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CONTENTS
List of Focus Boxes xi CHAPTER 4

Preface xiii
ENERGY AND HEAT 61
4.1 Heat and Thermal Energy 61
CHAPTER 1 4.2 Temperature 62
A CHANGING PLANET 1 4.3 Heat Transfer 63
1.1 Earth’s Beginnings 1 4.4 Heat Capacity and Specific Heat 75
1.2 Early Primitive Life 2 4.5 State Changes and Latent Heat 76
1.3 Evolution of Earth’s Atmosphere 4 4.6 Energy Quality 77
1.4 Aerobic Life 5 4.7 Entropy, Heat Engines, and the
1.5 Earth’s Changing Climate 7 Second Law of Thermodynamics 80
1.6 Earth’s Energy Endowment 9 4.8 Energy Quality, End Use,
1.7 The Human Era 14 and Cogeneration 84
Chapter Review 17 4.9 Refrigerators and Heat Pumps 87
4.10 Energy Overview 88
CHAPTER 2 Chapter Review 88
HIGH-ENERGY SOCIETY 20
2.1 Energy and Power 20 CHAPTER 5
2.2 Your Energy Servants 22 FOSSIL ENERGY 93
2.3 Your Energy Servants’ Jobs 25 5.1 The Origin of Fossil Fuels 93
2.4 Who Are Your Servants? 27 5.2 The Fossil Fuels 96
2.5 What Energy Servants Buy Us 28 5.3 Energy from Fossil Fuels 99
2.6 Policy Issue: Measuring 5.4 Fossil Energy Technologies 100
Quality of Life 30 5.5 Fossil Fuel Resources 111
Chapter Review 31 5.6 Will We Run Out? 117
5.7 Policy Issue: A Price on
CHAPTER 3 Carbon? 122
ENERGY: A CLOSER LOOK 34 Chapter Review 124
3.1 Forms of Energy 35
3.2 Electrical Energy: A Closer Look 37 CHAPTER 6
3.3 Quantifying Energy 43 ENVIRONMENTAL IMPACTS OF FOSSIL FUELS 128
3.4 Energy and Work 50 6.1 What’s Pollution? 128
3.5 Work and Kinetic Energy 53 6.2 Air Pollution 130
3.6 The Role of Friction 54 6.3 Other Environmental Impacts of
3.7 The Art of Estimation 54 Fossil Fuels 144
3.8 Wrapping Up 56 6.4 Policy Issue: The Clean Air Act 151
Chapter Review 57 Chapter Review 154
vii

viii / CONTENTS
10.4 Other Indirect Solar Energy Sources 287

CHAPTER 7
Chapter Review 289
NUCLEAR ENERGY 158
7.1 The Atomic Nucleus 160
7.2 Radioactivity 163 CHAPTER 11
7.3 Energy from the Nucleus 167 ENERGY CARRIERS: ELECTRICITY AND HYDROGEN 292
7.4 Nuclear Fission 169 11.1 Electricity 293
7.5 Nuclear Reactors 174 11.2 Toward a Hydrogen Economy? 308
7.6 The Nuclear Fuel Cycle and 11.3 Electric Cars 315
Uranium Reserves 180 Chapter Review 319
7.7 Environmental and Health Impacts
of Nuclear Energy 186 CHAPTER 12
7.8 Policy Issue: A Nuclear THE SCIENCE OF CLIMATE 323
Renaissance? 194 12.1 Keeping a House Warm 323
7.9 Nuclear Fusion 195 12.2 Keeping a Planet Warm 324
Chapter Review 201 12.3 In the Greenhouse 327
12.4 Earth’s Energy Balance 334
CHAPTER 8 12.5 A Tale of Three Planets 338
ENERGY FROM EARTH AND MOON 205 Chapter Review 341
8.1 The Geothermal Resource 205
8.2 Geothermal Energy Technology 209 CHAPTER 13
8.3 Environmental Impacts of FORCING THE CLIMATE 344
Geothermal Energy 214 13.1 Climate Forcing 344
8.4 Geothermal Heat Pumps 215 13.2 Climate Sensitivity 346
8.5 Tidal and Ocean Energy 220 13.3 Feedback Effects 348
Chapter Review 224 13.4 Natural and Anthropogenic
Forcings 350
CHAPTER 9 13.5 Carbon: A Closer Look 360
DIRECT FROM THE SUN: SOLAR ENERGY 227 Chapter Review 364
9.1 The Solar Resource 227
9.2 Photovoltaic Solar Energy 233 CHAPTER 14
9.3 Solar Heating 240 IS EARTH WARMING? 368
9.4 Solar Thermal Power Systems 247 14.1 Taking Earth’s Temperature 368
9.5 Other Solar Applications 251 14.2 Other Changes in Climate 378
9.6 Environmental Impacts of 14.3 Are We to Blame? 383
Solar Energy 252 Chapter Review 386
9.7 Policy Issue: Expanding
Solar Energy 253
CHAPTER 15
Chapter Review 256
FUTURE CLIMATES 389
15.1 Modeling Climate 389
CHAPTER 10 15.2 Climate Projections 396
INDIRECT FROM THE SUN: WATER, WIND, BIOMASS 260 15.3 Consequences of Global
10.1 Hydropower 260 Climate Change 400
10.2 Wind 268 15.4 Climate Change and Society 408
10.3 Biomass 276 Chapter Review 410

CONTENTS / ix
Appendix APP-1
CHAPTER 16
Glossary G-1
ENERGY AND CLIMATE: BREAKING THE LINK 414
Suggested Readings SR-1
16.1 Carbon Emissions: Where We’re
Answers to Odd-Numbered Questions
Going and Where We Need to Be 415
and Exercises ANS-1
16.2 Geoengineering 419
Credits and Data Sources CDS-1
16.3 Carbon Capture and Storage 422
Index I-1
16.4 Nonfossil Energy Sources 425
16.5 Using Less Energy 427
16.6 Strategies for a Sustainable Future 436
Chapter Review 442

FOCUS BOXES
BOX 1.1 BEING GRAPHICALLY LITERATE 16

BOX 2.1 END-USE AND PRIMARY ENERGY 26
BOX 3.1 CONVERTING UNITS 47
BOX 3.2 SI PREFIXES 48
BOX 4.1 HIGHER AND LOWER HEATING VALUES 78
BOX 4.2 COGENERATION AT MIDDLEBURY COLLEGE 85
BOX 5.1 CARBON VERSUS CARBON DIOXIDE 101
BOX 5.2 SHALE OIL, OR OIL SHALE? 116
BOX 5.3 EXPONENTIAL GROWTH 118
BOX 6.1 CHOOSE YOUR POISON: THE VW EMISSIONS SCANDAL 130
BOX 7.1 E = mc 2
171
BOX 8.1 THE GEYSERS 211
BOX 9.1 A TALE OF TWO STATES 255
BOX 12.1 A GREENHOUSE COMPLICATION 333
BOX 12.2 A BANKING ANALOGY 336
BOX 13.1 THE INTERGOVERNMENTAL PANEL ON CLIMATE CHANGE 351
BOX 14.1 THE WARMEST YEAR 371
BOX 14.2 ISOTOPIC TEMPERATURE RECONSTRUCTION 375
BOX 14.3 ICE-SHEET INSTABILITY 381
BOX 15.1 AN EARTH SYSTEM MODEL 393
BOX 15.2 MEANS AND EXTREMES 401
BOX 16.1 350: THE SCIENCE BEHIND THE NUMBER 417
BOX 16.2 NEGAWATTS: THE ROCKY MOUNTAIN INSTITUTE 432
xi

PREFACE
Many behaviors distinguish the human species from our fellow inhabitants of
Planet Earth. One of these behaviors—our use of energy in amounts far ex-
ceeding what our own bodies can produce—affects the environment in unprec-
edented ways. Centuries ago, pollution from coal burning was already a serious
urban problem. Despite regulatory and technological progress in pollution
control, diminished air and water quality continue to be major consequences
of our ever-growing energy consumption. Further environmental degradation
results as we scour the planet for fuels that contain the stored energy of which
we demand an unending supply. Our energy-intensive society also enables other
environmentally damaging developments such as sprawl, large-scale mecha-
nized agriculture, and massive deforestation. At the same time, energy brings
us higher standards of living and allows our planet to sustain a larger human
population.
In recent decades, a new and truly global impact of humankind’s energy
consumption has overshadowed the long-standing and still significant conse-
quences associated with traditional pollution, resource extraction, and energy-
enabled development. That impact is global climate change, brought about
largely by the emissions from fossil fuel combustion. Climate change is a prob-
lem that knows no national or even continental boundaries. It will affect us
all—although not all equally. It won’t have the civilization-ending impact of
an all-out nuclear war or a major asteroid hit, but climate change will greatly
stress an already overcrowded, divided, and combative world.
Achieving a healthier planet with a stable, supportive climate means either
using less energy or using energy in ways that minimize adverse environmen-
tal impacts. Here we have choices: To use less energy, we can either deprive
ourselves of energy’s benefits or we can use energy more intelligently, getting
the same benefits from less energy. To minimize environmental and especially
climate impacts, we can shift from fossil fuels to energy sources that don’t pro-
duce as much pollution or climate-changing emissions. Or we can learn how to
capture the emissions from fossil fuels and sequester them away from Earth’s
surface environment.
Earth’s energy resources are limited to a relatively few naturally occurring
stores of energy—fuels—and energy flows such as running water, sunlight,
wind, geothermal heat, and tides. A realistic grasp of our energy prospects
demands that we understand these energy resources. We need to know, first and
foremost, whether a given resource or combination of resources is sufficient to
meet humankind’s energy demand. For fuels, we need a good estimate of the
remaining resource and a time frame over which we can expect supplies to last.
xiii

xiv / PREFACE
We need to understand the technologies that deliver useful energy from fuels
and flows, to assess their environmental impacts, and to recognize that none is
without adverse effects. And we need to be realistic about the near-term and
long-term prospects for different energy sources in the economic context.
The oil shortages of the 1970s spawned a serious exploration of energy
alternatives. Governments and industries sponsored research programs, while
tax credits encouraged the installation of alternative energy systems. Vehicle
mileage and other measures of energy efficiency increased significantly. At the
same time, colleges and universities developed specialized courses in energy
issues and the relationship between energy and environment. These courses
emerged in traditional departments such as physics, chemistry, and engineer-
ing; in interdisciplinary programs dealing with technology and society; and in
the burgeoning new programs in environmental studies and environmental sci-
ence that sprang up with the emergence of a widespread environmental con-
science in the last decades of the twentieth century. Textbooks written for such
courses addressed the science and policy issues surrounding energy and the
environment.
Energy, Environment, and Climate also focuses on energy and its impact
on the environment. Unlike its predecessors, it’s built from the ground up on
the premise that climate change is the dominant energy-related environmental
issue of the twenty-first century. More traditional concerns, such as pollution
and energy resources, remain important, and they, too, are covered here. But a
full five chapters—about one-third of the book—are devoted to climate and the
energy–climate link.
Energy, Environment, and Climate begins with a survey of Earth’s history
and the origin of the planet’s energy resources. A quantitative look at past and
present patterns of human energy consumption follows, including a discussion
of the link between energy, economic development, and human well-being.
Chapters 3 and 4 provide an introduction to the science of energy, including the
all-important role of the second law of thermodynamics. Chapters 5 through
10 describe specific energy sources and their resource bases, the role each
plays in today’s global energy system, their associated technologies and pros-
pects for future technological development, and their environmental impacts.
Chapter 11 rounds out the coverage of energy with discussions of electricity
and hydrogen—both energy carriers, but emphatically not energy sources. The
energy chapters are organized around fundamental resources, including fossil
fuels, nuclear energy, geothermal and tidal energy, and direct and indirect solar
energy. Because fossil fuels dominate today’s energy supply, there are two chap-
ters dealing, first, with the fossil resource and fossil fuel technologies, and sec-
ond, with the environmental impacts of fossil fuels. Whereas other textbooks
have separate chapters on such energy-related issues as transportation, Energy,
Environment, and Climate includes these topics in the appropriate energy-
source chapters. For example, hybrid vehicles and combined-cycle power plants
appear in the fossil fuel chapters; fuel-cell and battery electric vehicles are dis-
cussed in the chapter that covers electricity and hydrogen as energy carriers;
and wind turbines are included in the chapter on indirect solar energy.

PREFACE / xv
Four chapters on climate follow the section on energy. Chapter 12 describes

the scientific principles that determine planetary climates, including the natural
greenhouse effect in the context of the planets Venus, Earth, and Mars. The
chapter ends with a discussion of the nature of scientific theories and of certainty
and uncertainty in science. Chapter 13 details the so-called forcings—both
natural and anthropogenic—that can upset the energy balance that ultimately
establishes Earth’s climate. Chapter 14 documents observations showing that
Earth is now undergoing unusually rapid climate change and explains why sci-
entists are convinced that most of that change is attributable to human activi-
ties. Chapter 15, which outlines projections of future climates, includes a look
at the workings of computer climate models and the role of climate feedbacks.
The final chapter brings together the two main themes of the book—energy and
climate—and explores how humankind might continue to enjoy the benefits of
energy use while minimizing its climate-changing impacts.
Energy, Environment, and Climate is written primarily from a scientific per-
spective. However, questions of policy and economics are never far behind the
science of energy and climate. The text therefore ventures occasionally into
policy and economic considerations—although to a far lesser extent than a
policy-oriented book would do. In particular, several chapters end with a sec-
tion specifically dedicated to a policy-related issue that grows out of the science
covered in the chapter.
Any serious study of energy and the environment has to be quantitative.
We need to understand just how much energy we actually use and how much
energy is available to us. It makes little sense to wax enthusiastic about your
favorite renewable energy source if it can’t make a quantitatively significant
contribution to humankind’s energy supply. Assessment of environmental
impacts, too, requires quantitative analysis: How much pollution does this
energy source emit? At what rate are we humans increasing the atmospheric
CO2 concentration? What’s the maximum CO2 concentration we can toler-
ate without incurring dangerous climate change? How long will nuclear waste
remain dangerous? How much waste heat does this power plant dump into
the river? How much CO2 results from burning a gallon of gasoline? What is
exponential growth and what are its consequences for future levels of energy
consumption, pollution, or carbon emissions? In dealing with such questions,
this book doesn’t shy away from numbers. At the same time, it isn’t a heav-
ily mathematical text with equations on every page. Rather, the text attempts
to build fluency with quantitative information—fluency that means being able
to make quick order-of-magnitude estimates, work out quantitative answers to
simple “how much” questions, and “read” numerical information from graphs.
The book doesn’t require higher mathematics—there’s no calculus here—but it
does demand your willingness to confront quantitative data and to work com-
fortably with simple equations. Anyone with a solid background in high-school
algebra can handle the material here. As for a science background, the text
assumes only that the reader has some familiarity with high-school-level chem-
istry and/or physics. Despite its scientific orientation, this book is written in a
lively, conversational style that students have welcomed in my other textbooks.

xvi / PREFACE
Energy, Environment, and Climate helps reinforce qualitative and quantita-

tive understandings of the text with its Chapter Reviews, which summarize
the big ideas presented in each chapter, invite you to consider the meaning of
new terms introduced in the chapter, and recap important quantitative infor-
mation and equations. Questions probe the concepts behind energy sources,
environmental impacts, and climate issues. Exercises provide practice in per-
forming calculations based on the material introduced in each chapter. Answers
to the odd-numbered Questions and Exercises are provided at the back of the
book. Research Problems send you to sources of contemporary data—usually
web-based—and allow for more detailed exploration of questions that may
be related to energy and environmental issues in your home state or country.
Given the discoveries quickly unfolding in this growing field, Research Prob-
lems may also ask you to update data presented in the book or to look more
deeply into quantitative data on global energy use and its impacts. Argue Your
Case questions ask you to formulate authoritative arguments for or against
propositions relevant to each chapter’s topics.
Energy, Environment, and Climate is illustrated with photos, line drawings,
and graphs. Line drawings describe the workings of energy technologies, the
flows of energy and materials throughout the Earth system, climate models
and feedback effects, pollution-control and waste-storage systems, and a host
of other content that’s best seen to be understood. Most of the photos show
actual energy systems, presented to give a sense of the technologies and their
scales. Graphs quantitatively describe everything from the breakdown of our
energy use by source or by economic sector to projections of future global tem-
peratures. Every graph is traceable to an authoritative source, provided in the
Credits and Data Sources at the end of the book.
Energy, Environment, and Climate deals with rapidly changing fields, and
this third edition is as up to date as possible. Nearly every graph and item of
numerical data has been updated through 2015 or later. Wind and solar energy
have seen great advances since the 2008 first edition and the 2012 second edi-
tion, including much-expanded deployment of large-scale grid-connected sys-
tems, and these developments are reflected here. Relatedly, the chapter on direct
solar energy has been reorganized to give greater emphasis to photovoltaic sys-
tems, and the indirect solar energy chapter includes new material on airborne
wind energy systems. A new section on hydraulic fracturing highlights the revo-
lutionary impact of fracking on oil and gas supplies, especially in North Amer-
ica. Another new section covers electric vehicles and compares the emerging
technologies of battery and fuel-cell vehicles. Discussions of cogeneration and
heat pumps have been expanded in light of these technologies’ increasing use.
The nuclear energy chapter includes new material on small modular reactors
and alternative approaches to nuclear fusion, as well as incorporating the latest
studies on the impacts of the Chernobyl and Fukushima accidents. Our under-
standing of climate change has advanced with the publication in 2013–2014
of the Intergovernmental Panel on Climate Change’s Fifth Assessment Report,
and the climate chapters have been appropriately updated with material from
IPCC 5 or later. Material on energy and climate policy and legislation, both

PREFACE / xvii
national and international, has been thoroughly updated; examples include an

expanded discussion of renewable portfolio standards and a look at the Paris
Climate Agreement that took effect in 2016. The text also describes new studies
suggesting ways to meet the climate challenge, including those from the Carbon
Mitigation Initiative and the Deep Decarbonization Pathways Project. Finally,
there’s a new section relating food, energy, and climate.
Other enhancements to the third edition include four new focus boxes
on diverse topics: the Volkswagen emissions scandal, the policies of differ-
ent U.S. states on photovoltaic solar energy, the difference between shale oil
and oil shale, and ice-sheet instability. The third edition continues the sec-
ond edition’s trend toward more international emphasis. However, the book
remains especially relevant to students in North America. For example, energy
data are often presented for both the entire world and for the United States,
while the new material on fracking for oil and gas emphasizes North America.
And policy considerations focus on such matters as the United States’ Clean
Air Act or British Columbia’s carbon tax. The Chapter Review Questions and
Exercises have been revised in light of instructor and student suggestions, and
the number of Exercises and Research Problems has increased.
Also complementing the main text are tables displaying important energy-
and climate-related quantities; some of the most useful of these also appear
inside the covers. An appendix tabulates relevant properties of materials, rang-
ing from R values of building materials to half-lives of radioactive isotopes to
global warming potentials of greenhouse gases. A Glossary defines all key terms
that appear in the book and includes acronyms as well as symbols for physi-
cal units and mathematical quantities. A list of Suggested Readings, includ-
ing authoritative web sites, is also provided. In addition, instructors teaching
from Energy, Environment, and Climate will find supplementary resources at
wwnorton.com/instructors, a password-protected web site that includes the
figures and tables appearing throughout the text, as well as an instructor’s man-
ual with solutions.
Energy, Environment, and Climate is not a book of environmental advo-
cacy or activism; it’s much more objective than that. I have my own opinions,
and I acknowledge that many—although not all—are in line with the views
of the broader environmental movement. But I pride myself on independent
thinking based on my own study of others’ writings and research, and I’d like
to encourage you to do the same. I’m also keenly aware that there’s a much
stronger scientific consensus on some issues, particularly climate change, than
the popular media, the general public, and even some governments—including
the current leadership of the United States—may acknowledge. I’ve been care-
ful to base my scientific statements on the consensus of respected scientists and
on peer-reviewed literature that’s available to you and everyone else for direct
examination. At the same time, I understand the uncertainties inherent in sci-
ence, especially in an area as complex as the interconnected workings of the
global environment. I openly state those uncertainties and quantify them when-
ever possible. That said, I would be pleased if the knowledge you gain from this
book inspires you to work toward change in our collective patterns of energy

xviii / PREFACE
consumption. I and the majority of my fellow scientists are convinced that such
actions are essential in the coming decades if we’re to avoid disruptively harm-
ful environmental impacts.
ACKNOWLEDGMENTS
No individual can be an expert on all the topics covered in a book like this one,
and during the writing process I’ve been fortunate to be able to call on special-
ists in many fields. They’ve contributed to making this book more authorita-
tive and timely than I, working alone, could have done. With appreciation, I
acknowledge the individuals who have given their expert opinion, read drafts
of individual chapters, or otherwise contributed advice and encouragement to
this project:
Climate experts Gavin Schmidt (NASA Goddard Institute for Space Stud-
ies) and Michael Mastrandrea (Stanford University) reviewed the climate
chapters and made many helpful suggestions. Dr. William Glassley (Lawrence
Livermore National Laboratory and California Energy Commission) reviewed
the sections on geothermal energy in Chapter 8; Dr. JoAnn Milliken (Acting
Program Manager, U.S. Department of Energy Hydrogen Program) reviewed
Chapter 11; and Roger Wallace (Vermont Wood Energy) and Greg Pahl (Ver-
mont Biofuels Association) reviewed sections of Chapter 10 on biomass. Oth-
ers who offered advice include Dr. William Ruddiman (University of Virginia),
Dr. Irina Marinov (University of Pennsylvania), the late Dr. Stephen Schnei-
der (Stanford University), Dr. Michael Mann (Pennsylvania State University),
Dr. Peter Vitousek (Stanford University), Dr. Robert Romer (Amherst College),
Dr. Mark Heald (Swarthmore College), Dr. Gary Brouhard (McGill Univer-
sity), Elizabeth Rosenberg (Argus Media), Dr. Eric Rignot (Jet Propulsion
Laboratory), and George Caplan (Wellesley College). Dr. James Williams of
the Monterey Institute provided a thorough review of Chapter 11’s material on
electricity. My Middlebury colleagues Sallie Sheldon (biology), Steve Sontum
(chemistry), Jon Isham (economics), Jeff Munroe (geology), Chris Watters (biol-
ogy), Bill McKibben (environmental studies), Grace Spatafora (biology), and
Molly Costanza-Robinson (chemistry and environmental studies) were kind
enough to share their expertise and encouragement. The new food–energy–
climate material in Chapter 16 was inspired by Barnaby Feder and Paul Stone
and was reviewed by my Middlebury/Monterey colleagues Molly Anderson
and Jason Scorse, while Middlebury political scientist Kemi Fuentes-George
reviewed the new coverage of the meetings that led to the Paris Agreement.
Finally, I thank my former student Peter Mullen for a thorough reading of the
manuscript, and I thank both Peter and Wendy Mullen for their support of this
and other projects.
In addition to those acknowledged above, I am grateful to the following
instructors of energy and/or climate courses who contributed reviews at the
request of W. W. Norton. Their comments, many based on instructor and student
experiences with the first and second editions, offered a blend of pedagogical and

PREFACE / xix
scientific expertise that has enhanced the readability, teachability, and author-
ity of this textbook. In addition to these formal reviews, I’ve benefited from
ongoing correspondence with a number of instructors using the book, especially
Daniel Prober (Yale University) and Mark Rosenberry (Siena College).
FIRST EDITION REVIEWERS

Cecilia Bitz (University of Washington)
Robert L. Brenner (University of Iowa)
F. Eugene Dunnam (University of Florida)
Dorothy Freidel (Sonoma State University)
Jonathan P. Mathews (Pennsylvania State University)
James Rabchuk (Western Illinois University)
Ljubisa R. Radovic (Pennsylvania State University)
Sunil V. Somalwar (Rutgers University)
SECOND EDITION REVIEWERS

Ani Aprahamian (University of Notre Dame)
Anand Balaraman (Georgia Southern University)
Mona Becker (McDaniel College [Western Maryland College])
David Branning (Trinity College)
Andrew J. Friedland (Dartmouth College)
Douglas Kurtze (St. Joseph’s University)
Gaytha Langlois (Bryant College)
Qi Lu (St. John’s University)
David Marx (Illinois State University)
Gary Pajer (Rider University)
Zoran Pazameta (Eastern Connecticut State University)
Doug Pease (University of Connecticut)
Alvin Saperstein (Wayne State University)
J. Scofield (Oberlin College)
S. Ismat Shah (University of Delaware)
Sam Tanenbaum (Claremont McKenna College)
Scott Wissink (Indiana University Bloomington)
Eric Woods (State University of New York at Albany)
THIRD EDITION REVIEWERS

Willem H. Brakel (American University)
Tom Carter (College of DuPage)
Ursula Gibson (Norwegian University of Science and Technology)
Richard Gran (University of Minnesota Duluth)
John Hassard (Imperial College London)
Barry Ness (Lund University)

xx / PREFACE
Finally, thanks to EJ Zita (Evergreen State College) for her efforts in check-
ing answers to the end-of-chapter exercises and preparing the online Solutions
Manual for the second edition of this text; much of her work has carried over
to the third edition.
I’m honored to be publishing this book with W. W. Norton, and I am
indebted to former Norton editor Leo Wiegman for inviting me to write the
first edition of this textbook. Inspired in part by this project, Leo left publishing
to start his own environmental work; this third edition was in the able hands of
editor Eric Svendsen and his assistants Rachel Goodman and Lindsey Osteen.
Project editor Carla Talmadge and copyeditor Norma Sims Roche spearheaded
an efficient and successful production process. I am grateful for all their efforts,
and it has been a pleasure to work with all of them.
Finally, I thank my family for their support and patience through the long
process of bringing this project to fruition.
ABOUT THE AUTHOR

Richard Wolfson is Benjamin F. Wissler Profes-
sor of Physics at Middlebury College, where he
also teaches environmental studies. He holds
a B.A. in physics and philosophy from Swarth-
more College, an M.S. in environmental studies
from the University of Michigan, and a Ph.D. in
physics from Dartmouth. His research involves
solar astrophysics and terrestrial climate, and
he has published nearly 100 papers in the sci-
entific literature. Wolfson’s other books include
several introductory physics texts as well as
Nuclear Choices: A Citizen’s Guide to Nuclear
Technology (1993) and Simply Einstein: Rela-
tivity Demystified (2003). He has produced five video courses for The Teaching
Company’s Great Courses series: Einstein’s Relativity and the Quantum Revo-
lution: Modern Physics for Nonscientists (1999); Physics in Your Life (2004);
Earth’s Changing Climate (2007); Physics and Our Universe: How It All Works
(2011); and Understanding Modern Electronics (2014).

ENERGY,
ENVIRONMENT,
AND CLIMATE
Third Edition

Chapter 1
A CHANGING PLANET
Earth was born some 4.6 billion years ago, and our planet has been changing
ever since. Earth’s evolution is driven by an interplay between matter—the
physical “stuff” that makes up the Universe—and energy, an equally impor-
tant universal “stuff” associated with motion, heat, and the fundamental forces
of nature. It’s energy that makes everything happen; without energy, the Uni-
verse would be a static, unchanging, lifeless place. In Earth’s case, agents of
change are astrophysical, geological, chemical, and biological. Astrophysical
events formed our planet and occasionally alter its history. Geological events
build mountains and wear them down, move continents, shake the solid
Earth, and spew gases into the atmosphere. Chemical reactions change the com-
position of rocks, soils, atmosphere, and oceans. Life appeared on Earth billions
of years ago, and soon biological processes were radically altering the planet’s
atmosphere and chemistry. Hundreds of millions of years ago, life emerged
from the oceans to colonize the land. Just a few million years ago, our human
species evolved and began the process of anthropogenic (i.e., human-caused)
environmental change. We’ve since become sufficiently plentiful and techno-
logically advanced that we’re now having a global impact on Planet Earth.
1.1 Earth’s Beginnings

The time is some 4.6 billion years ago; the place, a vast cloud of interstellar
gas and dust about two-thirds of the way out from the center of the Milky
Way galaxy. Most of the material in the cloud is hydrogen and helium, the lat-
ter having formed in the first 30 minutes after the Universe began in a colossal
explosion we call the Big Bang. But there are smaller amounts of oxygen, carbon,
nitrogen, silicon, iron, uranium, and nearly all the other elements. These elements
were formed by nuclear reactions in massive stars that exploded several billion
years earlier and spewed their contents into the interstellar medium.
Gravitational attraction among the gas and dust particles that make up the
cloud causes it to shrink, and—like ice skaters who approach, join hands, and
spin—the shrinking cloud begins to rotate. As it rotates, it flattens into a disk,
with all the matter in essentially the same plane. This collapse is remarkably
rapid, taking only about 100,000 years.
A massive accumulation develops at the disk’s center, and under the crushing
pressure of gravitational attraction, its temperature rises. Eventually the central
235693_01_001-019_r2_gk.indd 1 05/05/17 1:39 PM

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with in other places near London; it is in this country an introduced species, but is
found in many parts of the continent. It is a land-leech, and lives upon earthworms.
The genus Haemadipsa, which M. Blanchard places in a special sub-family,

contains a number of species which are for the most part land-leeches. Land-
leeches occur in many parts of the world, but chiefly in the tropics—in India, Ceylon,
Java, South America, etc. They lie in wait for their prey, upon the ground as a rule;
but they may ascend herbs and shrubs to gain a better outlook when they are aware
of an approaching footstep. A vivid account of the ferocity of these tiny Annelids in
Ceylon can be read in Sir J. E. Tennent's Natural History of Ceylon. They have been
said to be so pugnacious and so poisonous that persons surprised in their sleep by
the pests have succumbed to their united efforts. A whole battalion of English
soldiers decamped on one occasion from a wood which was overflowing with land-
leeches. The familiar misquotation "lethalis hirudo" might well be applied to this
species. Professor Whitman has written much upon the habits of the land-leech of
Japan (Haemadipsa japonica), which bites so softly that its presence cannot be
detected except for the stream of blood which trickles from the wound. While it is
feeding it emits from the pores of the nephridia a clear fluid, which, as it appears, is
used to keep the skin moist; when unduly dried the same phenomenon occurs. It is
curious that in this and other leeches the nephridia should play a part which in the
earthworm is played by the dorsal pores; in both animals the glands of the skin are
also concerned with the same duty.
The purely aquatic leeches swim by undulations, and also crawl by the help of the
two suckers, like a "Geometer" caterpillar. But when a land-leech is dropped into the
water it at once sinks to the bottom and crawls out; it does not swim, but can survive
immersion for a long period. In this it resembles the earthworms, which can also
survive a prolonged immersion, and even in the case of some are indifferent to the
medium, land or water, in which they live; the land-leech, however, is entirely
dependent upon damp surroundings; a dry air is fatal to it. The land-leech of Japan
leaves a slimy trail behind it as it crawls, in this respect recalling the land Planarian
Bipalium kewense.
GEPHYREA AND PHORONIS
BY
ARTHUR E. SHIPLEY, M.A.

Fellow and Tutor of Christ's College, Cambridge
CHAPTER XV
GEPHYREA
INTRODUCTION—ANATOMY—DEVELOPMENT—SIPUNCULOIDEA—PRIAPULOIDEA—
ECHIUROIDEA—EPITHETOSOMATOIDEA—AFFINITIES OF THE GROUP.
The animals included in the above-named group were formerly associated with the
Echinodermata. Delle Chiaje[468] states that Bohadsch of Prague in 1757 was the
first to give an accurate description of Sipunculus under the name of Syrinx, but
Linnaeus, who noted that in captivity the animal always kept its anus directed
upwards, re-named it Sipunculus. Lamarck[469] placed the Gephyrea near the
Holothurians; and Cuvier[470] also assigned them a position amongst the
Echinoderms. He mentions Bonellia, Thalassema, Echiurus, Sternaspis, and three
species of Sipunculus, one of which, S. edulis, "sert de nourriture aux Chinois qui
habitent Java, et qui vont la chercher dans le sable au moyen de petits bambous
préparés."
The name Gephyrea[471] was first used by Quatrefages, who regarded these
animals as bridging the gulf between the Worms and the Echinoderms. He included
in this group the genus Sternaspis (vide p. 335), now more usually classed with the
Chaetopoda.
The Gephyrea are exclusively marine. They are subcylindrical animals, which can
either retract the anterior end of their body—the introvert—carrying the mouth into
the interior; or are provided with a long flexible but non-retractile proboscis. The
latter is easily cast off. They usually bear spines or hooks of a hard chitinous
character, secreted by the epidermis or outermost layer of cells. The mouth is at the
base of the proboscis or at the end of the protractile part, the anus is at the other
end of the body or on the dorsal surface. The nervous system consists of a ring
round the mouth and of a ventral nerve-cord. A vascular system is present as a rule.
Nephridia are found which act as excretory organs, and in most cases also as ducts
for the generative cells. The Gephyrea are bisexual, and the male is sometimes
degenerate.
The group may be divided into four Orders:—(i.) Sipunculoidea; (ii.) Priapuloidea;
(iii.) Echiuroidea; (iv.) Epithetosomatoidea; of these the first is by far the largest,
both in number of genera and of species.
The Anatomy of Sipunculus nudus.
External Characters.—The body of S. nudus when fully extended may attain a

length of a foot, or even a little more; in this condition it is seen to consist of two
portions, the anterior of which is, however, retracted into the other when the animal
is disturbed. The retractile portion is sometimes termed the proboscis, but as its
nature is entirely different from that of the proboscis of the Echiuroidea, it is better to
refer to it as the introvert. Special retractor muscles are attached on the one hand to
the body-wall about half-way down the body, and on the other hand are fused into a
muscular sheath which surrounds the gullet, just behind the mouth. When these
muscles contract, they withdraw the introvert into the rest of the body or trunk in
much the same way as the finger of a glove may be drawn into the hand, by a
thread fastened to the inside of its apex. The introvert is protruded by the
contraction of the circular muscles of the body-wall. These exert a pressure on the
fluid which fills the body-cavity, and by this means the sides of the introvert are
forced forward until finally the head is exposed.
The introvert occupies about one-sixth or one-fifth of the total body length. It is
somewhat narrower than the trunk, and is covered by a number of small flattened
papillae, some of which lie with their free ends directed backward, overlapping one
another like tiles on a roof. In some other genera, as Phymosoma, the introvert
bears rows of horny hooks, which are apt to fall off as the animal grows old.
The trunk has from thirty to thirty-two longitudinal furrows, the elevations between
which correspond with a similar number of muscles lying in the skin. This
longitudinal marking is crossed at right angles by a circular marking of similar origin,
the elevations of which correspond with the circular muscles in the skin. These two
sets of markings thus divide the skin of the trunk into a number of small square
areas, very regularly arranged (Fig. 212).
The outline of the trunk is more or less uniform, but it is capable of considerable
change according to the state of contraction of its muscles. The circular muscles, for
instance, may be contracted at one level, thus causing a constriction at this spot.
The colour of S. nudus is a somewhat glistening greyish-white.
Fig. 211.—Right half of the anterior end of Sipunculus nudus L., seen from the inner
side and magnified. a, Funnel-shaped grooved tentacular crown leading to the
mouth; b, oesophagus; c, strands breaking up the cavity of the tentacular crown
into vascular spaces; c', heart; d, brain; e, ventral, and e', dorsal retractor
muscles; f, ventral nerve-cord; G, vascular spaces in tentacular crown.
The anterior end of the fully-expanded Sipunculus may be termed the head; here
the skin is produced into a frayed fringe which stands up in the shape of a funnel
round the mouth. This fringe is grooved on its internal surface with numerous little
gutters, all of them lined with cilia, which by their constant motion keep up a current
which sweeps food into the mouth. The fringe may be in the form of a simple ring
round the mouth, or the ring may be folded in at the dorsal side so as to take the
form of a double horse-shoe (Figs. 211 and 212).
Body-wall.—The glistening appearance of Sipunculus is due to the cuticle, a

chitinoid layer which is secreted by the external layer of cells, the epidermis.
Beneath this lies a layer of connective tissue, which is not always present in other
Gephyrea; within this lies a layer of circular muscles arranged in bundles, then
comes a very thin sheath of oblique muscular fibres, then a thicker layer of
longitudinal muscles, and finally a layer of peritoneal epithelial cells, which in
Sipunculus are for the most part ciliated.
Scattered over the surface of the body, and opening by narrow tubes which pierce
the cuticle, are a number of glandular bodies which may be either bi- or multi-
cellular. The glandular cells are apparently enlarged and modified epidermal cells;
they are arranged in a cup-shaped manner, with their apices directed towards the
orifice. They are crowded with granules, which are presumably poured out over the
cuticle, but the exact function of the secretion is entirely unknown. They have a well-
developed nerve supply.
Digestive System.—The mouth lies in the centre of the fringe, and is not provided
with any kind of jaw or biting armature; it leads directly into the thin-walled
alimentary canal, the first part of which is ciliated. The alimentary canal is not
marked out into definite regions, but passes as a thin-walled semi-transparent tube
to the posterior end of the body, and then turns forward again and opens to the
exterior by an anus situated about an inch below the junction of the introvert with the
trunk, on the median dorsal line. The descending and ascending limbs of the
alimentary canal are coiled together in a spiral, which may be more or less close in
different individuals. The whole is supported by numerous fine muscular strands,
which pass from the walls of the intestine to the skin, and by a spindle-muscle,
which runs from the extreme posterior end of the trunk up the axis of the spiral and
terminates in the skin close to the anus.
No glands open into the alimentary canal at any point of its course, but near the
anus a simple diverticulum, or pocket, of unknown function arises. The size of this
outgrowth differs enormously in different individuals. The alimentary canal near the
anus also bears two tuft-like organs, which, however, do not open into the intestine,
but probably have some function in connexion with the fluid in the body-cavity.
Along the whole course of the alimentary canal there runs a ciliated groove, into
which the food does not pass, but the cilia of which probably keep in motion a
current of water whose function may be respiratory.
Fig. 212.—Sipunculus nudus L., with introvert and head fully extended, laid open by
an incision along the right side to show the internal organs. × 2. a, Mouth; b,
ventral nerve-cord; c, heart; d, oesophagus; e, intestine; f, position of anus; g,
tuft-like organs; h, right nephridium; i, retractor muscles; j, diverticulum on
rectum. The spindle-muscle is seen overlying the rectum.
Vascular System.—On the dorsal surface of the anterior end of the alimentary
canal lies a contractile vessel, usually termed the heart. It is a tube about an inch
long, ending blindly behind, but opening in front into a ring-shaped space
surrounding the mouth and partially enveloping the brain. From this ring-like vessel
numerous branches are given off which pass into the fringe round the mouth, and
probably the chief function of the heart is by its contraction to force fluid into this
fringe, and so to extend it. The heart contains a corpusculated fluid. A similar but
shorter tube is found on the ventral surface of the anterior end of the alimentary
canal in the species in question; it also opens into the ring which surrounds the
mouth.
Respiratory System.—There are no special respiratory organs, and it has long

been a matter of dispute where the respiration of Gephyrea is carried on. The
oxygenation of the blood probably takes place to some extent through the walls of
the oral fringe, but the blood which receives its oxygen at this spot is limited in its
distribution, and could only supply the brain and head. It seems probable that the
remaining organs are supplied with oxygen by the fluid of the body-cavity, which
bathes them on all sides. This might obtain its oxygen from the blood in the heart, or
more probably, through the thin walls of the intestine, from the stream of water
which is maintained by the ciliated groove described above. Quite recently a form—
S. mundanus, var. branchiata—has been described[472] with thin-walled papillae
covering parts of the skin. These papillae are full of corpuscles, and are regarded by
their discoverer as branchiae.
Body-Cavity.—The pinkish fluid of the body-cavity contains numerous corpuscles,

the products of the reproductive organs (either ova or spermatozoa), and some
curious unicellular bodies known as "urns." The latter are shaped like a bowl with a
ciliated rim, and are formed from the budding of certain cells on the walls of the
dorsal blood-vessel.[473] Their function is unknown, but they resemble certain
multicellular bodies found in the body-cavity of Phascolosoma. The generative cells
found in the body-cavity are further considered below. The true corpuscles are
either biconcave round corpuscles coloured with a chemical substance, the
haemerythrin of Krukenberg, which apparently plays the same rôle as haemoglobin
in other animals; or amoeboid corpuscles, which, though rare in Sipunculus, are
very numerous in Phascolosoma.
Nervous System.—The nervous system of Sipunculus consists of a brain or

cerebral ganglion, a circumoesophageal ring surrounding the gullet, and a ventral
nerve-cord. The brain is a small bi-lobed nervous mass situated on the dorsal
surface of the oesophagus, in the angle between the right and left dorsal retractor
muscles close to their point of insertion. Numerous nerves arise from it, and pass to
the fringe surrounding the mouth and to neighbouring parts. At the sides, the brain
is continued into two stout nerve-cords which encircle the oesophagus, and
meeting, fuse together in the median ventral line to form the ventral nerve-cord (Fig.
211). The latter is of the same diameter throughout, and shows no signs of
segmentation; it is oval in section, and consists of small ganglion cells heaped up on
the ventral surface, i.e. next the skin, and of numerous fibres situated dorsally. The
cord gives off many nerves, which usually arise in pairs. These pass into the skin,
and forming rings, run round the body, and give off finer nerves as they go.
The nerve-cord is supported by numerous strands of muscle which pass to it from

the skin. These are especially long in the region where the introvert joins the trunk,
and thus allow free play to the nerve-cord when the former is being protruded or
retracted.
Sipunculus is not well provided with sense-organs, but in an animal which lives
buried in sand we should not expect to find these very highly developed. On the
introvert there are certain patches of epithelium bearing long stout cilia, which have
been regarded as tactile in function, and there is a tubular infolding reaching the
brain, which almost certainly has some sensory function. Ward[474] has termed this
"the cerebral organ." It consists of a duct lined with ciliated cells, which opens to the
exterior in the middle dorsal line outside the tentacular fringe. The duct leads down
to the brain, and expands at its lower end into a saucer-shaped space, covering that
portion of the brain where its substance is continuous with the external epithelium.
In Phymosoma this cavity is produced into two finger-shaped processes, which are
sunk into the brain and are lined by cells crowded with a dense black pigment.[475]
They are probably rudimentary eyes, perhaps distinguishing only between darkness
and light. The pits appear to be absent in Sipunculus nudus, but Andrews states
they are found, although without pigment, in S. gouldii.[476]
Excretory System.—The excretory organs or "brown tubes" are typical nephridia,

that is to say, they consist of tubes with glandular walls which open on the one side
to the exterior, and on the other by means of a ciliated funnel-shaped opening into
the body-cavity. In Gephyrea one wall of the tube is produced into a long
diverticulum or sac which hangs down into the body-cavity, and is usually supported
by muscle-fibres running to the body-wall. The lower end of the sac is broken up
into a number of crypts or pits, lined by large glandular cells crowded with brown
pigment. The pigment-granules are secreted into the cavity of the sac, and leave the
body through the external opening; they probably consist of the nitrogenous excreta
of the animal. The upper end of the sac, into which both the external and internal
orifices open, is usually enlarged, and its walls are very muscular. As in so many
other animals, the nephridia serve as ducts through which the reproductive cells
leave the body of the parent.
Reproductive System.—The Gephyrea are bisexual. In Sipunculus the testes and
ovaries are found in the same position in the two sexes, and are indistinguishable
without microscopic investigation. They each consist of small ridges situated at the
lower end of the ventral retractor muscles, just where the latter take their origin from
the longitudinal muscles of the skin. At this level the cells which line the body-cavity
on the inside of the skin are heaped up, and become modified in the one case into
ova or eggs, and in the other into the mother-cells of the spermatozoa. This method
of forming the reproductive organs from modified cells lining the body-cavity is very
common in the higher animals; but it is seen in its simplest and least modified form
in the Sipunculidae.
The eggs break away from the ovary in a very undeveloped condition, but whilst
floating about in the body-cavity they increase in size and secrete a thick membrane
around them. They have a well-marked nucleus, and are oval in outline.
The mother-cells of the spermatozoa also break away in an immature condition, and
complete their development in the nutritive fluid of the body-cavity. They divide into
a number of spermatozoa, usually eight or sixteen, which remain in contact. They
each develop a tail, which projects outwards, and aids the cluster in swimming
along. These clusters of spermatozoa are about the same size as the ova of the
female, and, like them, make their way into the "brown tubes." The exact way in
which this is accomplished is not very clear, but the cilia on the funnel-shaped
internal opening of the tube seem to have some power of selecting the generative
cells when they come within their reach, and of passing them on, whilst they reject
the much smaller corpuscles of the perivisceral fluid, which are never found in the
nephridia.[477] Once inside the internal opening, the clusters break up and the
spermatozoa escape singly into the sea. Here they meet with and fertilise the eggs
which have escaped from the body of the female.
Fig. 213.—Larva of Sipunculus nudus L. × 150. (After Hatschek.) a, Mouth; b, anus; c,

excretory organ; d, glandular appendage of oesophagus; e, wall of stomach over
which the retractor muscle runs; f, invaginated sense-organ at aboral pole.
Development.—Hatschek,[478] who investigated the development of Sipunculus
nudus at Pantano, an inlet of the sea near Messina, states that the spawning takes
place during the night, and ceases about July 10. The rate of development depends
upon the temperature, but the larvae usually free themselves from the egg-
membrane during the third day. When hatched the embryos lengthen out a good
deal, and take the form represented in Fig. 213. The larva swims actively by means
of a ring of stout cilia, which encircle the body just behind the mouth. Other shorter
cilia are found on the head, continuing into the lining of the mouth, and a little bunch
of them is situated at the extreme posterior end. The alimentary canal is already
formed, and is twisted, so that the anus lies dorsally, but not so far forward as it
does in the adult. A glandular structure opens into the mouth, and another body of
unknown function is connected with the oesophagus; both these disappear during
larval life. A pair of excretory tubules, the forerunners of the brown tubes, are found,
and the chief muscle tracts are already established. The nervous system is still in
close connexion with the skin, from the outer part of which it is derived; the cerebral
thickening bears two eye-spots.
The fluid of the body-cavity contains corpuscles, which are kept in active circulation
by the constant contractions of the body-wall, and by numerous tufts of cilia which
are borne on the inner surface of the skin. The dorsal blood-vessel is one of the
latest organs to arise.
The larva swims actively about for a month, during which time it increases greatly in
size; it then undergoes a somewhat sudden metamorphosis. The ciliated ring and
the structures related to the oesophagus begin to disappear, the distinction between
the head and the rest of the body is obliterated, and the head becomes relatively
small. The mouth changes its position, and becomes terminal instead of being
somewhat ventral, and the tentacular membrane begins to appear. At the same time
the larva relinquishes its free-swimming life, and sinks to the bottom; it begins
creeping amongst the sand by protruding and retracting the anterior part of its body,
and takes on all the characters and habits of the adult.
I. Order Sipunculoidea.
Besides the genus Sipunculus, the Order Sipunculoidea includes ten other genera.
A key to these, taken for the most part from Selenka's admirable monograph, is
given on page 424.
Phascolosoma contains, in comparison with Sipunculus, only small species, and it is

easily distinguished by the fact that the longitudinal muscles are fused into a
continuous sheath. As a rule the skin is smooth. A few species bear hooks, which
are generally scattered irregularly and not arranged in transverse rows, as in
Phymosoma (Fig. 214) and most of the other genera.
The fold which in S. nudus surrounds the mouth may be in the same species bent in
so as to take the form of a double horse-shoe, the opening of which is always
dorsal, just above the brain; in this case the mouth is crescentiform. In other genera
the fold is broken up into discrete tentacles, and these are variously arranged; in
Dendrostoma they are grouped together in four or six bundles round the mouth, but
the more usual arrangement is the horse-shoe-like row of tentacles which overhang
the crescentiform mouth, as in Phymosoma and some species of Aspidosiphon.
The ventral side of each tentacle is grooved and ciliated, and the grooves are
continued into the ciliated mouth. Their dorsal surface is pigmented, and in the
hollow of the horse-shoe lies a deeply pigmented epithelium covering the brain.
A blood-vessel courses up each tentacle, and usually two channels return the blood
to the vascular ring which surrounds the mouth. In those forms which possess
tentacles on the dorsal side of the mouth only, the ventral part of the vascular ring
lies in the lower lip, which is tumid and swollen. The brain supplies a nerve to each
tentacle.
When the introvert is retracted the tentacular ring is withdrawn and to some extent
collapsed; in this condition it would be almost touching the rough external surface of
the introvert. In some species of Phymosoma the delicate appendages of the head
are guarded from the hooks on the introvert by a thin membrane or collar,[479] which
completely ensheaths the retracted head.
Fig. 214.—A, Phymosoma granulatum F. S. Leuck. × 2. B, Head of the same. × 4. a,

Pigmented pit leading to brain. The crescentiform mouth on the lower side of the
figure is overhung by the tentacles.
When the introvert is fully extended the dorsal blood-vessel contracts and sends its
blood forward into the vascular ring, and thence into the tentacles or tentacular fold,
which are thus erected. In several species of Sipunculus, as S. nudus, S.
norvegicus, S. robustus, S. tesselatus, there is a ventral blind tube as well as a
dorsal, into which the blood is withdrawn when the head is retracted. In many other
species in various genera, such as Phymosoma weldonii and Ph. asser,
Dendrostoma signifer, S. vastus, the lumen of the dorsal vessel is increased by
numerous hollow blind processes which it bears, hanging freely into the body-cavity.
Three very small genera of Sipunculids—Onchnesoma, Petalostoma, and Tylosoma
—are devoid of all trace of vascular system and of tentacles; the mouth opens in the
centre of the anterior end of the introvert. In Onchnesoma the dorsal part of the lip is
somewhat produced, so that the head has somewhat the shape of a Doge's cap,
and in Petalostoma there are two leaf-like processes of the body-wall which guard
the mouth.
The extent to which the intestine is coiled varies very much even in the same
species; the axis of the coil is often supported by a spindle-muscle, but this is
sometimes absent. The caecum, which opens into the rectum of S. nudus, is again
a very variable structure, and when it is present varies remarkably in size.
The food of Sipunculids seems to consist almost entirely of sand, and their only
nourishment must be such small microscopic organisms or particles of animal and
vegetable débris as are to be found mixed with the sand. The alimentary canal is, as
a rule, quite full of sand, and yet in spite of the tenuity of its walls they never seem
to be ruptured. If the contents of the digestive tube be washed out with a pipette, it
will be found that it requires considerable force to dislodge many of the sand-
particles lying next the wall. These are more or less embedded in crypts or pockets
of the wall, and as the sand passes along the intestine they probably serve as more
or less fixed hard points, against which the sharp edges of the sand particles are
worn off. Amongst the sand are usually to be found pieces of shell, sometimes with
a diameter equal to that of the alimentary canal; these are usually rounded, but their
angles may have been removed by attrition before they entered the mouth of the
Sipunculid.
In S. tesselatus the sand is to some extent held together by a mucous deposit; in

those cases where there is no sand in the intestine, there is always a coagulum of
mucus, and the walls are contracted and thick; when full of sand the walls are
tensely stretched and very thin. This thinness of the wall of the alimentary canal
seems ill-adapted to a diet of sand, nevertheless it is also met with in other great
sand-eating groups of animals, such as the Echinids and the Holothurians.
The enormous amount of sand and mud which passes through the bodies of the
Sipunculids shows that they must take a considerable part in modifying the mineral
substances which form the bottom of the sea. Just as earthworms, as shown by
Darwin, play a considerable rôle in the formation of soil, so must these animals, in
conjunction with Echinids and Holothurians, effect considerable modifications in the
sand and mud which pass through their bodies. Mr. J. Y. Buchanan[480] is "led to
believe that the principal agent in the comminution of the mineral matter found at the
bottom of both deep and shallow seas and oceans, is the ground fauna of the sea,
which depends for its subsistence on the organic matter which it can extract from
the mud." The minerals at the bottom of the sea are exposed to a reducing process
in passing through the bodies of the animals which eat them, and subsequently to
an oxidising process due to the oxygen dissolved in the sea-water acting on the
minerals extruded from the animals' bodies.
The rate at which the sand passes through the body of Sipunculus is unfortunately
unknown, but that at any one moment a considerable quantity is contained in the
intestine is shown by the fact that the average weight of five specimens of S. nudus
from Naples, taken at random, was 19.08 grms., whilst the average weight of sand
washed out of their alimentary canal was 10.03 grms. The sand contained in five
other specimens of the same species measured respectively 6 c.c., 7 c.c., 6.5 c.c.,
7.5 c.c., and 7.5 c.c., giving an average of 6.9 c.c. for each individual.
Onchnesoma and Tylosoma have only one retractor muscle; Aspidosiphon and
Phascolion have, as a rule, two; Phymosoma and Sipunculus have four, and
perhaps this is the more usual number.
Phascolion, Tylosoma, and Onchnesoma have but one "brown tube"; in Phascolion
this is the right, in Onchnesoma it is sometimes the right and sometimes the left that
persists. Most other genera retain two, but there are many exceptions; for instance,
Phascolosoma squamatum has but one, and so has Aspidosiphon tortus, and in
both cases it is that of the left side. No Sipunculid has more than two. It has been
pointed out by Selenka that those species which have but one brown tube are, as a
rule, inhabitants of tubes or shells, and do not move actively about in the sand.
The eggs of all members of the family, with the exception of the genus Phymosoma,
are spherical, but those of the last-named genus are elliptical. They are always
surrounded by a thick membrane, the "zona radiata," pierced by numerous pores.
Aspidosiphon (Fig. 215) is easily recognised by the presence of two symmetrically-

arranged cuticular shields, one at each end of the trunk. These are formed by the
fusion of minute cuticular plates, such as exist in the skin of most Sipunculids. The
posterior shield is radially symmetrical, but the anterior is somewhat like the shell of
a Pecten, and symmetrical only about one plane. The introvert is protruded from the
acute angle of the anterior shield, and when extended lies almost at right angles to
the trunk, instead of being, as is usually the case, in the same straight line with it. In
many specimens, and these seem as a rule to be the older ones, a deposit of
calcium carbonate takes place over these shields, covering over and concealing
their external markings.
Cloeosiphon (Echinosiphon) has a calcareous ring, consisting of four or five rows of
lozenge-shaped calcareous bodies forming a close mosaic, arranged round the
base of the introvert, which when extended is in the same straight line as the trunk.
Each piece bears a brown spot, which is said to be the pore of a gland (Fig. 217).
Golfingia Lankester, has a cylindrical horny thickening at the anterior end of the
trunk and another at the posterior.
Fig. 215.—Aspidosiphon truncatus Kef. × 2. a, Introvert partially extended, but not

sufficiently to show the head.
Key to the Genera of Sipunculoidea.[481]
I. The longitudinal muscles in the body-wall divided into 17-41 distinct bundles.
Four retractor muscles.
A. Body covered with papillae. Numerous filiform tentacles which seldom (or
never?) surround the mouth, but stand above and dorsal to it in a horse-
shoe, with the opening dorsal. No rectal caecum. Hooks usually present.
Four retractors (in Ph. Rupellii only two?). Heart almost always without
caeca. Eye-spots always present. Eggs oval, flat, reddish. Almost entirely
small tropical species
1. Phymosoma
B. Body devoid of papillae. Tentacular membrane surrounds the mouth in a

circlet. Rectum with one or more caeca (except S. edulis?). Hooks absent
except in S. australis. Eggs spherical. The tentacular membrane contains a
vascular network. A ventral contractile vessel usually present in addition to
the heart. Mostly large forms. Found in all seas
2. Sipunculus
II. The longitudinal muscles in the body-wall form a continuous sheath, and are
not split up into bundles.
A. Two brown tubes. Numerous tentacles form a wreath round the mouth.
Alimentary canal forms a complete spiral, free behind except in Ph. Hanseni.
Spindle-muscle usually present. One or more ligaments present, but only on
the anterior convolutions of the intestine. Adhesive papillae always absent.
Hooks very frequently absent. Eggs spherical. Found in all seas.
3. Phascolosoma
B. Two free brown tubes. Only four or six plumed tentacles. A complete
intestinal spiral, not attached behind. Spindle-muscle always present. One or
more ligaments present, but only on the anterior convolutions of the
intestine. Hooks are present, but sometimes fall off early in life. Heart usually
bears caeca. Found only in the tropics.
4. Dendrostoma
C. Only one brown tube, that of the right side, present; it is attached to the
body-wall throughout its entire length. Numerous tentacles form a circle
round the mouth. The alimentary canal forms no spiral, or an incomplete
one. No spindle-muscle, but the intestine is attached to the body-wall
throughout its length by numerous ligaments. Adhesive papillae often
present. Not more than two retractors. Spherical eggs. Inhabits Mollusc
shells or tubes. Found in all seas
5. Phascolion
III. At both ends of the trunk a distinct horny shield, or tube-like cornification, or
a calcareous ring at the anterior end of the trunk. Hooks sometimes present.
Longitudinal muscles continuous or split up into bundles.
A. A shield at both ends of the trunk. Introvert excentric, arising from the
ventral side of the anterior shield. Tentacles small and few in number,
arranged in a horse-shoe above the mouth. A spindle-muscle, which arises
from the posterior end of the body, traverses the intestinal coil. Two
retractors only, these are the ventral; they are frequently fused together from
their point of origin.
6. Aspidosiphon
B. A calcareous ring surrounds the anterior end of the trunk, from the middle
of which the introvert is extruded. Longitudinal muscles continuous. Hooks
bifid. Tropical.
7. Cloeosiphon
C. A corneous ring, from which the introvert issues, surrounds the anterior
end of the trunk, and the posterior end of the trunk is produced into a
corneous spike. Six pinnate tentacles encircle the mouth. Four retractors.
Hooks present on the introvert. Longitudinal muscles continuous. Intestine
not coiled throughout in a spiral nor fastened posteriorly. Spindle muscle
present.
8. Golfingia
IV. No tentacles, but two leaf-like extensions of the body-wall guard the mouth.
Four retractors. Few intestinal loops, quite free. No vascular system.
9. Petalostoma
V. No tentacles, no vascular system. One retractor, and one segmental organ.
A. Introvert long. Body small, pear-shaped.

10. Onchnesoma
B. No introvert (?). Body cylindrical, thickly covered with papillae, which are
larger and more crowded at both ends of the trunk.
11. Tylosoma
Species of Sipunculoidea.—The genus Phymosoma (Fig. 214) contains more

species than any other genus of Sipunculoidea, and they are all of fair size. Twenty-
seven species are known, of which seventeen occur in the Malay Archipelago,
thirteen being found there alone. Phymosoma affects shallow water, the deepest
specimens being taken at a depth of about 50 fathoms; this may be due to the fact
that they flourish only in comparatively warm water. With very few exceptions, they
are found only in tropical seas, very often living in tubular excavations made in soft
coral rock.
The genus Sipunculus contains sixteen species. They are the largest and the most
conspicuous members of the group. They have a very wide distribution, some
species, as S. nudus (Fig. 212) and S. australis, being almost cosmopolitan. They
are most common in temperate and tropical seas, but S. norvegicus and S.
priapuloides are found far north, but always at considerable depths, 100 to 200
fathoms.
The following account of the habits of Sipunculus gouldii is taken from Mr.
Andrews'[482] paper on that species:—
"This Sipunculus is very abundant in certain small areas of compact, fine sand
darkened by organic matter and not laid bare at ordinary low tide. In such places,
only a few square metres in extent, they pierce the sand in all directions to a depth
of more than half a metre, making burrows with persistent lumen running from the
surface downward and then laterally, but with no regularity in direction.
"Kept in aquaria, the dependence of the animal upon the nature of the sand and its
method of locomotion may be readily observed. A vigorous individual buries itself in
a few moments in the following manner: Running out the introvert to nearly its full
extent, and applying it to the surface of the sand till some spot of less resistance is
found, the animal still further expands the introvert so that it penetrates the sand,
provided this is not too dense and firm, for then the body is merely shoved
backward. When the introvert is inserted, the contraction of the longitudinal muscles
of the body-wall brings the whole body forward somewhat, in case the introvert is
fixed in the sand. In case soft ooze was present, this fixation did not take place, and
the introvert was merely pulled out again, but when the sand was of the right
consistency the introvert was fixed by becoming much swollen at the tip, and then
constricted just posterior to this swollen area. This bulb-like area exerts lateral
pressure on the sand, as could be seen by movements of the grains. The swelling
of the anterior end of the introvert is brought about by the body-wall contracting
elsewhere, and forcing in liquid to distend that end. Owing to the curved form
assumed by the body in the normal contracted state when first removed from its
burrow, the entrance of the introvert may often be nearly vertical, and hence the
entire body is soon raised nearly upright in the water above the sand. If the body
has thus been warped forward sufficiently to become somewhat fixed in the sand,
the introvert is rolled in and again thrust forward from this new point of resistance,
and so on till the animal is entirely buried. This locomotion increases in speed as the
creature becomes more completely surrounded by sand, and is the only means of
moving from place to place.
"On a smooth surface, or on one not presenting the right degree of resistance, the
Sipunculus does not change its position, but remains till death finally occurs, rolling
its introvert in and out and contracting its body-wall to no purpose.
"The essential factors in the mechanism bringing about this hydrostatic locomotion
are an elongated contractile sac filled with liquid, and some means of definitely co-
ordinating the contractions of the sac.
"In natural environment the animals are found with sometimes one, sometimes the
other end nearer the surface of the sand: in the aquaria the same was observed, but
when the water became stagnant and impure the anterior end with expanded
branchiae was often protruded somewhat above the surface of the sand."
The genus Phascolosoma contains at least twenty-five species, for the most part
small. Ph. margaritaceum, however, measures[483] 10 cm. in length, and Ph.
flagriferum, 13 cm. The latter is produced at the hinder end of its trunk into a long
whip-like process, which recalls the horny spike of Golfingia. Most species live free,
but a few inhabit the shells of dead Gasteropods or of Dentalium, or the abandoned
tubes of worms. They occur in practically all seas.
Fig. 216.—Specimens of the Coral Heteropsammia cochlea, with Aspidosiphon
heteropsammiarum or A. michelini living in a state of commensalism with them.
(From Bouvier.)
Dendrostoma contains but five species, which are all found within the tropics in the
Pacific or in the West Atlantic. They are shallow-water forms, and some are found
between tide-marks.
Phascolion is a smaller genus, containing but ten species, which may have been
derived independently from different species of Phascolosoma, and in this case the
genus should be broken up. The members of this genus live in Mollusc shells, such
as Dentalium, Turritella, Buccinum, Chenopus (Aporrhais), Nassa, Strombus, and
generally acquire the coiled shape of their host. They are usually attached to the
shell by means of certain adhesive papillae found on their posterior end. Ph. strombi
fills its shell with mud, which must be kept together by some secretion of the animal.
The body lies in a tube in this mud, and the introvert projects from the small round
opening at the end of the tube, and explores the ground in every direction. They are
found in all seas, but more especially in the colder waters.
Fig. 217.—Cloeosiphon aspergillum Quatr. × ½. a, Introvert covered with spines and

partially extended, but not sufficiently to show the head; b, calcareous plates
surrounding the point of origin of the introvert.
The genus Aspidosiphon includes nineteen species, which are, with few exceptions,
exclusively confined to the Indian Ocean and neighbouring seas, including the Red
Sea. The exceptions are A. armatus from the Norwegian coast, and A. mülleri from
the Mediterranean and Adriatic. A. truncatus is also stated to occur at Panama, the
Bahamas, and at Mauritius. The remaining species almost all occur in the Malay
Archipelago and neighbouring islands, and as was the case with Phymosoma, this
part of the world seems to be the headquarters of the genus. A. mülleri lives in the
interstices of rocks and stones, and occasionally in disused Mollusc shells.
Two species of Aspidosiphon have been described by Bouvier[484] living in a state

of commensalism with two species of Madreporarian corals, Stephanoceris
rousseaui and Heteropsammia cochlea, which live on and surrounding the shells of
certain Molluscs at Aden (Fig. 216). Apparently the Gephyrean takes up its abode
within its house at a tender age, and according to Bouvier, it provides for its
increasing bulk by secreting a coiled calcareous tube, the outer surface of which
affords space for the growth of the coral.
The genus Cloeosiphon, the Echinosiphon of Sluiter, includes three species: C.

aspergillum (Fig. 217), C. molle, and C. javanicum. The first named occurs at
Mauritius, the Malay Archipelago, and neighbouring islands; the others are confined
to the last-named area, which thus again forms the headquarters of a genus.
Golfingia, described by Lankester from a single specimen, was dredged in St.

Andrews Bay, at the depth of 10 fathoms.
Petalostoma comprises but one species, P. minutum, which is found in the English
Channel.
Onchnesoma comprises two species, O. steenstrupii and O. sarsii, both found off
the coast of Norway at considerable depths between 200 and 300 fathoms.
Tylosoma comprises one species, T. lütkenii, also from the Norwegian coast. It is
dredged from stony ground in 50 to 80 fathoms.
II. Order Priapuloidea.
Anatomy.—This Order consists of the two genera Priapulus and Halicryptus. Both
are cylindrical animals with the mouth at one end and the anus at the other. The
introvert is short, and is covered with rows of chitinous spines, which are continued
to some extent over the body.
The skin is folded in a series of rings, and the body is usually somewhat swollen
posteriorly. P. caudatus bears a curious caudal appendage, beset with a number of
hollow lobes somewhat grape-like in appearance. This is situated ventral to the
anus; its lumen is continuous with that of the body-cavity, but it can be separated
from it by the action of a sphincter muscle. Two such appendages exist in P.
bicaudatus.
There cannot be said to be any head in the Priapuloidea; they have no tentacles or
tentacular fringe, no proboscis, and no distinct brain; simply a round aperture, the
mouth, which is surrounded by a groove in the skin, at the bottom of which the
circumoesophageal nerve-cord lies. The mouth leads into a very muscular pharynx
lined with stout chitinous teeth; this passes into an intestine, which is as a rule
straight, but in P. glandifer it has a single loop.
The Priapuloidea possess no vascular system and no brown tubes. Their skin has in
the main the same structure as that of the Sipunculids, with spines, glandular
bodies, and papillae with sensory hairs which resemble similar structures on
Phymosoma varians. Retractor muscles arise from the longitudinal muscles of the
skin, and are inserted into the pharynx; they are short and not constant in number.
The nervous system has retained throughout its primitive connexion with the
epidermis. In almost all animals the nervous system is formed from the epiblast or
outermost cellular layer of the embryo; it usually, however, breaks away from this
and sinks into the body. Thus in Sipunculus it lies within the body-cavity, and has
retained its primitive connexion with the outer layers of the skin only in the region of
the brain; but in the Priapulids the nervous system, which consists of a ring round
the mouth and of a ventral cord, lies embedded in the skin, and the nerve cells are
directly continuous with the cells of the epidermis. The nerve-ring lies at the base of
a groove in the skin, which forms a kind of gutter round the mouth; the ventral
nerve-cord is visible exteriorly as a light line which marks the ventral surface of the
animal. In no place is the ring or cord differentiated in any way, and there cannot be
said to be any brain or special sense-organs. Numerous nerves are given off from
the ring to the pharynx and intestine, and from the cord to the body-wall.
Fig. 218.—Priapulus caudatus Lam. Nat. size. a, Mouth surrounded by spines.
The sexes are distinct, but they differ from the other Gephyrea in the nature of their
reproductive organs. In mature specimens the ovaries or testes are easily
recognisable, lying to the right and left of the alimentary canal. The reproductive
glands are continuous with ducts, which act as oviducts and vasa deferentia
respectively. Both glands and ducts are attached to the body-wall by a mesentery.
The excretory function is performed in the Priapuloidea by the ducts of the

generative organs. These are primarily connected with a number of branching
canals of small size which project into the body-cavity. According to Schauinsland,
[485] one or more pear-shaped cells are found at the end of each branch, and each
is continued into a long cilium which hangs down into the lumen of the canal, and by
its movement produces a flickering motion. Beyond the free end of the large cilium
the canal is lined with ciliated cells. The remarkable resemblance this form of
excretory organ presents to that of the Platyhelminthes (vide p. 25) and of certain
Chaetopods is worthy of attention. In the young Priapuloidea the duct with its
branching canals is not masked by the generative organs, but as the animals
become mature, diverticula from the duct arise, and the cells covering these
become modified into ova in the female, and into spermatozoa in the male. The
presence of these follicles masks the excretory part of the gland. The ova and
spermatozoa escape through the ciliated ducts which open to the exterior one on
each side of the anus, and, contrary to what is the case with other Gephyrea, leave
the body without having ever been in the body-cavity.
Nothing is known of the embryology of either member of this family, but both genera
appear to be sexually mature from the end of May until October.
Classification.—The two genera which make up the Order Priapuloidea are

characterised as follows:—
Priapulus.—The body is continued into one or two caudal appendages, beset with
hollow papillae; these are ventral to the anus. The introvert forms ¼ to ⅓ of the total
body-length; it is covered with spines in conspicuous longitudinal rows, the rest of
the body being ringed. The retractor muscles are numerous, and are attached to the
body-wall, some anteriorly and some posteriorly.
The genus includes the following five species:—
P. caudatus Lam. (Fig. 218). Hab. Coasts of Greenland, Norway, Great Britain,
the North Sea, and the Baltic.
P. bicaudatus Dan. Hab. North Sea and Arctic Ocean.
P. glandifer Ehlers. Hab. Coast of Greenland, North Sea.

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(eBook PDF) Energy, Environment, and

Climate 3rd Edition

List of Focus Boxes xi CHAPTER 4

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10.4 Other Indirect Solar Energy Sources 287

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BOX 1.1 BEING GRAPHICALLY LITERATE 16

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Four chapters on climate follow the section on energy. Chapter 12 describes

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Energy, Environment, and Climate helps reinforce qualitative and quantita-

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national and international, has been thoroughly updated; examples include an

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FIRST EDITION REVIEWERS

SECOND EDITION REVIEWERS

THIRD EDITION REVIEWERS

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ABOUT THE AUTHOR

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1.1 Earth’s Beginnings

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Fam. 2. Herpobdellidae.—Pharynx without denticulate jaws, with three unarmed

A characteristic genus of this family is Trocheta, which is so common at the

The genus Haemadipsa, which M. Blanchard places in a special sub-family,

ARTHUR E. SHIPLEY, M.A.

The Anatomy of Sipunculus nudus.

External Characters.—The body of S. nudus when fully extended may attain a

Body-wall.—The glistening appearance of Sipunculus is due to the cuticle, a

Respiratory System.—There are no special respiratory organs, and it has long

Body-Cavity.—The pinkish fluid of the body-cavity contains numerous corpuscles,

Nervous System.—The nervous system of Sipunculus consists of a brain or

The nerve-cord is supported by numerous strands of muscle which pass to it from

Excretory System.—The excretory organs or "brown tubes" are typical nephridia,

Fig. 213.—Larva of Sipunculus nudus L. × 150. (After Hatschek.) a, Mouth; b, anus; c,

Phascolosoma contains, in comparison with Sipunculus, only small species, and it is

Fig. 214.—A, Phymosoma granulatum F. S. Leuck. × 2. B, Head of the same. × 4. a,

In S. tesselatus the sand is to some extent held together by a mucous deposit; in

Aspidosiphon (Fig. 215) is easily recognised by the presence of two symmetrically-

Fig. 215.—Aspidosiphon truncatus Kef. × 2. a, Introvert partially extended, but not

Key to the Genera of Sipunculoidea.[481]

B. Body devoid of papillae. Tentacular membrane surrounds the mouth in a

V. No tentacles, no vascular system. One retractor, and one segmental organ.

A. Introvert long. Body small, pear-shaped.

Species of Sipunculoidea.—The genus Phymosoma (Fig. 214) contains more

Fig. 217.—Cloeosiphon aspergillum Quatr. × ½. a, Introvert covered with spines and

Two species of Aspidosiphon have been described by Bouvier[484] living in a state

The genus Cloeosiphon, the Echinosiphon of Sluiter, includes three species: C.

Golfingia, described by Lankester from a single specimen, was dredged in St.

II. Order Priapuloidea.

Fig. 218.—Priapulus caudatus Lam. Nat. size. a, Mouth surrounded by spines.

The excretory function is performed in the Priapuloidea by the ducts of the

Classification.—The two genera which make up the Order Priapuloidea are

The genus includes the following five species:—

P. bicaudatus Dan. Hab. North Sea and Arctic Ocean.

P. glandifer Ehlers. Hab. Coast of Greenland, North Sea.

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