used
by people, and they have been extensively studied and
conserved because of their importance as food sources.
Many specific features distinguish molluscs from other tro-
chozoans. They have an unsegmented soft body (mol-
lusc is derived from a word meaning soft) covered by a
layer of tissue called the mantle. The mantle secretes a
calcium carbonate shell into which the animal can with-
draw by means of a retractor muscle. Respiration occurs
in the mantle cavity, usually through gills. Their well-
developed head has specialized sense organs, and their
coelom is reduced by a thick, muscular body wall which is
developed on the ventral (belly) side as a muscular foot
used in locomotion. The digestive system is complex, with
a radula (a hard, toothed instrument for rasping food) in
the mouth. They have an open circulatory system with a
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2-chambered heart and the respiratory pigment hemo-
cyanin (giving the blood a greenish color) or hemoglobin.
There are one or two kidneys, a nervous system, usually
with nerve ganglia, nerve cords, and an extensive nerve
network. Variations of these features are seen in the three
major classes of molluscs, Gastropoda (snails, limpets,
abalones and nudibranchs), Bivalvia (clams, oysters and
mussels) and Cephalopoda (squids, cuttlefishes, octo-
puses and Nautilus), which are described in your textbook.
Ecdysozoa
Superphylum Ecdysozoa is a taxonomic level created
quite recently to group together several phyla thought to
have common ancestry, based on morphological and mo-
lecular sequencing evidence. It comprises the animals that
shed their exoskeleton (ecdysis = molting). These
animals have a three-layered cuticle, forming an exoskel-
eton composed of organic material (usually chitin and
other substances), which is shed periodically as the animal
grows. This tends to limit the size of the animal, but also
opens other developmental options. In animals with mineral
skeletons, like molluscs and vertebrates, growth occurs
only as more mineral is added, limiting the animals form,
whereas molting makes it possible for the animal to dramati-
cally alter its form. Insects, in particular, have exploited this
opportunity by undergoing metamorphosis of the larval
forms into very different adult forms. Ecdysozoa include
two major phyla, Arthropoda (insects, crustaceans, spi-
ders and relatives) and Nematoda, the roundworms, plus
several minor phyla, including Loricifera and Kinorhyn-
cha (Fig. 12). We covered the last three phyla in Lesson 7.
Phylum Arthropoda (insects, crustaceans, spiders, etc.):
More species have been identified in this phylum than in any
other. The vast majority are insects, with about 800,000
known species, and probably many more undiscovered (es-
timates range into the tens of millions!). In marine environ-
ments, however, insects are rare. Like the angiosperms of
the plant kingdom, insects are relative late-comers which
diversified explosively on land. The living arthropods may
be divided into several subphyla. The largest, Uniramia
(unbranched appendages) includes insects, millipedes and
centipedes, and has few marine representatives. The wa-
ter strider Halobates, which skates on the surface, is the
only insect in the open ocean (C&H Fig. 15.17, p. 344).
The second, Crustacea, has a huge variety of marine
forms, including the familiar lobsters, shrimp, crabs and
barnacles. A third group, Chelicerata, includes spiders
and scorpions and some marine forms, the horseshoe
crabs and sea spiders. A fourth group, the trilobites,
and possibly many other arthropod lineages, are extinct.
In addition to their exoskeleton, arthropods have a seg-
mented body and were previously thought to be related
to the segmented worms, the annelids. In primitive forms,
each segment was equipped with a pair of jointed ap-
pendages (arthropoda means jointed foot). In living
arthropods, many of these appendages are highly modi-
fied or lost, and segments are fused into head, thorax and
abdomen regions. Movement of the appendages is con-
trolled by a complex muscular system, including smooth
and striated muscles, like ours. The coelom is reduced,
and most of the body cavity is an open hemocoel where
the tissues are bathed directly by blood, in an open cir-
culatory system. Respiration is by tracheae, or gills in
marine forms. The nervous system is similar to an anne-
lids, but with a more complex brain. Most have single and
compound eyes. Fertilization is internal in most groups.
Subphylum Crustacea are the predominant marine rep-
resentatives of the arthropods, with approximately 68,000
species identified, which may only be a fraction of their true
diversity. In most crustaceans, the exoskeleton is hard-
ened by the addition of calcium carbonate and fused in
the head and thorax region, forming a shield called a cara-
pace (crustacean means shelled ones). Unlike insects,
crustaceans have biramous appendages, branching into
inner and outer arms. Their appendages include two sets of
antennae (one more than insects), a set of mandibles
and two or three sets of maxillae or maxillipeds on the
head, several pairs of walking legs (pereopods) or other
types of specialized appendages on the thorax, and usually
reduced appendages on the abdomen. These appendages
are specialized to perform a variety of functions, including
food capture and manipulation, chewing, swimming, walking,
breathing, copulation, egg brooding, and sensory reception.
Crustaceans have a special larval form called the nauplius.
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This subphylum is divided into about six classes. By far
the largest, Malacostraca, includes the familiar (and deli-
cious) Order Decapoda ("ten-footed") - lobsters, shrimp
and crabs. Smaller, inconspicuous malacostracans are ubiq-
uitous and important, e.g., amphipods, isopods and krill,
which are extremely abundant as zooplankton in polar seas
and the exclusive food source for some whales, birds and
fishes. A second class, Maxillopoda, includes copepods,
tiny crustaceans which are so common among the zoo-
plankton that they also are contenders for the title of most
abundant animals on Earth. They are the grazers of the sea,
feeding on phytoplankton and forming a vital link between
the producers and animals at higher trophic levels, as we
will see later in this course. Barnacles also are included
in this class. Shelled forms that resemble bivalves, they live
sessile lives attached head-first to the substrate. A third
class, Branchiopoda, includes fairy shrimp and water fleas.
Subphylum Chelicerata is the third group of living ar-
thropods, in addition to Uniramia and Crustacea. They are
named for their pointed appendages, called chelicerae,
which are used to grasp food, instead of the chewing man-
dibles used by most other arthropods. Most chelicerates are
unable to ingest solid food and instead drink blood or inject
digestive enzymes into their prey. This group is mostly com-
posed of arachnids, the terrestrial spiders, scorpions and
mites. Many marine chelicerates dating back to the Cambrian
are now extinct, with horseshoe crabs and sea spiders
the only remaining marine representatives of this group.
Deuterostomia
The groups we have described so far are protostomes, in
which the mouth develops from the first in-pocketing of the
embryo. The remaining animal phyla are deuterostomes, in
which the embryonic mouth forms from a second indentation.
Grouped into the Superphylum Deuterostomia, these
phyla include Chaetognatha (arrow worms), Echinoder-
mata (sea stars, sea urchins, etc.), Hemichordata (acorn
worms), and Chordata (vertebrates and their relatives).
Phylum Chaetognatha (arrow worms): It is not re-
ally clear how these strange creatures should be classified.
Small, torpedo-shaped and nearly transparent, they dart
about swiftly, like arrows. They have side and tail fins, well-
developed muscles and nervous system, and compound
eyes. Voracious marine predators, they have teeth and
spines extending from the head, which snap shut to cap-
ture prey and are covered by a hood when not in use. They
have no respiratory, excretory, or circulatory organs, and
move materials through the body cavity by means of cilia,
and exchange gases and wastes directly through the body
wall. There are only about 100 species, but they can oc-
cur in staggering numbers, especially in warm shallow seas.
Phylum Echinodermata (sea stars, sea urchins, sea
cucumbers, etc.): Although radially symmetrical as adults,
echinoderms have bilaterally symmetrical larvae, which fol-
low the deuterostome path of embryonic development.
They are the only animals to make this switch from pri-
mary bilateral symmetry to secondary radial sym-
metry in the adult. Some echinoderms even switch again,
to a tertiary bilateral symmetry. This is an old and suc-
cessful phylum, with about 7,000 living species, and tens
of thousands of fossil species described, all marine. They
were well established near the beginning of the Cambrian
Period and have developed many unique features during the
past 550600 million years. Although there are fewer echi-
noderms today than in the past, they are still widespread
and can be extremely abundant in some areas. In parts of
the Bering Sea, for example, there are more sea stars and
heart urchins than any other organism of comparable size.
Echinoderms are pentaradial (or pentamerous) - they
have a five-fold organization of the skeleton and most organ
systems, which can be seen in the five arms of a starfish
or the five petals on a sand dollar. Their unique hydraulic
system, the water-vascular system, has extensions called
tube feet which are used for movement, respiration and
sensation. They bear some similarity to chordates, in how
the mesoderm and coelom develop, and in their calcium
carbonate endoskeleton. But the echinoderm coelom
is uniquely divided into complex partitions, and the endo-
skeleton is made of tiny ossicles with a unique sponge-like
microstructure, which are modified into various plates and
spines (echinodermata means spiny-skinned). Like the
more primitive Radiata, they are unsegmented and lack
a head, and they have an oral surface where the mouth
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lies, and an opposite aboral surface. Most organ systems
are simple, and excretory organs are absent. Typically, the
larvae swim about as part of the pelagic zooplankton, and
then settle into a benthic lifestyle as adults. This combi-
nation of primitive, advanced and unique features makes
echinoderms one of the most distinctive animal phyla. They
include five classes - the Asteroidea, Ophiuroidea, Ech-
inoidea, Holothuroidea and Crinoidea - each distinc-
tive in its own right, which are described in your textbook.
Phylum Hemichordata (acorn worms and relatives):
This small phylum of worm-like creatures is infrequently
seen living in tubes in the mud, or among rocks and sea-
weed. It is of interest primarily as a possible link between
our own group, the chordates, and other animal phyla such
as echinoderms. Hemichordates (half chordates) share
some, but not all, of the chordate features. Their coelom
develops in a way seen only in deuterostomes, and is di-
vided into three regions, as in vertebrates. They have gill
slits (some fossil echinoderms also show signs of gill slits),
a structure reminiscent of the nerve cord of chordates,
and a rudimentary structure called a stomochord, which
somewhat resembles the notochord of chordates. The
larvae of some hemichordates, and the results of some
DNA studies, also show a close affinity to the echinoderms.
Phylum Chordata (tunicates, lancelets and ver-
tebrates): Chordates are distinguished from
other animals by four features, each of which is
present at some time during their development:
gill slits (or pharyngeal slits) - a series of open-
ings connecting the inside of the throat to the ex-
terior, often used as gills and for filter feeding.
a dorsal, hollow nerve cord - a bundle of nerve fibers,
often enlarged at the forward end to form the brain, and
running along the back with pairs of nerves branching off at
intervals, connecting the brain to the muscles and organs.
a notochord - a flexible cartilaginous rod ly-
ing underneath, and supporting, the nerve cord.
a post-anal tail - an extension of the body wall mus-
culature, nerve cord and notochord, behind the anus.
Three subphyla share all of these traits: Urochordata,
Cephalochordata and Vertebrata. In vertebrates, the
embryonic notochord becomes surrounded and replaced
by a backbone, a column of articulating bones or cartilage
(vertebrae). They are the subjects of the next three lessons.
Subphylum Urochordata (tunicates): Appearanc-
es are deceiving in the case of urochordates, or tuni-
cates. They include about 3,000 marine species of sea
squirts, larvaceans and salps. At some point in their
life history, they are enclosed in a sac called a tunic.
As adults, sea squirts are sessile filter-feeders, simple
animals resembling sponges. Enclosed in their slimy tu-
nics, they have an incurrent siphon where water en-
ters, and an excurrent siphon where it exits (and will
squirt out if they are squeezed!). Their larvae, however,
are tadpole-like animals with a strong, muscular tail for
swimming and a clear notochord, nerve cord and gill
slits. The name Urochordata (uro = tail) comes from the
fact that the notochord is present only in the tail. The lar-
vae soon settle on the sea bed, attaching head-first and
undergoing their dramatic metamorphosis into the sim-
ple adult form. The notochord and tail are absorbed, and
the gill slits become the openings in the sieve-like tunic.
Tiny, transparent tunicates, called Larvaceans, retain the
pelagic larval form. They secrete a gelatinous house or
net, sometimes more than ten times their body size, for filter-
ing out food particles. When the filter becomes clogged, they
can escape through a door. New nets are secreted rapidly
and frequently, every few hours in some species. Salps also
are planktonic tunicates, with barrel-shaped bodies within
transparent tunics. Some are colonial, including Pyrosoma
spinosum, sometimes seen in the warm waters off New
Zealand in brilliantly luminescent, sock-shaped colonies up
to 60 feet long! Tunicates are widespread and often abun-
dant, and have been much studied for the insights they can
provide into the ancestry of higher chordates, like ourselves.
Subphylum Cephalochordata (lancelets): This small
group, with only about two dozen species, also is impor-
tant primarily for clues to understanding the development
and evolution of higher chordates. Small eel-like animals,
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they usually lie half buried in shallow, sandy environments
with the head poking up, filter feeding through their gill
slits. The other chordate features also are well devel-
oped throughout life. The name Cephalochordata (cephalo
= head) comes from the fact that the notochord extends
into the head. But, unlike vertebrates, their brain is tiny
and sense organs are poorly developed, and they lack
a backbone. The best known lancelets are a group called
amphioxus (pointed at both ends), once thought to be
the archetypal form from which higher chordates evolved
(becoming the subject of legend and song!). With more
information, however, this once again appears to be an
oversimplification, and the ancestry of vertebrates (and of
chordates in general) is still unclear. The different classes
of vertebrates are the subjects of the next three lessons.
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your textbook.
protostome and deuterostome
Lophotrochozoa, Ecdysozoa and Deuterosto-
mia
lophophore
trochophore larva and tadpole larva
ontogeny and phylogeny
recapitulation principle
metameric/segmented and pentaradial/pen-
tamerous
ecdysis/molting and metamorphosis
obligate symbiosis
open and closed circulatory system
endoskeleton, exoskeleton, carapace and os-
sicles
epidermis and cuticle
chitin and calcium carbonate
parapodia and pereopods
setae
siphon
proboscis
compound eyes
biramous
opisthosoma and trophosome
mantle, foot, radula, pen, byssal threads
respiratory pigment: hemoglobin and hemo-
cyanin
gills and tracheae
water-vascular system and tube feet
tunic
protochordate
gill/pharyngeal slits, nerve cord, notochord
and post-anal tail
backbone (vertebrae)
glossary terms
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a. protostome and deuterostome
b. Lophotrochozoa and Ecdysozoa
c. peanut worms and acorn worms
d. parapodia and pereopods
e. tube feet and mollusc's foot
f. pentamerous and metameric
g. ossicles and cuticle
h. molting and metamorphosis
i. horseshoe crabs and horseshoe worms
j. mantle and tunic
2. Many diverse groups of animals are covered in this lesson. Choose one of the more important phyla, and briefly de-
scribe its principal characteristics. Name the major subphyla or classes included in that phylum, and some organisms
belonging to each subphylum/class.
3. Both innkeeper worms and cone shells use a proboscis for feeding, but they use it in different ways. Explain this dif-
ference. Which phylum does each organism belong to, and how closely related are they to each other?
4. Lingula, barnacles and clams look remarkably alike at first glance, since Lingula and clams have a calcium
carbonate shell with two valves, and barnacles also are protected by calcareous plates. Does this mean that these
organisms are closely related, or is this a case of convergent evolution, in which unrelated species evolve similar
structures because of similar lifestyles? Explain your answer, including identifying the phylum to which each of these
animals belongs.
5. What is the recapitulation principle? Explain how, by studying their larvae and applying the recapitulation principle,
we can better understand the phylogeny of each of the following groups: trochozoans, echinoderms and tunicates.
(discuss each group separately)
6. Briefly describe the four distinguishing features of chordates.
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marine fishes
Fishes are the oldest and simplest vertebrates, and gave rise to all the other vertebrates.
They remain the most numerous and diverse vertebrate animals, and are the most
economically important marine organisms..
1. Be familiar with the major groups of fishes, some aspects of their evolutionary his-
tory, and the features distinguishing them as vertebrates.
2. Understand how fishes are anatomically and physiologically adapted to their environ-
ment.
Biology 9th Ed.).
eLearning & Distance Education..
Reading assignment
C&H: Chapter 8 (pp. 151175)
9
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Fig. 13 fish Classification
Phylum Chordata
Subphylum Vertebrata
Agnatha (jawless vertebrates)
Hagfishes
Lampreys
Gnathostomata (jawed vertebrates)
Chondrichthyes (cartilaginous fishes)
Holocephali: chimaeras/ratfishes
Elasmobranchii: sharks, skates, rays
Osteichthyes (bony fishes)
Actinopterygii (ray-finned fishes)
Chondrostei: sturgeons, paddlefishes, birchirs
Neopterygii
gars
bowfins
teleosts: most fishes, e.g., salmon, herring, perch
Sarcopterygii (lobe-finned fishes and tetrapods)
coelacanths
lungfishes
tetrapods
Domain Eukarya
Kingdom Metazoa
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marine fishes
I
In the previous lessons, we learned that animals are
the most diverse multicellular organisms, and that in-
vertebrates account for most of that diversity, with
more than 97% of animal species. We surveyed some of
the major animal phyla, among the 35-40 currently de-
scribed. Phylum Chordata, we learned, includes 3 sub-
phyla, 2 of which are protochordates: the tunicates
and lancelets. The third group, Subphylum Vertebrata,
constitutes the other 3% of animal species and is the
subject of the remaining lessons in Part 2 of this course.
We already know some things about vertebrates. We know
that their tissues are highly organized into organs and or-
gan systems, they are bilaterally symmetrical, have a true
coelom, and develop as deuterostomes. We also know that
vertebrates display 4 features that set chordates apart
from other animals: gill slits, a dorsal nerve cord, a
notochord and a post-anal tail. The key feature that is
unique in vertebrates, setting them apart from all other ani-
mals, is a result of the further evolution of the notochord.
The notochord is a rod derived from mesoderm cells, lying
between the nerve cord and the gut. It is stiff enough for
muscles to attach to, but more flexible than the rigid endo-
skeletons and exoskeletons of non-chordates. In adult ver-
tebrates, the notochord is replaced by a series of hollow
elements made of bone or cartilage, called vertebrae,
which are separated by mobile vertebral joints. This se-
ries of vertebrae forms the vertebral column, also called
the backbone or spine, which encloses and protects the
nerve cord. A bony or cartilaginous skull also evolved to
protect the nerve cord where it extends into the head to
form the brain. The skull and spine (and, later, ribs) make
up the axial skeleton. Additional bony or cartilaginous
elements make up the appendicular skeleton of the ap-
pendages (paired fins or limbs). Together, they form the
endoskeleton, which provides support and protection,
and allows for more efficient locomotion in vertebrates.
Vertebrates include fishes and tetrapods, the four-
footed amphibians, reptiles, birds and mammals.
Vertebrates are by far the most diverse group of chor-
dates, and fishes, the subject of this lesson, are by far
the most numerous and diverse of the vertebrates. They
are the oldest and simplest vertebrates, and gave rise to
all other vertebrates. Refer to Fig. 13 here, and Fig. 8.1
in your textbook, for an outline of fish classification, and
use the table at the end of Chapter 8 to review the char-
acteristics of the various groups of fishes. Also look at the
Geologic Timescale, in the Appendix of this Course Guide.
First Fishes
The oldest known fish fossils were just discovered in 1999,
in some 530 million-year-old rocks in China, at the site of
an ancient sea bed. This places them at the time of the
Cambrian Explosion, much older than any previously
known vertebrate fossils. Paper-clip sized, they lacked a
bony skeleton, teeth and jaws, but had other features set-
ting them apart from lower chordates, including sophis-
ticated gills and musculature, fins, a well-defined head,
paired eyes, and a skull and other endoskeletal struc-
tures made of cartilage. It took another 30 million years
for some fishes to develop the ability to accumulate cal-
cium phosphate in their bodies to form bones, teeth
and scales, and 50 million years more for jaws to evolve.
Fishes were relatively rare during this time, while inverte-
brates flourished. Then a profusion of jawless fishes, pos-
sessing both a bony endoskeleton and an exoskeleton
of bony plates or scales, appeared during the Silurian
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Period (450415 mya). These armored forms, called os-
tracoderms (shell-skinned), were generally less than a
foot long and were probably slow, bottom-dwelling animals,
feeding on detritus and small slow-moving prey. Ostraco-
derms declined and eventually became extinct, with the
expansion of jawed forms during the Devonian Period
(415375 mya), sometimes called The Age Of Fishes.
The evolution of jaws allowed these fishes to become ac-
tive predators, and they diversified rapidly. Placoderms
(plate-skinned) were prominent during this period, includ-
ing many heavily armored forms, some growing to more than
10 feet. Instead of teeth, they had bony plates on the jaws,
sometimes with razor-like, self-sharpening edges. Devonian
placoderms included some of the first freshwater fishes.
Ancestors of the group Chondrichthyes (cartilaginous
fishes), including sharks, also flourished during the Devo-
nian. They are believed to have descended from bony placo-
derms which later reverted to a cartilaginous skeleton. Rep-
resentatives of the other modern fish group, Osteichthyes
(bony fishes) also appeared at this time, including both
ray-finned (Actinopterygii) and lobe-finned (Sar-
copterygii) forms. Lobe-finned fishes dominated during
the Devonian and gave rise to the first tetrapods, about
370 mya. The highly successful and diverse placoderms
became extinct at the end of the Devonian, lasting only
about 50 million years. The once-abundant jawless fishes
are now represented by two small groups, the lampreys
and hagfishes, and cartilaginous fishes and lobe-finned
fishes have also greatly declined. At the same time, a single
branch of ray-finned fishes, the teleosts, have become by
far the most diverse and abundant of all vertebrates, and
the most economically important of all marine organisms.
The Fish Family Tree
Traditionally, vertebrates are divided into two major cat-
egories, Agnatha ("without jaws"), represented today only
by the hagfishes and lampreys, and Gnathostomata
(jaw-mouth) for all other vertebrates, including both the
jawed fishes and the tetrapods. Fishes with jaws fall
into two groups, according to their skeletal material, car-
tilaginous fishes (Chondrichthyes) and bony fishes
(Osteichthyes). Chondrichthyes are further divided into
two subgroups, Holocephali for the chimaeras or rat-
fishes, and Elasmobranchii for the sharks, skates and
rays. Bony fishes also are divided into two groups: Sar-
copterygii (flesh fins) or lobe-finned fishes, including
the primitive coelacanth and lungfishes, the closest living
relatives of the tetrapods; and Actinopterygii (ray fins),
for all other bony, jawed fishes. Ray-finned fishes include
several small groups of primitive, mostly freshwater fishes,
plus one huge groupthe teleosts. Teleosts make up the
vast majority of marine and freshwater fishes (and verte-
brates) alive today. The following survey highlights some of
the advances in vertebrate evolution made by these groups.
AgnathaJawless Fishes
Agnatha are the most primitive living fishes, eel-shaped
animals including hagfishes and lampreys, with only
about 50 species between them. Besides being jawless, they
also lack paired fins, scales and a stomach, and they have
a cartilaginous skeleton. Their smooth skin is covered with
slime. They are sometimes called cyclostomes, because
of their circular, muscular mouth. Hagfishes exist mostly
by scavenging, and many lampreys are parasites. Despite
these similarities, lampreys are now thought to be more
closely related to jawed fishes than to hagfishes, and hag-
fishes are often left out of Subphylum Vertebrata altogether.
Hagfishes: Pinkish and almost blind, these exclusively ma-
rine fishes dine on polychaete worms and dead or dying fish,
sometimes diving right in and eating from the inside out.
Sensing tentacles surround the mouth, which is equipped
with tooth-like rasps for tearing flesh. They can tie them-
selves into knots, sliding the knot against a hunk of flesh
to tear off a bite. They have a slow metabolism and can go
up to seven months without eating. Also called slime eels,
these lovely creatures must sometimes sneeze just to clear
their nostrils of their own secretions. Unusual in that they
have no larval stage, hagfishes lay large leathery eggs which
bear miniature young. They are more advanced than lower
chordates in that they have a partial skull, a cartilaginous
shelf which supports a five-part brain and three types of
sensory organs: paired eyes (probably lost secondarily), an
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olfactory (smelling) organ, and paired inner ears (for
hearing, balance and perception of position of movement)
with one semicircular canal (we have 3, used for balance).
Their gills are used for respiration, rather than feeding,
and they have a ventral heart. Hagfishes have no vertebrae,
however, and are sometimes not classified as vertebrates.
Lampreys: These fishes are anadromous, migrating into
fresh water to spawn (deposit eggs). The young remain in
a larval stage most of their life (up to 7 years), then meta-
morphose and go to sea as adults for one or two more
years, before returning to fresh water for spawning. They
have a large, round sucker surrounding the mouth and a
single nostril on top of the head. Many are parasites, using
the sucker to attach to other fishes and scraping their skin
with a rasping tongue to suck their blood. The sucker is also
used to attach to stones as they make their way upstream
during spawning. Lampreys have a more complete skull
than hagfishes, and skeletal supports for the gills, called
gill arches, with spine-like processes. They also have eyes,
with a lens and extrinsic (outside) eye muscles, two semi-
circular canals, and a rudimentary lateral line system,
a special component of the inner ear in fishes which detects
vibrations and currents in the surrounding water. Lamprey
populations are declining as a result of water pollution.
GnathostomataJawed Vertebrates
Gnathostomes are characterized by jaws (a verti-
cally biting device made up of endoskeletal elements
equipped with teeth), and a bony or cartilaginous spine,
gill arches internal to the gills, paired fins or limbs,
and paired nostrils (nares) leading to paired olfac-
tory organs. They include cartilaginous fishes, ray-
finned fishes, lobe-finned fishes, and the tetrapods.
Chondrichthyes
These are the cartilaginous fishes: chimaeras or rat-
fishes, skates, rays and sharks). Their cartilaginous
skeletons may be a secondary, or derived trait, since
they are thought to have evolved from bony placoderm
ancestors. Small tooth-like enameled scales called plac-
oid scales cover the skin, protecting and giving it a rough
texture. The teeth form continuously in the mouth, with old
teeth falling out to be replaced by new ones. Behind each
eye is a small respiratory opening, the spiracle. Cartilagi-
nous fishes have paired pectoral fins (side fins toward the
front of the body), for efficient swimming, an asymmetrical
(heterocercal) tail, and a ventral mouth. The pelvic
fins (side fins toward the rear of the body) are modified
with claspers in males, which function like a penis for in-
ternal fertilization, as in mammals. They lack a swim blad-
der, and lift is provided by their streamlined shape, pectoral
fins and internal oil bodies which are lighter than water.
Their olfactory senses and lateral line system are well
developed. There are about 1000 recent species (and
many more fossil forms) of cartilaginous fishes, including
two very different groups, the more primitive Holocephali
(chimaeras or ratfishes) and Elasmobranchii (sharks,
skates and rays), which are described in your textbook.
Osteichthyes
These are the bony fishes, which include the vast majority
of fishes, with about 23,000 species. Their bony skeleton in-
cludes rooted teeth, large bones covering the head (cra-
nial bones), a pectoral girdle (shoulder bones), fin rays
supporting the fins, and otoliths (bones in the inner ear).
The fins are more refined than those of cartilaginous fishes,
allowing them greater maneuverability, and the jaws have
more freedom of movement. Thin, flexible cycloid or cte-
noid scales, made of bone and covered with skin and mu-
cus, protect the body and reduce drag by water friction. An
operculum, a bony plate covering and protecting the gills,
enables these fishes to breathe when not swimming. They
also have a swim bladder for controlling buoyancy. Their
terminal mouth is positioned at the front of the head, and
the tail is symmetrical (homocercal). There are two groups
of bony fishes, Sarcopterygii (lobe-finned fishes, plus
the tetrapods) and Actinopterygii (ray-finned fishes).
Lobe-finned fishes, which dominated during the Devonian,
are today represented by only a few species of lungfishes
and coelacanths. Ray-finned fishes include several small
groups, and the huge teleost group, with more than 20,000
species which have expanded into nearly every aquatic niche.
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Sarcopterygii (lobe-finned fishes) are characterized
by rounded paddle-like fins containing bones and muscles,
which in the case of lungfishes (and tetrapods) may be
used to support the body on land. During the Devonian,
many species of marine lungfishes occurred worldwide but
today only a few freshwater species occur in South America,
Australia and Africa. Their swim bladder is a lung which ab-
sorbs oxygen and eliminates wastes. Lungfishes are among
the most ancient fishes present today, and are believed to
be the closest living relatives of tetrapods. Like tetrapods,
they have four similarly-sized appendages, enameled teeth,
and separation of blood flow to the body and the lungs. The
coelacanth, the other type of living lobe-finned fish, also
is an ancient form. In fact, it was thought to be extinct until
1938, when one of these strange creatures was found off the
coast of South Africa (C&H p. 171: "A Fish Called Latimeria").
About 200 more have since been caught in the Indian Ocean.
Actinopterygii (ray-finned fishes): The fins of ray-
finned bony fishes are webs of skin supported by spines
called "rays." With about 23,000 species, they make up
the vast majority of fish diversity, about 96%, and account
for nearly half of all vertebrates. One group, the tele-
osts, is responsible for most of this diversity, with more
than 20,000 species. They include most fishes found from
the surf zone to the ocean depths, in lakes and rivers (and
the aquarium and dinner table). The full suite of advanced
features of bony fishes are displayed by this huge group.
Common teleosts in waters around Alaska include Pacific
Herring, Pacific Halibut, Rockfish and five species of
Pacific Salmon: Chinook (King) Salmon, Coho (Silver) Salm-
on, Sockeye (Red) Salmon, Pink (Humpback) Salmon, and
Chum (Dog) Salmon. Pacific salmon are anadromous (like
lampreys and sturgeons), and have extremely complex life
histories. They may travel thousands of miles from their natal
(birth) streams, spending their adult lives in the North Pacific
and returning precisely to the natal stream to spawn and die.
During spawning they undergo many anatomical and physio-
logical changes including, in some cases, a hooked nose which
gives the genus its name: Oncorhynchus. Salmo, the genus
of Atlantic Salmon, also is anadromous but these salmon
often survive to spawn again. Salmon populations are endan-
gered by human activities, as are many other fish species.
Ray-finned fishes also include several small groups of
bizarre-looking living fossils. They are mostly fresh-
water fishes, like the birchirs, paddlefish, gars and bow-
fin. Sturgeons are found in both fresh and salt water.
The Beluga Sturgeon of the Caspian Sea is prized for its
caviar. The Pacific Sturgeon may weigh 400 pounds and
reach more than 20 ft long, the largest freshwater fish.
These fishes tend to have some primitive features, such
as a cartilaginous skeleton, asymmetrical tail, or lungs.
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your textbook.
vertebrae, backbone/spine/vertebral column
endoskeleton, axial skeleton and appendicular
skeleton,
ostracoderm and placoderm
Agnatha and Gnathostomata
Chondrichthyes and Osteichthyes
cartilaginous fish and bony fish
Holocephali and Elasmobranchii
Actinopterygii and Sarcopterygii
ray-finned fish and lobe-finned fish
teleost
Oncorhynchus and Salmo
placoid scale and cycloid/ctenoid scale
spiracle and operculum
heterocercal and homocercal tail
dorsal fin, tail/caudal fin, anal fin, pectoral
fins, pelvic fins
swim bladder
rectal gland and chloride cells
inner ear, semicircular canal, otolith, lateral
line and neuromasts
olfactory organ/sac
nares
nictitating membrane
ampullae of Lorenzini
gill arch, gill filaments and lamellae
erythrocyte, hemoglobin and myoglobin
countercurrent system of flow
myomeres
chromatophore and iridophore
warning coloration and cryptic coloration
disruptive coloration and countershading
ovoviviparous, oviparous and viviparous
anadromous and catadromous
demersal and schooling
secondary/derived trait
ichthyology
glossary terms
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a. ostracoderms and placoderms
b. Agnatha and Gnathostomata
c. Chondrichthyes and Osteichthyes
d. Holocephali and Elasmobranchs
e. Sarcopterygii and Actinopterygii
f. Oncorhynchus and Salmo
g. semicircular canals and lateral line system
h. spiracle and operculum
i. placoid scales and cycloid (ctenoid) scales
j. heterocercal tail and homocercal tail
k. pectoral fins and dorsal fins
l. oviparous and ovoviviparous
m. anadromous and catadromous
n. chloride cells and rectal gland
2. Describe four uses of coloration in fishes.
3. Discuss three differences between sharks and bony fishes, in the ways they achieve buoyancy and locomotion.
4. Describe four aspects of the circulatory and respiratory systems of fishes, which help them to obtain oxygen ef-
ficiently.
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marine reptiles
and birds
The first tetrapods were four-limbed amphibian-like creatures that emerged
from the water onto land. All tetrapods descended from these newly terrestrial
creatures. Thus, marine tetrapods have made an evolutionary round-triptheir
ancestors came from the sea, onto land, and back to the sea. This lesson provides
an overview of the tetrapods and an introduction to the marine reptiles and birds.
1. Know the distinguishing features of the four tetrapod groups and some of their
evolutionary history.
2. Be familiar with the marine representatives of reptiles and birds, how they are
adapted for a marine existence, and some of their ecological relationships.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 9, pp. 177182 ("Marine Reptiles" and "Seabirds") and Figures 12.12
and 12.13 (p. 277)
10
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Fig. 14 Classification of Marine Reptiles and Birds
Phylum Chordata
Tetrapoda
Amphibia
Amniota
Synapsida (mammals)
Anapsida
Order Chelonia: sea turtles
Diapsida
Order Squamata: sea snakes, marine iguana
Order Crocodilia: saltwater crocodile
Birds:
Order Sphenischiformes: penguins
Order Porcellariiformes: albatrosses, fulmars, shearwaters, petrels
Order Pelecaniformes: pelicans, boobies, cormorants, frigatebirds
Order Charadriiformes: gulls, terns, skimmers, alcids, shorebirds
Order Anseriformes: ducks, geese
Order Ciconiiformes: herons, egrets, bitterns
Order Gaviiformes: loons
Order Podicipediformes: grebes
Order Gruiformes: rails, coots
Domain Eukarya
Kingdom Metazoa
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marine reptiles and birds

I
n our survey of marine organisms, we have arrived at
the tetrapods. We have followed many branchings of
the Tree of Life to get here, moving from the Eukary-
ote branch, to animals, to deuterostomes, to chordates, to
vertebrates, to jawed vertebrates, to sarcopterygians, and
finally to tetrapods. Tetrapods ("four-footed") include am-
phibians, reptiles, birds and mammals. They evolved
from lobe-finned fish ancestors akin to present-day lung-
fishes, probably around 350370 mya during the Devonian
Period. In some lobe-finned fishes, bony supports in the fins
are arranged much like the bones in the limbs of tetrapods,
with one upper bone, two lower bones and many small bones,
like those in the leg, ankle and foot. Skull bones and other
parts of the skeleton also are similar in some lobe-finned
fishes and tetrapods. Fossils of a fish with features inter-
mediate between lobe-finned fishes and tetrapods, which
may represent a link between the two groups, was recently
discovered (C&H p. 80: When Fishes Stepped on Land).
Thus, marine tetrapods have made an evolutionary round-
trip. Their ancestors emerged from the sea onto land, and
all tetrapods descended from these newly terrestrial crea-
tures. Marine tetrapods then made a return trip, back to a
marine existence. Early tetrapods adapted to the new chal-
lenges of living on landlike how to support themselves,
move, breathe and keep from drying outusing the raw
material of their lobe-finned fish ancestors. Marine tetra-
pods, in turn, had to reinvent themselves for aquatic living,
using the raw material of their land-adapted ancestors.
Keep this evolutionary perspective in mind, as you read
about these remarkable animals and the adaptations they
have made to fulfill the basic requirements of life in their
marine environments. This lesson provides an overview of
the tetrapods and an introduction to the marine reptiles
and birds. Marine mammals are covered in the next lesson.
Tetrapods
Tetrapods may be divided into two general groups, the am-
phibians and the amniotes (Fig 14). Amphibians include
frogs and toads, salamanders and newts. They have lungs
and a 3-chambered heart, with one ventricle and two
atria which separate the systemic blood flow (to and from
the body) from the pulmonary blood flow (to and from the
lungs). They also have skin glands to help prevent des-
sication (drying out), eardrums to conduct sound waves,
eyelids and eyes adapted for vision outside of water, and
limbs with more substantial, articulated (jointed) bones to
support their weight on land. The name of this group means
double-life, as they have partially avoided the problem of
dessication on land by returning to the water for part of their
life cycle. Their vulnerable eggs usually are laid in water and
the larvae have gills and swim about, wriggling their tails
and looking a lot like fish. No amphibians are entirely marine.
Amphibian ancestors gave rise to the Amniotes. They
evolved a new type of egg with a shell and four fluid-filled
membranes (the amnion is one) surrounding the embryo.
These structures prevent dessication and provide a stable
environment, allow for gas exchange and waste removal,
and provide food for the developing embryo. Living amni-
otes are commonly classified as reptiles, birds and mammals.
Reptiles (to creep) are cold-blooded ectotherms
and poikilotherms with dry skin and scales, unlike the
soft permeable skin of amphibians. Most have a 3-cham-
bered heart. They have internal fertilization and
various means of reproduction, including ovovivipary.
Their eggshells can be relatively soft or hard, different
species having different degrees of calcification. They
undergo direct development, without a larval form.
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Reptiles have evolved many modes of locomotion, and
include turtles, snakes, lizards, and crocodilians.
Birds are endotherms and homeotherms that lay eggs
with hard shells of calcium carbonate. Their forelimbs
are fashioned into wings. They have feathers derived
from skin cells and made of keratin (like our fingernails
and hair), and a bony beak without teeth. Their bones are
light yet strong, and in some cases hollow, with a keeled
sternum (breastbone) to support powerful flight muscles.
Many bones are fused or lost in birds. They evolved a
4-chambered heart with two atria and two ventricles for
more efficient circulation and complete separation of sys-
temic and pulmonary blood flow. Their lungs are supple-
mented by a complex system of air sacs extending into
the long bones, allowing air to be circulated unidirec-
tionally and continuously. This is a more efficient system
than the tidal flow (inhalation-exhalation) of other tetra-
pods, and allows birds to sustain a higher metabolic rate.
Mammals, like birds, are endotherms and homeotherms
with a 4-chambered heart. They are distinguished by
their hair or fur, and milk-producing mammary glands.
They also have a large brain with a unique region called the
neocortex, which is involved in higher functions (sensory
perception, motor coordination, spatial reasoning). Mono-
tremes (the platypus and echidna) lay eggs, but all other
mammals are viviparous. The egg membranes are modified
into the placenta and umbilical cord, nourishing the em-
bryo and removing wastes, as it develops inside the mother.
A common classification scheme for reptiles, birds and mam-
mals was used in Figure 14 to classify the marine forms of
these animals. They are assigned to the groups, Anapsida,
Diapsida and Synapsida, based partly on their skull struc-
ture. Turtles, the most primitive reptiles, are the only living
representatives of the anapsids, whose skulls are completely
covered in solid bone. Dating from the age of the dino-
saurs, anapsids were previously very diverse. All the other
reptiles, plus the birds, are diapsids. Their skulls include
many fused bones with openings, making them lighter and
more flexible for feeding. In the past, diapsids were amaz-
ingly diverse, with slithering, crawling, running, swimming
and flying forms. They are still the most diverse amniotes,
with over 14,600 species. Mammals are the only surviving
descendants of a diverse group of reptile-like animals that
ruled the land during the Permian Period, the synapsids.
Marine Reptiles
Although a mere shadow of their former diversity, the liv-
ing reptiles are successful terrestrial vertebrates, with
about 7,000 species distributed worldwide. Only a hand-
ful of species have returned to the sea, but these in-
clude representatives of the three major orders: Che-
lonia (turtles), Squamata (snakes and lizards), and
Crocodilia (crocodiles, alligators, caymans and gavial).
Chelonia (turtles): These ancient reptiles are character-
ized their shell, made up of a dorsal carapace and ven-
tral plastron. The shell developed from scales, ribs, ver-
tebrae and elements of the shoulder and hip girdles of
their ancestors. In addition to protection, the shell serves
as a calcium bank from which calcium can be withdrawn
for making eggshells and for buffering the blood during hi-
bernation, when metabolic acids accumulate in the blood.
Like all tetrapods, they breathe air through their lungs, but
turtles get additional gas exchange through the throat
and cloaca (common exit for the excretory and reproductive
tracts). They also have special muscles that aid in inflating
and deflating the lungs, which are enclosed in a rigid rib-
cage. Turtles lack teeth, but can clamp down with powerful
jaws. There are more than 200 terrestrial species, found on
all continents except Antarctica. Only 9 speciesthe sea
turtlesare marine, with limbs modified into flippers
for swimming. Female sea turtles still return to shore to
lay their eggs, sometimes traveling more than a thousand
miles, thus leading a double life opposite to that of amphib-
ians! The hatchlings are easy prey for terrestrial and ma-
rine predators. All sea turtles are considered threatened
or endangered, the victims of human use, abuse and
carelessness (C&H p.180: The Endangered Sea Turtles).
Squamata (snakes and lizards): These are the most nu-
merous and diverse of living reptiles. They are also the most
numerous marine reptiles, including about 55 species of sea
snakes found mostly in tropical waters, and the marine
iguana. The elongate, limbless snake body is pre-adapted
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for efficient swimming, and sea snakes have improved on
this body plan, with laterally flattened bodies and paddle-
shaped tails. The right lung stretches almost the length of
the body, increasing their oxygen stores when underwater.
Like turtles, sea snakes have enhanced gas exchange,
as they are able to absorb additional oxygen through the
skin, and through the throat by swallowing water and then
ejecting it. Excess salt is eliminated by a salt-excreting
gland that sits under the tongue. Like their terrestrial coun-
terparts, they are carnivorous and use venom to subdue
their prey. Most sea snakes are entirely marine and ovovi-
viparous, but some species still return to land to lay eggs.
Some species are becoming rare, as they are hunted for
their skins and are inadvertently caught in trawl fishing nets.
Marine iguanas inhabit the Galapagos Islands, and spend
their days diving into the cold water to graze on seaweed,
and basking on the warm rocks. They use their tails to swim
and cling to the bottom by their claws, diving as deep as 40
ft and remaining underwater for as long as 30 minutes. Salt-
excreting glands located above the nostrils help them to
cope with the high salt content of their diet. They also sneeze
frequently to expell salt, which gives their black bodies a
whitewashed look. Marine iguanas warm themselves behav-
iorally by flattening their bodies against the warm rock sur-
face, and cool down by elevating the torso and exposing it to
the cooling breeze, opening and closing blood vessels in the
chest to control heat exchange. They can also pant to cool
off. Marine iguanas have few natural predators and are pro-
tected in the Galapagos Islands, a national park of Ecuador.
Crocodilia: These ponderous, armored descendants of
the once ruling reptiles are endowed with some surpris-
ingly advanced features. Unlike other reptiles, the amphibi-
ous crocodilians have a complicated 4-chambered heart
(capable of functioning like a 3-chambered heart during
dives), a muscular diaphragm to assist with breathing, and
a secondary palate (bony separation of the mouth and
nasal cavities) which allows them to continue breathing while
submerged with only the nares above water. They have well-
developed senses and are efficient predators, with conical
teeth set in powerful jaws. They use their massive tail for
swimming and to sweep prey into deep water, and can move
quickly on land by belly-crawling, walking and even gallop-
ing on all four legs. They are oviparous, laying their large
eggs on land. The females of some species protect the nest
and care for the young, a unique behavior among reptiles.
One marine species occurs, the saltwater crocodile, which
reaches lengths of 2030 ft, and is found in coastal waters
around Australia, Indonesia and some western Pacific islands.
Marine Birds
Also descendants of the ruling reptiles, birds are the most
diverse tetrapods, with about 10,000 species. Homeo-
thermy and the ability to fly have allowed birds to invade
many environments. Birds in marine environments represent
9 out of about 30 orders of birds (C&H Fig. 9.1), and about
3% of all bird species. They are often divided into two unof-
ficial categories, seabirds and shorebirds. All birds nest
on land, but seabirds spend much of their lives on or above
the ocean, except when breeding. Shorebirds, on the other
hand, tend to stick to the coast, using bays, estuaries and
mudflats for feeding and shelter, especially during migration.
Seabirds: This category includes a number of unrelated
groups, but they share some adaptations to a marine lifestyle,
as a result of convergent evolution. The excess salt in-
gested by seabirds is extracted by salt glands located near
the eyes, and excreted through the nasal cavity or mouth.
Like other birds, they have oil glands for waterproofing
their plumage, but seabirds are more protected from the
wet and cold by denser feathers and a layer of down. Cor-
morants are an exception. These ancient birds lack the oil
gland responsible for waterproofing, reducing their buoyancy
and allowing them to dive deeper in search of food. Terns
also lack waterproof plumage, since they catch their prey at
or above the surface, rarely touching the water. The plum-
age of seabirds is mostly restricted to black, white and grey
colors, to provide camouflage. Their legs are set farther
back on the body than in land birds, a more efficient posi-
tion for swimming and diving. Most seabirds have webbed
feet. Some use them for propulsion when paddling or diving.
Others, like puffins, murres and shearwaters use them
as stabilizers during diving, while albatrosses use their
webbed feet to provide leverage when pushing off against
the water. Most seabirds nest in colonies. They tend to live
longer, breed later, have fewer young, and invest more time
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and energy into raising them, than do other birds. Because
of this greater investment, and because foraging for food
may take place far from the nest site and require long peri-
ods away, both parents usually are involved and pairs are of-
ten monogamous. The reproductive strategies of seabirds
may have evolved in response to the challenges of life at sea,
but also a relative lack of predators compared to life on land.
While they share some traits due to convergent evolution,
seabirds also vary greatly in how they use the ocean. Some
are pelagic, breeding on sea cliffs and small islands and win-
tering on or in the open ocean, totally dependent on the sea
for food. Pelagic seabirds may be further divided into aerial-
ists like the tubenoses (albatrosses, shearwaters, fulmars
and petrels, and open-water swimmers like puffins and
murres (members of the alcid family) and penguins. Other
seabirds are more coastal, but still strongly dependent on
the ocean for food. Coastal seabirds include gulls and terns,
cormorants, pelicans, and frigatebirds. Some species move
inland to nest, like the tundra-nesting skuas. Some species
migrate, as far as thousands of miles each year, to nest and
rear their young. The Arctic Tern holds the record, traveling
25,000 miles to breed in the Arctic and winter in the Antarctic!
Seabirds also differ in the specific ways they fly, swim or dive,
which is reflected in their different wing types. Albatrosses
have long, slender powerful wings (wingspans can exceed 7
ft) for long-distance, nearly effortless gliding. Murres have
much shorter wings with massive breast muscles (account-
ing for one-third of their body weight), which propel them in
short, direct flights or deep dives. Terns and jaegers have
pointed wings for swooping and dipping, while the more all-
purpose wings of gulls are intermediate in length and width.
A variety of modifications of their wings, feet, beak and body,
allows different species to adopt different feeding strategies
and fill different niches. Many birds feed at the water sur-
face, where ocean currents may concentrate prey such as
krill, fishes and squids. Some surface feeders catch prey
while swimming (e.g., shearwaters). Others catch their
prey on the wing, by dipping (gulls), skimming (skim-
mers), pattering (storm-petrels), or snatching (frigate-
birds and some terns). Other species catch prey below the
surface, using a variety of diving styles, and still others
engage in aerial pursuit. Frigatebirds and skuas steal the
food of other seabirds. Skuas, gulls and giant petrels scav-
enge dead animals and prey on the eggs or chicks of other
seabirds, or even on small adult birds. In this way, multiple
species coexist in the same habitat through specialization,
utilizing different parts of the available resources. This is
called resource partitioning (C&H Figs. 9.7 and 9.8.).
Humans have had a long association with seabirds, and con-
tinue to affect their populations in many ways. Traditionally,
people have harvested seabird eggs and hunted birds for
food, oil and feathers. In many cases this has contributed
to the decline of populations, or caused local extinctions,
even the complete extinction of the Great Auk. Hunting and
egging continues today, although less intensely and with
some controls. Seabirds also have played a traditional role
in guiding fishermen to fish, and sailors to land. The rela-
tionship between seabirds and the fishing industry today
is complicated, but in many cases seabirds are negatively
affected. One serious problem is the loss of an estimated
100,000 albatrosses each year, as a result of entangle-
ment in nets set out for tuna and swordfish by longline
fisheries (C&H Fig. 17.6, p. 389). Seabirds also are victims
of oil spills, toxins and pollution, and the introduction of de-
structive species, like rats and feral cats. The establishment
of bird sanctuaries and refuges, an international ban on
drift nets, and other fisheries regulations and conserva-
tion measures are helping to reduce some of these losses.
Shorebirds: Shorebirds belong to the Order Charadri-
iformes, which also includes some seabirds like gulls, terns,
skuas and the alcids. True shorebirds include sandpip-
ers, plovers, stilts and avosets. They are mostly small with
pointed beaks and long legs without webbed feet, for wading
along beaches or in coastal wetlands, while hunting for small
invertebrates and fishes in the sediment or shallow water.
Their relatively long, narrow pointed wings carry them on
migrations as long as 15,000 miles, at speeds of up to 50
mph. Like seabirds, shorebirds have evolved diverse feed-
ing strategies which allow them to exploit various niches and
partition their resources. Probing into the sand or mud, and
gleaning at the surface are two general strategies. Prob-
ers are further divided according to beak length, feeding
on prey at different depths within the sediment (C&H Fig.
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12.12, p. 277). The Long-billed Curlew, for example, has
the longest beak (up to 9 inches), and probes deeper than
the relatively short-billed Willet. Gleaners also divide their
habitat according to leg length, wading different distances
into the surf as they scurry along the beach in search of
food. Other shorebirds have evolved unique strategies, e.g.,
oystercatchers open the shells of mussels and clams using
their thick triangular bill, and turnstones wander the beach,
turning over stones and debris, in search of choice mor-
sels. These are all good examples of resource partitioning.
Members of five other orders of birds, besides Charadri-
iformes, are connected to the sea in ways similar to those of
true shorebirds. These include long-legged waders such
as herons, egrets, bitterns and rails, and swimmers and div-
ers like ducks, geese, loons, grebes and coots. Prized for
their elegant feathers, millions of herons and egrets were
killed each year for the fashion industry, in the late 1800s.
This provoked the first public environmental action, and
resulted in the foundation of the Audubon Society, and
passage of the Federal Migratory Bird Treaty Act of
1918 and the Migratory Bird Conservation Act of 1929.
Migratory animals are particularly vulnerable to disturbances
such as oil spills and pollution, which have destructive ef-
fects on habitats and food webs. Sadly, shorebird popula-
tions are still declining today, with continued human develop-
ment of coastal habitats. Many species are threatened or
endangered. The Endangered Species Preservation
Act of 1966, followed by the Endangered Species Act
of 1973, which is administered by NOAA and the U.S. Fish
and Wildlife Service, were enacted to protect both endan-
gered species and the ecosystems upon which they depend.
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your textbook.
tetrapod
amphibian and amniote
Reptilia, Aves and Mammalia
Anapsida, Diapsida and Synapsida
Chelonia, Squamata and Crocodilia
seabird and shorebird
Charadriiformes, alcids and tubenoses
amnion
ectotherm and endotherm
poikilotherm and homeotherm
atria and ventricles
systemic and pulmonary blood flow
tidal flow and unidirectional flow
direct development
oviparous, ovoviparous and viviparous
convergent evolution and resource partition-
ing
carapace and plastron
keratin and calcium carbonate
salt gland and oil gland
air sacs
diaphragm
cloaca
guano
threatened and endangered species
glossary terms
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Written Assignment for Lesson 1o
a. amniote and amphibian
b. anapsid and diapsid
c. poikilotherm and homeotherm
d. Charadriiformes and Chelonia
e. carapace and feathers
f. seabird and shorebird
g. tubenoses and alcids
h. oil glands and salt glands
i. pulmonary and systemic blood flow
j. convergent evolution and resource partitioning
k. threatened and endangered species
2. Tetrapoda means four-footed. Describe how the limbs of sea turtles, sea snakes and seabirds differ, and how they
are adapted to the lifestyles of these groups.
3. Describe the differences between the eggs of amphibians, reptiles and birds.
4. Compare adaptations for gas exchange in sea turtles, sea snakes and birds.
5. Discuss some of the differences in feeding strategies among birds (choose either shorebirds or seabirds), which
result in different niches for different species, allowing them to coexist.
6. Describe an example of convergent evolution that you learned about in this lesson.
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marine
mammals
In this lesson we conclude our survey of marine organisms with marine mammals, animals
so well-adapted for life in the ocean that it is difficult to believe they are members of our
own class.
1. Be familiar with the various groups of marine mammals, their distinguishing charac-
teristics and representative species.
2. Understand how marine mammals are anatomically and physiologically adapted for
life in the sea.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 9, pp. 186209 ("Marine Mammals" )
11
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Fig. 15 Classification of Marine mammals
Phylum Chordata
Tetrapoda
Amniota
Class Mammalia
(Subclass Prototheria/Monotremataegg-laying mammals)
(Subclass Metatheria/Marsupialiaopossums, kangaroos, etc.)
Subclass Eutheria/Placentaliaplacental mammals
Order Carnivora
(Suborder Feliformia"cat-like"cats, civits, hyenas, etc.)
Suborder Caniformia"dog-like" (dogs, skunks, racoons, etc.)
Family Ursidae: Polar Bear
Family Mustelidae: marine otters
Superfamily Pinnipedia
Family Odobenidae: Walrus
Family Otariidae: sea lions, fur seals (14 species)
Family Phocidae: true seals (19 species)
Superorder Ungulata (horses, elephants, deer, cattle, etc.)
Order Sirenia
Family Dugongidae: Dugong
Family Trichechidae: manatees (3 species)
Order Cetacea
Suborder Mysticetibaleen whales
Family Balaenidae: Bowhead, right whales (4 species)
Family Balaenopteridae: rorquals (10 species)
Family Eschrichtiidae: Gray Whale
Family Neobalaenidae: Pygmy Right Whale
Suborder Odontocetitoothed whales
Family Delphinidae: dolphins, killer whales (37 species)
Family Monodontidae: Beluga, Narwhal
Family Phocoenidae: porpoises (6 species)
Family Physeteridae: sperm whales (3 species)
Family Ziphidae: beaked whales (21 species)
(Superfamily Platanistoidea: river dolphins)
Domain Eukarya
Kingdom Metazoa
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marine mammals
I
n this lesson we conclude our survey of the tetrapods,
and of all marine organisms, with marine mammals.
Mammals have independently made the return-trip to
a marine existence at least five times, with the evolution
of cetaceans, sirenians, pinnipeds, marine otters
and polar bears. They are found in every ocean and on
every continent, in a variety of ecological roles, includ-
ing herbivores (sirenians), filter feeders (baleen whales),
carnivores (pinnipeds) and top carnivores (killer whales
and polar bears). In some cases, they are so well-adapt-
ed for life in the ocean that it is difficult to believe they are
members of our own class. But, like all mammals, they
are air-breathing homeotherms with hair or fur (at
some time during life, even if just a few whiskers), mam-
mary glands that produce milk for their young, and a
large brain including a neocortex. All marine mammals
belong to the subclass of placental mammals, nourish-
ing the embryo through a placenta and bearing live young.
They are long-lived, and have a slow reproductive rate..
Marine mammals are large (including the largest animal ever
to inhabit the Earth, the Blue Whale, growing to over 100 ft
and 150 tons) and tend toward the spherical, being well-
endowed with fat or blubber, all of which gives them a small
surface-to-volume ratio. In most, the external ears are
small or absent, and external genitalia and nipples are hidden
in slits or depressions. The limbs are absent or partly con-
tained within the body, and are greatly modified into flippers
and flukes. These anatomical features help to minimize heat
loss, and maximize streamlining and propulsion in the water.
Marine mammals also have made extraordinary physiological
adaptations to life in the ocean. Their young grow more quick-
ly on milk with a fat content as high as 4050% (human milk
is 3% fat). They breathe more efficiently, store more oxygen
and use it more efficiently, and their tissues are better able
to function anaerobically (without oxygen). These adapta-
tions allow marine mammals to spend long periods underwa-
ter, holding their breath. Breath-holding in marine mammals
also is greatly extended by a physiological adaptation called
the diving response, in which heart rate slows dramatically
(bradycardia) and blood flow to nonessential tissues is re-
duced. Blood flow to essential tissues like the brain and heart
is maintained, and blood oxygen stores last much longer (in
some cases, for more than an hour). The diving response
of marine mammals (and other diving animals) is a highly
developed form of a basic protective response shared by all
vertebrates. Some marine mammals also can dive to great
depths (more than 7,000 ft, in the case of sperm whales)
without getting the "bends," because their collapsible ribs
and lungs limit the absorption of nitrogen into the blood.
The ability to selectively control blood distribution also is used
by marine mammals for thermoregulation. They can con-
serve or dissipate heat in the extremities (even in the tongue,
in the case of the great whales), as needed. The ability to
utilize water present in food, inspired air and blubber, and
to excrete urine that is saltier than seawater, enables marine
mammals to conserve water. Excellent hearing, acute tactile
senses, the ability to communicate underwater with sound,
and the use of sound waves to locate, and perhaps stun, prey
(echolocation) are other examples of the extraordinary
adaptations marine mammals have made to life in the sea.
A classification scheme for marine mammals is shown in Figure
15. Once again, it begins with their Domain and summarizes
major branching points along the long evolutionary paths by
which these animals came to be. This scheme differs from
the one your textbook uses (Fig. 9.1), in that pinnipeds are
included in Order Carnivora, just another example of science
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as an ongoing quest, subject to debate. The marine mammal
groups, including all members of the orders Cetacea and
Sirenia, and some members of Carnivora, are introduced
below, and are discussed in greater detail in your textbook.
Order Cetacea
Cetaceans: The approximately 90 species of whales,
dolphins and porpoises who comprise the Order Cetacea
(from Greek ketos meaning sea monster) are all com-
pletely aquatic mammals. Their ancestors (surprisingly, re-
lated to cloven-hoofed animals like cattle and deer) began
returning to the sea about 50 million years ago. Cetaceans
feed, mate, give birth and suckle their young in the water.
They are the most specialized mammalian swimmers and div-
ers, with flippers for forelimbs, no external hindlimbs, hori-
zontal flukes that propel with a vertical motion, and a dor-
sal fin. Their highly compressed neck vertebrae and a thick
layer of blubber add to their streamlining, and their nostrils
are on top of their head, forming a blowhole. Some ceta-
ceans are capable of swimming at 25 mph, others can dive to
over 7,000 ft and remain submerged for close to two hours.
Cetaceans include two groups, Mysticeti or baleen whales
(e.g., bowhead whales and gray whales) and Odontoceti or
toothed whales (e.g., sperm whales, dolphins and porpois-
es). Mysticetes filter huge mouthfuls of water through their
baleen plates, trapping krill and small fish, except the gray
whales, which are benthic feeders. Odontocetes have peg-like
teeth used to strain or grasp larger prey, usually fish or squid.
Cetaceans are long-lived, highly intelligent social animals
with a low reproductive rate. They are well known for their
vocalizations, and can hear sounds many miles away, and
discern their direction. Some odontocetes also are capable
of sophisticated echolocation, perhaps best developed in
bottlenose dolphins, which are trained by the Navy to detect
and mark underwater mines. The sensitivity of cetaceans
to sound makes them vulnerable to noise pollution in the
water, for example from ship propellers. Stranding deaths
have been linked to high intensity sonar bursts during na-
val operations. Cetaceans may be harmed by all types of
marine pollution, oil spills in particular. Many species are
threatened or endangered, primarily because of over-
hunting of whales and entanglement of dolphins in fishing
gear. The formation of the International Whaling Com-
mission in 1946, passage of the United States Marine
Mammal Protection Act of 1972, and other attempts to
regulate human activities and conserve whale populations
have helped some to recover, but harmful practices continue.
Order Sirenia
Sirenians: So named because they were once mistaken for
mermaids, sirenians are the only other entirely aquatic group
of mammals. Their lineage is as old as cetaceans', and they
share many similar adaptations (another example of conver-
gent evolution). Unlike other marine mammals, they are
herbivores, like their closest living relatives, the elephants.
Slow, passive creatures with a low metabolism, they do not
tolerate cold water despite their large size and blubber. They
are restricted to warm, shallow waters with adequate veg-
etation, along coasts and in estuaries and rivers. Commonly
called manatees, dugongs or sea cows, only four species
remain, and all are endangered. They were slaughtered in
the past, and today face habitat loss, pollution, collisions with
boats and other impacts related to human population growth
and coastal development. A fifth species, the Stellers Sea
Cow, was hunted to extinction as recently as about 1770.
Order Carnivora
Pinnipeds (seals, sea lions and walrus): More recently
returned to the sea than cetaceans or sirenians, pinnipeds
(winged feet) still retain some ties with land. They come
ashore or onto the ice to mate, give birth and suckle their
young, or to molt. They are awkward on land, but are su-
perb swimmers and divers. Pinnipeds share some charac-
teristics with both their terrestrial carnivore relatives (they
are most closely related to bears), and with other marine
mammals. They are sleek-bodied and rather large, with four
webbed flippers, have both fur and blubber, and sen-
sory organs adapted to function both in air and in water
(sea lions have been trained by the Navy to locate objects
in deep, murky water). All are carnivorous, eating primar-
ily fish, shellfish and squid. They range from the polar ice
packs to the tropics, and from coasts to the open ocean.
About 35 species belong to three families: Otariidae
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(eared seals), Phocidae (earless seals) and Odo-
benidae (the walrus). Also over-hunted for their skin,
meat and blubber, many species remain endangered.
Marine Otters: These members of the Carnivora or-
der are the largest representatives of the weasel fam-
ily (Mustelidae), and the smallest marine mammals. At
least 6 of the 13 existing otter species are fully or par-
tially adapted to a marine existence. They all occur at high
latitudes. The sea otter, which inhabits the North Pacific
Ocean and the chungungo, which is found in the South-
east Pacific Ocean, are the only otters that feed exclusively
at sea. Chungungos often go ashore to eat their prey,
rest, give birth and rear their young, whereas sea otters
rest, mate, give birth and suckle their young in the water.
As relative newcomers to life at sea, marine otters lack
many of the adaptations seen in other marine mammals.
The sea otter has a long, flat tail and webbed hindfeet
used for swimming and making relatively short, shallow
dives. Their front paws lack webbing and have retractable
claws used for grooming and feeding. They have no blubber,
but have a relatively high metabolic rate, and the densest
fur of any mammal. Additional insulation is provided by air
trapped in the fur, which must be kept clean for their sur-
vival, so sea otters spend about 10% of their time groom-
ing. Demand for its pelt nearly led to the sea otters ex-
tinction, and today the species remains endangered.
Sea otters are also subject to disease, habitat loss, oil
spills, entanglement in plastic trash, and conflict with aba-
lone fisheries. Previously, they ranged as far south as Baja
California, but today can be found only in the coastal wa-
ters of the northern Pacific. Sea otters feed primarily on
benthic invertebrates like clams, mussels, urchins and
crabs. One of the only animals besides primates known
to use tools, they use rocks or shells to pry their prey
off the rocks, and to hammer or pry open their shells.
Polar Bear: The largest and most carnivorous bear (fam-
ily Ursidae), the polar bear is the marine mammal least
adapted to an aquatic existence, yet is totally reliant on
sea ice as its primary habitat. Distributed throughout
the ice-covered waters of the circumpolar Arctic, polar
bears depend on sea ice as a platform for travel and ac-
cess to land, for seeking mates and breeding, and for hunt-
ing their primary prey, ringed seals (which also depend
on sea ice). They are particularly dependent on polynas,
areas of open water surrounded by ice (Lesson 3), which
are often richly productive and attractive to marine mam-
mals and seabirds. They may travel long distances on
the ice or in the water, in search of food or mates. Males
overwinter out on the pack ice, while pregnant females
dig dens on land or on pack ice, where they give birth.
Weighing up to 1,700 pounds, these huge bears have a
short snout and small ears, and are covered with dense
white fur with water-repellent guard hairs, and a layer of
fat up to 5 inches thick. Their feet also are huge and fur
covered on the bottom, acting as oars in the water and dis-
tributing their weight on the ice, like snowshoes. They are
equipped with extremely stout, curved claws for clambering
onto the ice and gripping prey. Polar bears have an extraor-
dinary ability to go for long periods without food or water,
entering a walking hibernation when food is scarce.
Circumpolar peoples have long hunted polar bears, and
there has been concern about over-harvesting of this spe-
cies. At the top of the trophic pyramid, they also are espe-
cially vulnerable to pollutants, which become concentrated
with each step up the food chain, a phenomenon called
biomagnification. Natural substances also become con-
centrated, and eating polar bear liver can cause "hyper-vi-
taminosis A" due to toxic levels of Vitamin A. However, polar
bears now face an even greater, unprecedented threat, with
the rapid deterioration of their sea-ice habitat as a result of
climate change. The extent and thickness of sea ice in the
Arctic has decreased drastically, and it has been breaking
up earlier in recent decades, as we learned in Lesson 3.
The deterioration of sea ice is correlated with a decline in
the physical and reproductive health of polar bears. Loss
of their habitat, and their declining populations and slow re-
productive rate, which limit their ability to replace individu-
als, are cause for grave concern for the future of this spe-
cies. In the U.S., they were listed as threatened in 2008.
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your textbook.
Cetacea, Sirenia and Carnivora (and Pinnipe-
dia)
Mysticetes and Odontocetes
whales, dolphins and porpoises
baleen whales and toothed whales
rorquals and great whales
manatees, sea cows and dugongs
Otariidae, Phocidae and Odobenidae
eared seals, earless seals and walrus
Mustelidae and Ursidae
marine otters and sea otter
mammary glands and milk
uterus and placenta
neocortex
fur, guard hairs and blubber
flippers and flukes
blowhole and spout/blow
baleen plate
melon, spermaceti organ and ambergris
viviparous
homeotherm and endotherm
thermoregulation
diving response
bradycardia/bradychardia
apneustic breathing
the bends
lactic acid
vocalizations and echolocation
pods
breaching
stranding/beaching
migrations
delayed implantation
International Whaling Commission and Marine
Mammal Protection Act
glossary terms
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1. Name three orders of mammals that include marine mammals, and two marine species from each order. ("species"
are unique organisms - for example, "whale" is not a species)
2. Heat is conducted away from warm bodies 23 times faster in water than in air. Describe four anatomical adaptations
for minimizing heat loss in marine mammals.
3. List the three families of pinnipeds and, for each family, describe two distinguishing characteristics and one repre-
sentative species. (again, the name of a unique organism)
4. To which of the following is a sea otter most closely related: walrus, polar bear, dugong, ringed seal? To which of the
following are you most closely related: octopuses, insects, sea squirts, jellyfishes?
5. Like land mammals (and other tetrapods), marine mammals are air-breathers. Describe four physiological adap-
tations of the respiratory systems of marine mammals that increase the efficiency of gas exchange in a marine
environment.
6. Briefly describe the feeding habits of one species of marine mammal.
7. Describe one unique behavior of a marine mammal that interests you.
8. Describe one example of convergent evolution involving unrelated species of marine mammals. Describe another
example of convergent evolution, involving marine mammals and some non-mammalian animals.
9. What reasons can you think of for there being so many endangered and threatened species of marine mammals?
It's time for your second exam. Locate a proctor in your area and complete the Exam Request Form on the fol-
lowing pages. (If you live in the Fairbanks area there is no need to complete the form. Come to our office during
business hours to take your exam.) Allow 4 hours for completing the exam.
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proctors.
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INSTRUCTIONS
itself.
907-479-3444 fax: 907-479-3443
Student Name:
Course Title:
Mailing Address:

Proctors signature:
Student's Address:

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Part three
Putting It All
Together: Marine
Ecosystems
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Life At The Top:
The Epipelagic
Realm
Although the epipelagic has been accessible to humans since they began
sailing the seas, and has been studied at least since Darwins time, we still
have much to learn about this realm.
1. Understand the physical characteristics of the epipelagic environment, and
the challenges and opportunities they present for the organisms living there.
2. Know the types of organisms found in the epipelagic, and their adaptations
for life in this environment.
3. Be familiar with some of the important trophic relationships among the vari-
ous members of epipelagic communities.
1. Read the information for this lesson in the following pages of this Course
Guide.
2. Complete the Reading Assignment (below) in C&H (Castro & Huber, Ma-
rine Biology 9th Ed.).
3. Complete the Written Assignment (last page of the lesson) and submit it
to UAF eLearning & Distance Education.
Reading assignment
C&H: Chapter 15 (pp. 332-359) and reread "Eye on Science" p. 431 ("Effects of
Climate Change on Arctic Peoples")
More information, including interactive quizzes, flashcards, videoclips and links,
can be found at the publisher's Online Learning Center: www.mhhe.com/
castrohuber9e
12
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Life At The Top: The Epipelagic Realm
N
ow that we have described the general character-
istics of the marine environment, and surveyed the
major groups of marine organisms, we are ready
to explore specific marine habitats and the unique biotic com-
munties they support. In this lesson, we explore life near the
surface of the ocean, in the epipelagic zone. In the next
lesson, we move to the deeper ocean and, in Lessons 14 and
15, we will explore coastal environments, including the inter-
tidal and subtidal zones and estuaries. Each of these environ-
ments has distinct physical characteristics to which organisms
are specifically adapted, interacting with each other and with
the environment to form complex and unique ecosystems.
The sunlit epipelagic is teeming with life, all supported
by vast numbers of mostly microscopic phytoplank-
ton. Primary production by the epipelagic phytoplank-
ton supports consumers not only at the surface but in
other environments as well. Organic matter produced
at the surface sinks down, supporting life in the deeper
ocean (and many deep ocean creatures venture up into
the epipelagic at night to feed). It also is carried by cur-
rents into coastal marine environments, and is exploited
by land-dwelling birds and mammals, including humans.
Although the epipelagic has been accessible to humans since
they began sailing the seas, and has been scientifically stud-
ied at least since Darwins time, we still have much to learn
about this realm. Recall (Lessons 5 and 6) that vast numbers
of minute cyanobacteria, Synechococcus and Prochlo-
rococcus, were unknown until fairly recently. Yet they appear
to constitute the majority of all phytoplankton, and rank as
the most numerous forms of life on Earth. They are mem-
bers of the tiny picoplankton (C&H Fig. 15.2, Table 15.1),
too small to be caught in standard plankton nets. The pi-
coplankton also include some archaea, an entire domain
of life unrecognized until a few decades ago. And we have
recently discovered that marine viruses outnumber the pi-
coplankton (and all other living things) by 10:1 to 100:1.
This necessitated the addition of another category of plank-
ton even smaller than picoplankton, the virioplankton, or
femtoplankton. Although viruses are not considered to
be living organisms by most scientists, we are learning that
they play an extremely important role in the marine food web.
Phytoplankton cells infected by these vast numbers of ma-
rine viruses (about 10 million in a drop of seawater) release
dissolved organic matter (DOM) into the ocean, which is re-
cycled back into the marine food web through the microbial
loop (C&H Figs. 15.4 and 15.25). We now realize that DOM
accounts for as much as half of all the epipelagic primary
production, so the microbial loop is a very significant part of
the epipelagic food web. Other members of the picoplankton,
minuscule heterotrophic bacteria and archaea, perform
the vital function of consuming DOM, bringing it back into
the food web. Somewhat larger, but still microscopic, proto-
zoan members of the nanoplankton complete the micro-
bial loop by consuming these bacteria and archaea, which
are too small to serve as prey for larger predators. The en-
ergy from DOM enters the rest of the epipelagic food web
when larger net zooplankton consume the protozoans.
Our understanding of epipelagic communities has been
significantly altered by these discoveries, and continues to
evolve as more forms of life are discovered and the complex
interactions within these communities become more clear.
We also have much to learn about the complex physical and
biological interactions between the epipelagic, the deeper
ocean and the atmosphere. A better understanding of these
regional and global relationships is critically important now
that conditions appear to be in a state of rapid change.
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Life In And Around Sea Ice
A specialized but extremely important part of the epipelagic
environment is sea ice, whose physical characteristics we
studied in Lesson 3. Surprisingly, these are some of the most
productive environments on Earth. They are also among the
least well known because of the difficulties of accessing and
working in areas dominated by ice. Polar environments are
among the most extreme on Earth and are subject to the
most extreme seasonal fluctuations. They are also the en-
vironments where climate change is most progressed. Spe-
cies superbly adapted for life in these extreme environments
are faced with rapidly changing conditions, and the future of
many is uncertain. How these changes will affect the complex
interactions that sustain communities and ecosystems (and
global weather systems and climate) is of great concern.
Unique communities of organisms, called sympagic com-
munities, can be found living within sea ice and at the ice-
water interface. Sea ice is infiltrated by a network of min-
ute brine channels formed during the process of brine
rejection. These are inhabited by viruses, a multitude of
microbes (bacteria, diatoms, flagellates), and an assortment
of ice meiofauna, including tiny flatworms, nematodes,
crustaceans and rotifers. Diatoms (more than 200 species
have been identified from Arctic sea ice) and other types of
ice algae are the major producers in sympagic food webs.
Insulated from the cold air, algae are able to grow in the
lowermost ice layers. Mats of algae grow on the underside
of the ice, and algal filaments may extend several meters
into the water column. In the Arctic, ice algae can contrib-
ute better than half of the total marine primary production.
Many organisms thrive on the underside of the ice, some
grazing on algae, others preying on grazers or on each oth-
er. Some species are endemic (unique) to sympagic com-
munities, for example some gammaridean amphipods,
which graze in large numbers on the underside of Arctic sea
ice. Amphipods, copepods and other zooplankton are the
major food source of Arctic cod (Boreogadus saida), a
key species in Arctic marine food webs. These small, oily fish
are extremely abundant and have a widespread, circumpolar
distribution. Ranging farther north than any other fish spe-
cies, they carry antifreeze glycoproteins (or AFGPs) in
their blood, which prevents damage from freezing. (Some
unrelated Antarctic fishes produce similar AFGPs, another
example of convergent evolution.) Arctic cod are an im-
portant food source, for larger fish (e.g., Atlantic cod, Green-
land halibut and Arctic char), seabirds (e.g., gulls, kittiwakes,
fulmars, murres and guillemots), seals (e.g., ringed, spotted
and ribbon) and whales (e.g., beluga and narwhal). They
even supplement the diet of polar bears, Arctic foxes and
local human populations, in areas where they are washed
ashore in large numbers by winter storms, and they are
taken by some fisheries. Thus, energy is transferred from
life within the ice to the water column, by grazers eating ice
algae and themselves being eaten by Arctic cod, which in
turn fall prey to animals at higher trophic levels. Additional
trophic levels are added by fisheries netting consumers of
Arctic cod (e.g., salmon fisheries), polar bears consum-
ing their primary prey (ringed seals), killer whales preying
on seals and other whales, seabirds that eat the chicks
and eggs of other seabirds (e.g., glaucous gulls) and na-
tive peoples subsisting on fish, seabirds, seals and whales.
During the Arctic spring and summer, the melting seasonal
ice pack releases fresh water, in contrast to the salt re-
leased by brine rejection during freezing. As fresh water
spreads from the melting ice edges, it creates a layer of less
dense water that floats at the surface, producing a halo-
cline and setting the stage for a phytoplankton bloom. The
melting sea ice releases ice algae, which seeds the bloom.
All the accumulated organic debris of sympagic life is re-
leased as well, fueling the bloom. If the water is not disturbed
by a strong vertical mixing event, stratification keeps the
phytoplankton at the surface where they grow quickly in this
sunlit, nutrient-rich environment. These ice-edge blooms
are critical for both pelagic and benthic life in the Arctic. If
the ice melts too early in the spring, when there is sufficient
sunlight for photosynthesis but the water is too cold for many
zooplankton to be present, then most of the energy from
an ice-edge bloom sinks, becoming fuel for the benthos. If
the ice melts later in the spring when the water has warmed
up enough for zooplankton, then the ice-edge bloom sup-
ports a rich pelagic food web. Zooplankton such as krill and
copepods graze the phytoplankton and are eaten in turn by
larger invertebrates, like jellyfish, and by fishes, seabirds
and mammals including ringed seals and bowhead whales.
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Bowheads are huge, weighing as much as 75100 tons.
More than a third of this bulk is taken up by the head, with
an enormous bow-shaped mouth carrying 460 to 720 of
the longest baleen plates of any whale (up to 4 m long).
With perhaps the biggest mouth of any animal ever to ex-
ist, they are able to attain their tremendous size by engulf-
ing huge mouthfuls of water and straining out vast num-
bers of tiny zooplankton. They may grow more slowly and
live longer (more than 100 years) than any other mammal.
This Arctic food web is amplified by smaller fishes, called
forage fishes, which eat zooplankton and then become
prey to larger fishes, seabirds, seals, and whales like nar-
whals and belugas. Pelagic forage fishes include Arctic cod,
smelts, capelin and sand lance, as well as the juveniles of
larger species, like salmon. Feeding relationships can be
quite complex. For example, adult salmon eat adult smelts,
while some adult smelts prey on juvenile salmon, and juve-
nile salmon eat the smaller juvenile smelts (and the adults
of many fish species prey on juveniles of their own kind).
Most pelagic fishes migrate in huge schools, so both for-
age fishes and larger fishes are worthwhile targets for
even the largest predators, like sea lions, killer whales
and humpback whales, as well as for human fisheries.
An ice-edge bloom does not occur if the seasonal ice retreats
too early in the spring, before there is sufficient sunlight to
fuel photosynthesis, or if no seasonal ice formed at all. There
may be a later open-water bloom with large numbers of
zooplankton quickly consuming the phytoplankton and trans-
ferring energy to the pelagic food web. However, unlike ice-
edge blooms, open-water blooms do not benefit from the
nutrients and stratification associated with melting sea ice.
Ice-edge blooms also concentrate their production, in terms
of both space and time, which allows consumers to forage
more efficiently. Foraging fish, seabirds and mammals all fol-
low the retreating ice edge, which may move more than a
thousand kilometers during the season. Benthic communities
also benefit from the organic matter that sediments down
from the flourishing pelagic life supported by an ice-edge
bloom. This trickle-down effect is particularly important
where the ice edge advances and retreats over shallow con-
tinental shelf regions, for example in the Bering Sea. These
regions can be hugely productive, teeming with jellyfish, mol-
luscs, crustaceans, fishes, seabirds and marine mammals.
Large populations of both pelagic fishes (salmon) and ben-
thic species (halibut, pollock) support some of the most lu-
crative fisheries in the world. The Bering Sea pollock fishery
is the largest single species fishery in the world today (a ma-
jor source of the fish sticks in your local supermarket freez-
er). All this productivity ultimately depends on the presence
of the seasonal ice pack, and the timing of its annual retreat.
Some benthic communities benefit from another phenom-
enon associated with sea ice: polynas, areas of open wa-
ter in the ice pack where prevailing winds keep blowing ice
away from the leeward side of coasts and islands. These
features become ice-making machines, as the exposed
water freezes, then the ice is blown away, exposing more
water to freezing temperatures, and so on. The repeated
formation of ice at polynyas produces a surface layer of
very dense salty water (through brine rejection) which
sinks, setting up a current that carries surface nutrients
down to benthic environments. The benthos in some re-
gions may depend on these currents, generated at large
polynyas that tend to form in the same place year after year.
Sea ice is a critical habitat for many organisms. The envi-
ronment just under the ice serves as a nursery ground for
Antarctic krill, Arctic cod, and other invertebrates and fishes,
while ice seals and walruses give birth on top of the ice
pack. Pagophilic (ice-loving) seals include ringed, spotted,
ribbon, hooded, bearded and harp seals in the Arctic, and
Weddell, Ross, crabeater and leopard seals in the Antarc-
tic. Most ice seals nurse their pups on the ice, and in many
cases the pups are unable to swim until weaning, which takes
almost two months for ringed and Weddell seals. The young
of most pagophilic seals are born with soft downy furthe
lanugowhich is white in some species, like ringed seals.
Ringed seals carve out lairs in snowdrifts or pressure ridg-
es, reducing the exposure of the pups to weather and preda-
tion by polar bears. Seals and walruses need to haul out on
the ice to rest, molt, and avoid marine predators like killer
whales. They can dive under the ice to escape from other
predators, like polar bears and humans. Polar bears are ex-
cellent swimmers but require sea ice to travel and rest on,
and to access their primary prey, ringed seals. When the sea-
sonal ice pack is gone, they come ashore and usually fast un-
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til the ice forms again in the fall. They breed on the ice pack,
and females make their winter dens on sea ice as well as on
land. Arctic foxes venture out onto the ice, preying on ringed
seal pups and scavenging the remains of seals killed by polar
bears. The bowhead whale spends its entire life closely asso-
ciated with sea ice, in leads or polynyas or at the edge of the
ice pack, where belugas and narwhals also winter. All three
cetaceans are beautifully adapted for life in this environment,
lacking a dorsal fin, insulated by a thick layer of blubber and
capable of breaking through the ice to breathe. Finally, sea
ice provides access to food for the Inuit peoples inhabit-
ing the coastal Arctic, who have survived for thousands of
years by hunting seals, walruses, whales and polar bears.
Sea-ice environments offer other critical advantages to the
animals living there. Compared to land areas available for
hauling out, the ice provides a vast amount of space and a
variety of different habitats, with less competition for food,
and isolation from most predators and other disturbances.
Crowding is reduced and the environment is cleaner. The ice
shields the water below from the extremely harsh weather in
these regions, and pressure ridges and snowdrifts provide
shelter above the ice. Sea ice allows easy access to food,
especially where it is concentrated at the edge of the ice
pack and in polynys, or in areas shallow enough for benthic
feeders like walruses and bearded seals to reach the bottom
when diving from the ice platform. Drifting ice also is used by
animals for migration and transport to new feeding areas.
Sea ice is melting, as we learned in Lesson 3. With global
warming happening twice as fast at the poles as on the rest
of the planet, there has been a dramatic decline in the ex-
tent and thickness of sea ice, and the ice pack is freezing
later and breaking up earlier. These changes have enormous
local implications for the organisms living in these regions.
In parts of the Antarctic where sea ice is declining, popu-
lations of Antarctic krill, which require sea ice for their
nursery grounds, have greatly decreased and are being re-
placed by salps, which typically inhabit warmer waters. Some
krill-dependent predators already are declining. Among the
hardest hit are Emperor and Adelie penguins, which
feed almost exclusively on krill. The location of their nest-
ing colonies, and the timing of their breeding cycles, have
evolved to coincide with the proximity of their food source
at the ice edge, which now is receding farther and earlier.
These changes are surely producing others as they ripple
through the community in a trophic cascade, affecting the
predators, prey and competitors of the altered populations.
There is also great concern for Arctic sympagic communities,
and the pelagic and benthic communities supported by ice-
edge blooms, in view of the dramatic declines in Arctic sea
ice that we are witnessing. Some changes are already being
seen. As sea ice has been thinning, forming later and break-
ing up earlier, walruses have been crowding onto shore in
unprecedented numbers (up to tens of thousands), at earlier
times in the year, and in areas where they have not been seen
before on the northwest coast of Alaska. On land, animals are
subject to greater stress, and they must travel greater dis-
tances to reach foraging areas. During the summer of 2007,
several thousand mostly young walruses died in stampedes,
when they were hauled out along the Chukchi and Bering Sea
shores. In 2009, 131 died in a similar event. Walruses need
an ice platform thick enough to support their great weight,
and it needs to be positioned over water shallow enough that
they can dive down to the bottom to feed. The ice edge also
carries them great distances over fresh foraging grounds,
as it retreats north in the spring. Now, the ice is becom-
ing thinner and is receding over deeper and deeper water.
Ice seals also require ice that is thick enough and persists
long enough for them to complete the critical stages of birth-
ing and rearing of pups, and the annual molt. Ringed seals,
the most abundant and widespread marine mammal in the
north and the principal prey of polar bears, also depend
on snowdrifts for their lairs. Snowdrifts have been melting
earlier, when the pups are still dependent on these protec-
tive caves. Longterm studies of polar bears in Hudson Bay
are revealing that their condition and reproductive rates also
are decreasing, along with the decreasing seasonal ice pack.
Some scientists fear that the majority of the worlds polar
bears, including all those off the coast of Alaska, will be gone
by mid-century, or sooner (in 2008, the U.S. Fish and Wildlife
Service officially listed them as threatened). Arctic seabirds
such as the black guillemot also are imperiled by shrinking
ice conditions. Like the Antarctic Adelie penguin, they must
nest within range of the productive ice edge so they can feed
themselves and their chicks. Longterm studies of black guil-
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lemots nesting on an island in the western Beaufort Sea have
shown a drastic decrease of this population in recent years.
Declining populations of ice-dependent animals threatens
the subsistence livelihood of the Inuit people, as well. The
receding ice also forces them to travel farther away from
shore and has resulted in rougher water, making hunting
more difficult and dangerous. Greater storm surges are
fast eroding the coastlines and have already forced the in-
habitants of some Arctic coastal villages to relocate. At the
same time, the shrinking ice is opening these areas to in-
creased shipping, and oil and gas exploration and extrac-
tion. Although economically advantageous, these activites
also create disturbances and pollution that pose serious
risks to marine life. Arctic cod, for example, have buoyant
eggs with an unusually thin membrane. These adapta-
tions are suited to the calm under-ice environment where
spawning occurs, but makes the eggs particularly vulner-
able to rough water and to pollution. Arctic cod and other
marine organisms may be harmed by pollution, distur-
bance and loss of habitat, at any stage of their life history.
High latitude communities are particularly vulnerable be-
cause they tend to have relatively low species diversity,
compared to temperate and tropical communities. Breaking
one link in a food chain can have a more profound effect on
the whole food web because fewer chains, and fewer links
in each chain, are involved. Loss of an anchoring spe-
cies like Arctic cod, which many other species depend on for
food, can have a particularly devastating effect. The rapid-
ity of change at the poles is also a factor. It is much more
difficult for organisms to adapt to change when it comes
quickly and in large doses. Since the study of ice-dominated
regions is still in its infancy, it is possible that some forms of
life unique to these environments may be lost before they
are even discovered. Continued study is crucial to our un-
derstanding of these ecosystems, how they are changing,
and the regional and global implications of these changes.
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your textbook.
epipelagic zone and photic zone
neritic zone and oceanic zone
phytoplankton and zooplankton
megaplankton, macroplankton, mesoplankton,
microplankton, nanoplankton, picoplankton
and femtoplankton/virioplankton
holoplankton and meroplankton
Prochlorococcus, Synechococcus, and Tricho-
desmium
buoyancy and drag
neuston and pleuston
DOM, detritus and marine snow
microbial loop
suspension feeders
herbivorous/grazing zooplankton and carnivo-
rous zooplankton
planktivorous nekton
forage fishes
protective coloration, countershading and
self-shading
rete mirabile
vertical migration
algal blooms, spring blooms, red tides, HABs,
and eutrophication
ice-edge bloom and open-water bloom
limiting factor, light-limited, nutrient-limited
and limiting nutrient
stratification and overturn
coastal upwelling, Antarctic upwelling and
equatorial upwelling
biomixing and biological nutrient pump
El Nio-Southern Oscillation (ENSO) and La
Nia
sympagic and pagophilic
brine rejection and brine channels
albedo
critical habitat
nursery ground
endemic species
trophic cascade
anchoring species
glossary terms
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a. plankton and nekton
b. phytoplankton and zooplankton
c. picoplankton and net plankton
d. holoplankton and meroplankton
e. Trichodesmium and Prochlorococcus
f. copepods and krill
g. DOM and detritus
h. microbial loop and rete mirabile
i. self-shading and countershading
j. red tides and ice-edge blooms
2. By definition, plankton can't swim against the current. Describe three ways that plankton manage to avoid sinking
out of the productive epipelagic zone.
3. Protozoans, microscopic animal-like protists (Lesson 5), play a critical role in the epipelagic food web. Describe this
critical function they serve, and name several types of protozoans found in the epipelagic environment.
4. Planktivorous nekton come in a huge range of sizes. Briefly describe this category of epipelagic organisms and give
several examples of planktivorous nekton.
5. Most of the epipelagic nekton are not planktivorous. Describe three adaptations shared by many epipelagic nekton,
which make them very efficient carnivores. Who are some of the most efficient epipelagic predators?
6. The epipelagic, coinciding with the photic zone, is the place to see and be seen. Describe three adaptations used by
epipelagic organisms to avoid being seen and eaten.
7. A typical Antarctic epipelagic food web is depicted in Figs. 10.14 and 10.15 in your textbook (pp. 222 and 223), and
a North Sea food web topped by herring is shown in Fig. 15.24 (p. 349). Using these as a model, draw a food web
for the Arctic, identifying at least five trophic levels, including (but not necessarily limited to) the following organisms:
Arctic cod, beluga, black guillemot, bowhead whale, diatoms, gammaridean amphipods, Inuit people, jellyfish, killer
whale, polar bear, ringed seal, salmon.
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The
Netherworld:
the mesopelagic
& deep-sea
realms
Pelagic organisms found in the midwaters and deep sea are fascinating and otherworldly,
bizarre-looking creatures displaying amazing adaptations for coping with their harsh
environment.
1. Understand the physical characteristics of the diverse environments encountered
below the epipelagic: the mesopelagic, deep-sea pelagic, and deep-sea benthic,
including deep-sea hot springs and cold seeps.
2. Be familiar with the types of organisms found in mesopelagic and deep-sea environ-
ments, and their anatomical, physiological and behavioral adaptations.
3. Understand some of the trophic relationships within mesopelagic, deep-sea pelagic
and deep-sea benthic communities.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 16 (pp. 361381) and "Deep-Water Coral Communities" (p. 321)
13
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The Netherworld:
The Mesopelagic & Deep-Sea Realms
B
elow the epipelagic zone lies the most vast and least
known environment on Earththe mesopelagic
(or midwaters), a twilight world, and the deep
sea, where no sun at all penetrates. The deep sea realm
includes the bathyal and bathypelagic, abyssal and
abyssopelagic, and the hadal and hadopelagic zones.
The fact that we can cover all these habitats in just one les-
son is a reflection of our ignorance of this immense and
relatively inaccessible environment (much as grouping mi-
croorganisms together into one unit is a reflection of our
lack of knowledge of these hugely abundant, diverse organ-
isms). The ocean depths seem a forbidding place for life,
cold and dark and under extreme pressure. Indeed, the
concentration of organisms here is much less than in the
epipelagic, or the other shallow and well-lit environments we
will explore in the next two lessons. Recent exploration has
shown, however, that the mesopelagic and deep sea, the
largest environment on Earth, also contains an abundance
of life. One inhabitant of the ocean depths, a toothy little
fish called the benttooth bristlemouth (Cyclothone ac-
clinidens), is the prime candidate for most abundant ver-
tebrate on Earth, although very few people have seen it.
Biodiversity in the ocean depths is proving to be even
more astonishing. What little sampling has been done on
the deep ocean floor suggests that the deep-sea benthos
may be the most diverse group of organisms on Earth. As
much as 90 percent of the specimens pulled up from the
bottom by some expeditions have never been seen before,
and every two weeks, on average, a new deep-sea spe-
cies has been described since the 1980s. Extrapolating
from these findings, some scientists have estimated the
number of deep-sea species to be as high as 10 million!
Until the HMS Challenger expedition of the 1870s, it was
commonly assumed that no life could survive in the deepest
sea. The thousands of specimens dredged up by Sir Thom-
son and his crew from depths of up to almost 5 miles proved
otherwise, though many were badly mangled in the process.
The first to observe deep-sea life in its natural environment
was the American naturalist, William Beebe. In the 1930s,
he descended half a mile down in a tethered metal bathy-
sphere less than 3 feet wide, and was astonished to see
the "dancing" lights of so many bioluminescent creatures.
In 1954, the Swiss scientist Auguste Piccard, who had pre-
viously made the first flight into the stratosphere, realized
his dream of descending untethered into the deep sea. He
reached a depth of 4,000 m by using weights attached to
his spherical metal bathyscaphe, then dropped the weights
and ascended by means of a gasoline-filled balloon attached
to the bathyscaphe. In 1960, Piccard's son Jacques and U.S.
Navy Lieutenant Don Walsh made their historic trip to the
deepest place on Earth, 7 miles down into the Challenger
Deep of the Mariana Trench, in the Trieste, the second
generation bathyscaphe designed by Auguste. In 1964, the
first maneuverable deep-diving submersible, the Alvin, was
driven over the sea floor, by controlling the angle and speed
of its large aft propeller. Alvin and the French submersible,
Cyana, were used in Project FAMOUS (French-American
Mid-Ocean Undersea Study) during which the Mid-Atlantic
Ridge was observed for the first time, in 1972. A few years
later, expeditions were made to the floor of the Pacific. Hy-
drothermal vents, and the astonishingly lush communities
surrounding them, were observed for the first time in 1977.
These discoveries spurred the pace of deep-sea exploration,
conducted by American, French, Canadian, Russian, Japanese
and Chinese scientists, using both manned submersibles and
tethered and untethered robots. Still, less than one percent
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your textbook.
mesopelagic, bathypelagic, abyssopelagic and
hadopelagic zones
midwater and deep-sea
dysphotic zone and aphotic zone
oxygen minimum zone (OMZ)
deep scattering layer (DSL)
main thermocline
Trieste and Alvin
vertical migrator and non-migrator
bioluminescence and photophores
counterillumination
tubular eyes and bilobed eyes
hermaphrodites and male parasitism
pheromones
deep-sea gigantism
chemosynthetic prokaryote
meiofauna and macrofauna
epifauna and infauna
baitfall
hydrothermal vent, black smoker and white
smoker
cold seep
seamount
deep-water corals and bioherm
biodiversity
glossary terms
of the sea floor has been observed. Unfortunately, many
deep-sea environments are being damaged by overfishing,
trawling and other destructive practices. Some forms of life
unique to these environments may be lost before they can even
be discovered. Each of the 30,00050,000 seamounts,
the vast majority of which are unexplored, may be home to a
unique community of organisms. Unique deep-water coral
communities also have been found at depth. Lacking zoo-
xanthellae in their lightless environment, these deep-water
corals form mounds known as bioherms which can grow to
more than 1,000 ft high and can be more than 4,000 years
old. Unique deep-sea communties such as these, holding an
incredible wealth of biodiversity waiting to be discovered, are
threatened by human activities. Uncovering the full extent
to which life has evolved in this dark mysterious world de-
pends not only on scientific exploration and investigation, but
also on our ability to manage how we exploit its resources.
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1. The mesopelagic zone is a twilight world, whereas no sunlight at all penetrates down to the deep sea. Many of the
inhabitants of these realms have evolved adaptations for producing their own light, a phenomenon called biolumi-
nescence. Briefly discuss three differences between mesopelagic and deep-sea animals, in their use of biolumines-
cence.
2. Mesopelagic organisms are largely dependent on the leftover crumbs of primary production that sink down from the
epipelagic zone. Nearly all midwater fishes, however distantly related, share certain anatomical characteristics which
allow them to cope with this scarcity of foodanother example of convergent evolution. Briefly describe four of
these characteristics and how they benefit these organisms.
3. The DSL (deep scattering layer), discovered in the course of sonar soundings during World War II, changes position
from day to night as a result of a habit engaged in by most mesopelagic organisms. What is this habit, and why does
it occur?
4. The oxygen minimum zone, which we learned of in Lesson 3, occurs in the mesopelagic zone. What are two adapta-
tions organisms have made for living under these oxygen-starved conditions? What global ocean phenomenon is
responsible for replenishing oxygen in deep-sea waters and for returning nutrients, accumulated in deeper waters,
back to the surface?
5. Very little food is available to organisms living in the deep sea. Benthos on the deep-sea floor have a slight advan-
tage because material that reaches the bottom stays put, and they are occasionally treated to a food bonanza in the
form of "whale falls" and other baitfalls. What is a baitfall, and what are the usual foods/feeding habits of the deep-
sea benthos (discuss both meiofauna and macrofauna)?
6. The deep sea is vast, dark and sparsely populated. Describe three evolutionary adaptations of deep-sea organisms
that enable them to find mates and accomplish reproduction under these conditions.
7. Perpetual darkness, cold and extremely high pressure make all aspects of life difficult in the deep sea, and the abun-
dance of organisms living there is low. Is the diversity of species also low in this environment? Briefly discuss the
biodiversity of the deep-sea benthos.
8. Some deep-sea benthic habitats do support life in abundance. Where are these flourishing communities found (de-
scribe at least two)? Who are their primary producers? Describe several of the animals living in these communities,
directly or indirectly supported by these primary producers.
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on the shelf:
Continental
shelf
communities
The subtidal zone is home to a variety of communities living on the continental shelf's
mostly unvegetated, sandy or muddy surface, as well as kelp forests and seagrass beds,
and most of the world's coral reefs.
1. Understand the physical characteristics of benthic environments in the subtidal
zone, and the challenges and opportunities they present for organisms living in
these environments.
2. Be familiar with the subtidal benthos, including coral reef communities, and how they
are adapted for life in these environments.
3. Understand some of the trophic relationships of coral reefs and other subtidal
benthic communities.
Biology 9th Ed.).
Reading assignment
C&H: Chapters 13 and 14 (pp. 287-305 and 307-330)
14
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on the shelf: Continental shelf communities
B
enthic communities in the subtidal (or sub-
littoral) zone, the part of the marine environ-
ment over the continental shelf below the low
tide line, are the subject of this lesson. We learned about
pelagic life in the water column over the shelf (the ne-
ritic zone) in Lesson 12. Communities more closely
associated with the shoreline, in the intertidal zone
and estuaries, will be covered in the following lesson.
The subtidal zone is home to a variety of communities living
on the shelf's mostly unvegetated, sandy or muddy surface,
as well as kelp forests and seagrass beds and most of
the world's coral reefs. The subtidal is a much smaller zone
than those lying beyond the shelf break, which we studied
in the previous lessons. It is the richest part of the ocean,
however, containing most of the world's fishing grounds,
and major oil and mineral resources. The coral reefs found
here are the most diverse and complex of all marine eco-
systems, rivaling the richness of tropical rainforests. Since
the advent of scuba technology, the subtidal environment
has become generally accessible. Its proximity to land also
makes it especially vulnerable to exploitation by humans,
and to pollution and other damage from human activities.
Most of the world's major fisheries and many continental
shelf communitiesin particular, coral reefshave already
been damaged as a result of these activities and face an
uncertain future, as we will explore further in Lesson 16.
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your textbook.
subtidal/sublittoral zone
low tide line
continental shelf and shelf break
tidal currents and turbulence
stratification
sedimentation
substrate
lithogenous and biogenous sediments
benthos
demersal
epifauna, infauna and meiofauna/interstitial
animals
deposit feeder and suspension feeder
passive suspension feeder and filter feeder
bioturbator
encrusting algae, coralline red algae, and
calcareous green algae
epiphyte
kelp bed and kelp forest
canopy, understory and algal turf
holdfast, stipe, frond and pneumatocyst
sporophyte and gametophyte
anthozoans and hydrozoans
hermatypic and ahermatypic
scleractinians
fringing reef, barrier reef, atoll and lagoon
patch reef, coral knoll, pinnacle, key/sand cay
and algal ridge
reef flat, reef crest, reef slope, back-reef
slope and fore-reef slope
buttress/spur and groove formation
mesenterial filaments and sweeper tentacles
planula
zooxanthellae
bleaching
bioerosion
eutrophication
ocean acidification
ENSO/El Nio
recruitment
competition and competitive exclusion
facultative symbiont and obligate symbiont
trophic cascade
glossary terms
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1. The continental shelf accounts for a relatively small part of the marine environment, compared to the epipelgic,
mesopelagic and deep sea, but it is the richest part of the ocean. Describe three physical characteristics of the
continental shelf environment, explaining how each contributes to its high productivity.
2. Most inhabitants of unvegetated soft-bottom environments in the subtidal zone dine on detritus. Describe how two of
these organisms, one epifauna and one infauna, make use of detritus.
3. Seagrasses, as we learned in Lesson 6, are the only truly marine representatives of the angiosperms, or flowering
plants. Their roots grow into the sand or mud of soft-bottom subtidal environments, where they can form thick sea-
grass beds. While few organisms feed directly on seagrasses, seagrass beds provide in many other ways. Describe
three ways that seagrass beds benefit their inhabitants or visitors.
4. Although plant-like in appearance, seaweeds lack true roots and require a hard substrate on which to attach. Major
primary producers in hard-bottom subtidal communities, they are subjected to wave action, variable temperature
and light conditions, grazing, and competition. Describe three adaptations that seaweeds have evolved to contend
with these challenges.
5. Kelps grow in cold nutrient-rich waters, along rocky coasts of temperate and subpolar regions. They are the most
conspicuous members of complex, highly productive kelp communities. When one set of feeding relationships in
these communities is disturbed, the effects can ripple throughout the entire community, causing a "trophic cascade"
effect. Describe the trophic cascade that has been studied in kelp forests around the Aleutian Islands of Alaska,
involving sea urchins, sea otters, killer whales and kelps.
6. Reef-building corals obtain nourishment in a variety of ways. Briefly describe four ways that these organisms feed
themselves. Which way is most critical for their reef-building ability?
7. Growing and sustaining a coral reef is a "team effort." Describe how three organisms, other than reef corals, make
critical contributions to this process.
8. The richest and most complex of all marine ecosystems, tropical coral reefs are teeming with life. Describe two spe-
cific ways in which organisms compete with each other (for space or food), and two ways that they assist each other,
to be successful members of these highly competitive communities.
9. Although they have lots of help, reef corals are the chief designers and builders of coral reefs. Reef corals require
very specific physical conditions for their growth and well-being, and are very sensitive to disturbances in their
environment, which put coral reefs at great risk. Describe three physical requirements of tropical reef corals, and
discuss one example of how their environment has been disturbed and the health of the coral reef ecosystem has
been negatively affected.
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between two
worlds:
The intertidal
zone & estuaries
In this lesson, we explore habitats with widely varying physical conditions, because of their
location at the intersection of two very different environments.
1. Understand the physical characteristics of intertidal and estuarine environments,
and the challenges and opportunities they present for the organisms living there.
2. Know the types of organisms found in the intertidal zone and in estuaries, and their
adaptations for life in these environments.
3. Be familiar with some of the important trophic relationships within intertidal and
estuarine communities.
Biology 9th Ed.).
Reading assignment
C&H: Chapters 11 and 12 (pp. 244-266 and pp. 268-285)
15
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between two worlds:
the intertidal Zone & estuaries
I
n this lesson, we explore habitats with widely varying
physical conditions, because of their location at the in-
tersection of two very different environments. The in-
tertidal zone, the most accessible and familiar part of the
marine environment, is alternately immersed in saltwater and
exposed to the air. Its inhabitants have evolved a wide variety
of adaptations for coping with this extremely dynamic envi-
ronment. Estuaries also are exposed to unique and vari-
able conditions where the land meets the sea, at the outflow
of rivers. Estuarine habitats include salt marshes, mud-
flats and mangrove forests, as well as seagrass beds,
which we studied in the previous lesson, and oyster reefs.
These are the marine environments most directly, and
most severely, affected by human activities. They have
been subjected to dredging, dumping, draining, dam-
ming, construction, farming, logging, mining, pollution and
invasive species, in addition to the damage done by ero-
sion, drought, storms and rising sea levels. The extent of
these coastal wetlands has been drastically reduced and
is still declining. We are belatedly learning how important
they are. By destroying these environments and commu-
nities, we are damaging our own species, more than half
of whom are concentrated within 60 miles of a coast.
Coastal wetlands provide critical habitats for many spe-
cies. They also serve as air conditioners, cooling the days
and warming the nights, and as the Earth's "kidneys," filter-
ing and cleansing the water that runs through them. In the
wake of violent storms like Hurricane Katrina in 2005 and
Superstorm Sandy in 2012, and devastating tsunamis such
as the 2004 Indian Ocean tsunami and 2011 Japan tsunami,
it has also become clear that they provide vital protection to
our coastal communities. Estuaries buffer waves and storm
surges and act like a giant sponge, soaking up excess water
and reducing flooding. Many efforts are now underway to
repair and restore wetlands, but it is proving very difficult.
These are among the most complex ecosystems on Earth
and, once lost, it may not be possible to duplicate them.
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your textbook.
intertidal/littoral zone
active margin and passive margin
low tide, high tide, spring tide and neap tide
diurnal tide, semidiurnal tide and mixed tide
headland and bay
wave refraction
wave shock
tide pool
immersion and emersion
desiccation
dispersal
vertical zonation
upper intertidal, middle intertidal and lower
intertidal
substrate
rocky bottom and soft bottom
epifauna and infauna
meiofauna/interstital animals and bioturba-
tors
deposit feeder, suspension feeder and filter
feeder
sessile
holdfast, pneumatocyst, and byssal threads
interstitial water
anoxic
hemoglobin
transplantation, removal and caging experi-
ment
ecological succession and climax community
keystone predator
competitive exclusion and resource partition-
ing
coastal plain estuary (drowned river valley)
wetlands
bar-built estuary, sand bars and barrier
islands
tectonic estuary and fjord (glacier-carved
estuary)
mangrove forest (mangal), tidal/salt marsh,
mudflat, and oyster reef
negative estuary
salt wedge
tidal current and tidal bore
outwelling
brackish water
euryhaline and stenohaline
osmoconformer and osmoregulator
active transport and osmosis
anadromous and catadromous
pneumatophore
invasive species
biological clock
eutrophication
glossary terms
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1. Complete the crossword puzzle on the following page, and submit it (or a list of the answers) with this assignment.
(All the answers in the crossword puzzle are included in the "Glossary Terms" for this lesson.)
2. The intertidal is the smallest zone within the marine environment, but it is the best known because of its accessibility.
At low tide, we can easily observe rocky shore intertidal organisms as they deal with a unique challenge: emersion.
Describe two of the problems caused by emersion and, for each of these problems, give two examples of how inter-
tidal organisms have evolved a solution.
3. Water movement due to waves, tides and currents is another challenge of life in the rocky intertidal. Describe how
two organisms cope with wave shock.
4. One obvious difference between rocky and soft-bottom coastal environments is the opportunity for organisms to
burrow into the substrate of soft-bottom habitats. Describe how this is accomplished by three types of infauna in
soft-bottom intertidal or estuarine communities.
5. Rocky intertidal habitats display a distinct vertical zonation. For each of the three vertical zones, briefly describe how
one organism typically found there is adapted for life in that zone.
6. Estuaries also are subjected to extreme interactions where the land meets the sea. What physical factor fluctuates
most widely in estuarine environments? Describe the strategies adapted by three different organisms for coping with
this widely varying condition in estuarine environments.
7. Primary producers, directly or indirectly, provide the energy and raw materials needed by all other members of a
community. Describe some of the most important producers that support life in the following intertidal and estuarine
environments: rocky intertidal, mudflats, and salt marshes. (If organic matter is supplied indirectly, by producers
outside the community, also include that in your discussion.)
8. Estuaries are home to relatively few species. Those species are abundantly represented by large numbers of indi-
viduals, however, and the productivity of estuaries is very high. Many nonresident organisms partake of this bounty,
on a daily or seasonal basis, or at some point in their life cycle. Describe two of these visitors and how they make
use of estuarine environments.
9. Estuaries are extremely important to us because they filter and cleanse outflow, and buffer the effects of storms
along our coasts. Unfortunately, we have destroyed a large percentage of estuarine environments. Describe two hu-
man activities which have contributed to the destruction of these vitally important ecosytsems.
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crossword puzzle for Lesson 15
(Tear out and submit for grading)
Lesson 15 Crossword
1
2 3 4 5
6 7
8
9
10
11
12
www.CrosswordWeaver.com
ACROSS
7 "high and dry" in the intertidal (opposite of
"immersion")
9 pattern of banding in rocky intertidal
communities (2 words, no space)
10 impact from water crashing on shore, in the
intertidal zone (2 words, no space)
11 process by which surplus detritus is carried
from estuaries out into the open ocean
12 water of intermediate salinity
DOWN
1 tolerant of a wide range of salinities
2 extensions of mangrove roots, growing up
into the air
3 final stage of ecological succession (2
words, no space)
4 carnivore that makes a disproportionate
impact on the community (2 words, no
space)
5 devoid of oxygen
6 dense mangrove forests
8 collective term for marshes and other
coastal environments, partially or
completely flooded at high tide
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the human
factor

We are only beginning to recognize the impacts of human activities on the world ocean, just
as we are only beginning to explore this vast realm and to understand its significance in the
global ecosystem that is Earth's biosphere.
1. Be familiar with the resources of the sea and how they are used by humans.
2. Recognize some of the scientific, economic, social and political aspects involved in
managing marine resources.
3. Understand the impacts of humans on marine environments and marine life, and
efforts to conserve and restore them.
4. Grasp the global significance of marine environments and ecosystems, the extent of
their destruction, and the importance of their conservation.
Biology 9th Ed.).
Reading assignment
C&H: Chapter 17 (pp. 383404), Chapter 18 (pp. 406423) and "Special Report: Our
Changing Planet" (pp. 231-242)
16
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The human factor
T
hroughout this course, there have been numerous
references to the effects of human activities on ma-
rine environments: the break-up of ice shelves and
the retreat of sea ice as a result of global warming (Les-
sons 3 and 12); damage to continental shelf habitats due to
a variety of factors, including pollution, increased sediment
levels and global warming (Lesson 14); and the destruction
of important estuarine environments, as a result of coast-
al development and other human activities (Lesson 15).
We have also noted some of the disastrous consequences
suffered by marine organisms as a result of our activities:
destruction of seagrass beds, mangrove forests and coral
reefs (Lessons 14 and 15); damage to fish populations from
overfishing (Lesson 9); the threatened or endangered status
of all species of sea turtles and some sea snakes due to
over-hunting, and the loss of seabirds and shorebirds as a
result of oil spills and other types of pollution, destruction
of habitat, and indiscriminate fishing practices (Lesson 10);
and the threatened or endangered status of many marine
mammals from over-hunting, pollution and loss of habitat,
including deteriorating sea-ice conditions (Lessons 11 and
12). Some well-known species have been pushed to the brink
of extinction, such as some of the great whales. Others,
like the great auk and Steller's sea cow, have already been
exterminated (Lessons 10 and 11). Far greater numbers
of less conspicuous forms of life are disappearing, some
even before being discovered, such as deep-sea benthic
communities subjected to trawling (Lesson 13) and sympagic
communities threatened by global warming (Lesson 12).
We have seen that relationships among the members of a
community are intricately interwoven, and that interfering
with one species can have far-reaching effects, such as the
trophic cascade effect of removing a predator from the
food web in kelp forests around the Aleutian Islands (Les-
son 14). Disturbing the physical environment also sets in
motion a series of consequences. Increased greenhouse
gases cause global warming, resulting in the loss of sea
ice. Sea ice provides critical habitat for many organisms
and its formation helps to drive the great ocean conveyor,
which in turn supplies oxygen to deep waters and nutri-
ents to surface waters, as well as profoundly influencing
global climate. Disturbances can escalate unexpectedly
when positive feedback occurs, as with the accelerated
warming and melting seen at the poles (Lessons 3 and 12).
Another consequence of increasing greenhouse gases is
ocean acidification (Lesson 3). The decreasing pH of
seawater over the last 200 years corresponds to a 30% in-
crease in hydrogen ion concentration. If this trend continues,
a 3-fold increase in the concentration of hydrogen ions may
occur by the year 2100, higher than has occurred for hun-
dreds of millenia, and at a much faster rate of change than
at any time during this period. Acidification changes other
aspects of ocean chemistry, by increasing the weathering of
rocks and altering the availability of nutrients and the ef-
fects of toxins. Its most direct impact on marine life may be
to impede the growth of organisms with calcium carbonate
shells, including diatoms and krill, key links in the food chains
of many marine ecosystems, many species of crustaceans
and molluscs that we rely on for food, and the reef-forming
corals which support the richest of all marine ecosystems.
In addition to altering the pH of the ocean, human activi-
ties are now responsible for 70% of the reactive nitro-
gen (forms of nitrogen useable by primary producers) in
the environment. The amount of nitrogen that we pump
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into the air and ocean increases the incidence of can-
cer and other diseases, acid rain, and explosive algal
blooms which inundate coral reefs, cause paralytic shell-
fish poisoning, and create dead zones in the ocean.
Attempts to lessen or reverse some of the damage caused
by human activities have included the enactment of the Fed-
eral Migratory Bird Treaty Act, the Endangered Spe-
cies Act, the Marine Mammal Protection Act, and the
formation of the International Whaling Commission
(Lessons 10 and 11). A key objective of the biodiversity as-
sessment carried out by the Census of Marine Life (COML)
project is to target conservation efforts (Lesson 5, C&H p.
216). UNCLOS (United Nations Convention on the Law of
the Sea), the 1982 Law of the Sea treaty not yet ratified by
the U. S., is the global effort to regulate use of the world's
oceans, including conservation of marine habitats and life
(Lesson 1). Comprehensive information on marine eco-
systems in polar regions is being provided in the north by
BEST (the Bering Sea Ecosystem Study) and in the south
by SO-GLOBEC (Southern Ocean GLOBEC) and ICED (Inte-
grating Climate and Ecosystems Dynamics in the Southern
Ocean), parts of U.S. GLOBEC, the U. S. Global Ocean Eco-
system Dynamics Program. It is vital that we work to better
understand and monitor these high-latitude systems, which
are undergoing the most rapid changes and which are criti-
cally important for nutrient cycling and productivity in the
world ocean, and powerful influences on global climate.
In this lesson, we will explore further the impacts of hu-
man activities on marine ecosystems, and our attempts to
manage these precious resources more carefully and intel-
ligently. We are only beginning to fully recognize the con-
sequences of human activities, just as we are only begin-
ning to explore the sea and to understand its significance
in the global ecosystem that is Earth's biosphere. It is
increasingly clear that we are imperiling the health of the
world ocean, and that this will have far-reaching conse-
quences for all of Earth's inhabitants, including ourselves.
Our own future depends on our understanding of these
complex systems and, more importantly, on our actions.
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your textbook.
anthropogenic impacts
finfish and shellfish
clupeoid fishes and demersal species
stock/population
catch and by-catch
fisheries and industrial fisheries
fishing effort, optimal effort and overfishing
maximum sustainable yield
gill net, longline, purse seine, trawl and
ghost fishing
fish flour (FPC), fish meal and fish oil
renewable resource and nonrenewable re-
source
UNCLOS, EEZ, COML, U. S. GLOBEC, SO-GLOBEC,
ICED and BEST
Sustainable Fisheries Act of 1996 and Pew
Oceans Commission
Marine Mammal Protection Act and Interna-
tional Whaling Commission
Federal Migratory Bird Treaty Act
Endangered Species Act
aquaculture/mariculture
salmon ranching and salmon farming
seeding
genetic engineering
marine natural products
polymetallic nodules
tidal energy, ocean thermal energy conversion
(OTEC)
methane hydrate (clathrates)
biodegradable, non-biodegradable and per-
sistent substance
desalination and saline brines
sewage treatment and sludge
heavy metals, radioactive waste and solid
waste
hydrocarbons, chlorinated hydrocarbons and
PCBs
dispersants
thermal pollution
reactive nitrogen, nitrogen cascade and eutro-
phication
hypoxic/dead zone
carbon emissions, greenhouse gas and green-
house effect
ocean acidification
biological magnification
global distillation
native species and invasive species
rare species, threatened species and endan-
gered species
extinction and extermination
marine protected areas, habitat restoration
and artificial reef
glossary terms
After completing your written assignment for this lesson it will be time for your final exam. Locate a proctor in
your area and complete the Exam Request Form following the appendix. (If you live in the Fairbanks area there
is no need to complete the form. Come to our office during business hours to take your exam.) Allow 4 hours for
completing the exam.
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a. finfish and shellfish
b. demersal fishes and clupeoid fishes
c. trawls and purse seines
d. optimal catch and by-catch
e. renewable resources and nonrenewable resources
f. EEZs and PCBs
g. biodegradable and persistent
h. biological magnification and global distillation
i. sludge and saline brines
j. eutrophication and ocean acidification
k. polymetallic nodules and marine natural products
l. invasive species and native species
2. Describe the relationship between the size of a population ("stock") and the growth rate of the population. What
range of population size, and what range of fishing effort, is most conducive to a maximum sustainable yield, or
optimal catch? What has been the actual fishing effort of most of the world's fisheries in recent decades, and what
impacts have fisheries made on fish stocks?
3. Describe three factors, other than fishing effort, which have adversely affected fishery resources. (factors could
include other human activities, or physical phenomena)
4. Given the decline of fish stocks, and a growing human population, the practice of mariculture is gaining interest.
Describe three potential problems associated with this industry.
5. Human activities greatly affect marine ecosystems, not only as a result of what we remove, but also what we add to
the sea. Briefly discuss the effects of three types of marine pollutants.
6. Of all the impacts humans make on marine life and marine environments, briefly discuss one that has made a par-
ticular impression (positive or negative) on you.
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proctors.
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INSTRUCTIONS
itself.
907-479-3444 fax: 907-479-3443
Student Name:
Course Title:
Mailing Address:

Proctors signature:
Student's Address:

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appendix
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Assembled from the International Commission on Stratigraphy 2004 report A Geologic Time Scale and from information on the
Univeristy of California Museum of Paleontology website (http://www.ucmp.org). This work is licensed under the Creative
Commons Attribution-NonCommercial License. To view a copy of this license, visit http://creativecommons.org/licenses/by-
nc/2.5/ or send a letter to Creative Commons, 559 Nathan Abbott Way, Stanford, California 94305, USA.
Geologic Time Scale
Eon Era Period Millions
of years
ago
Major biological events
Neogene Today
23
The beginning of the Neogene is when the first hominids
(early humans) appeared. Modern humans appeared and
developed civilization by the end of this period. Mammoths,
sabre-toothed cats, and giant camels dominated until
10,000 years ago when many large mammals went extinct.
C
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o
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c

E
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a
Paleogene 23
65
Rise of the mammals and birds. Rodents, primates, pigs,
cats, dogs, bears and whales appear. Flowering plants
spread across the globe.
Cretaceous 65
145
Dinosaurs continue to dominate the land. Marsupials,
modern sharks, bees and butterflies appear. Flowering
plants appear. Period ends with the mass extinction of the
dinosaurs and many plants.
Jurassic 145 200 Dinosaurs dominate the land. Mammals are common but
small. Feathered dinosaurs and birds appear. The most
common land plants are ferns, palm-like trees called
cycads, and grasses. M
e
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z
o
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c

E
r
a
Triassic 200 251 The few survivors of the Permian extinction go on to
populate the land and oceans. New species like mammals,
dinosaurs and crocodiles appear.
Permian 251 299 Amphibians dominate the land. Early cone-bearing plants
like pine trees appear. Period ends with the largest mass
extinction known with 95% of all marine species and 50%
of all animals going extinct.
Carboniferous 299 359 Many swamps on land and sponge reefs in the oceans.
Reptiles appear. Early winged insects and cockroaches
appear.
Devonian 354 417 Fish spread across the oceans. Amphibians appear. The
first trees and other plants spread across the land creating
the first forests.
Silurian 417 443 Spiders, scorpions, insects, complex plants, and fish with
bony jaws appear. Fish adapt to living in rivers and fresh
water for the first time.
Ordovician 443 488 First land plants appear. Primitive fungi and sea weed
appears. The oceans are full of corals, mollusks, worms,
primitive fish, and echinoderms like starfish.
P
h
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n
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a
Cambrian 488 543 A large number of new animal species appear in a relatively
short time. First fish appear. No known life on land yet.
Proterozoic Eon 543
2,500
First multi-celled organisms like sponges appear. Earliest
complex life forms are algae from 1.4 billion years ago.
Oxygen begins to accumulate in the atmosphere.
Archean Eon 2,500
3,800
Earliest life on Earth are bacteria from 3.5 billion years ago.
Earth is very different from today. The atmosphere is
mostly methane and ammonia. The continents only just
have begun to form.
Hadean Eon 3,800
4,570
No known life. The Earths crust cools and solidifies. The
moon forms.
retrieved from http://www.mysciencebox.org/timelines under creative commons license.

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Introduction To Marine Biology

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An Independent Learning Course

The Benthic Environment

The Pelagic environment

Marine Invertebratesthe lower phyla

marine reptiles and birds

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